The Birds of Africa Volume VII 9781472927033, 9780713665314

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vi

CONTENTS

PLATES Plate 1

Facing Page

18

Sparrows (grey-headed, Chestnut, golden), petronias . . . .

. .

10

2

Sparrows (rufous, Cape, Desert)

. .

11

3

Buffalo-weavers, sparrow-weavers, speckled weavers, Rufous-tailed Weaver

26

4

Malimbes

27

5

Social weavers, Sociable Weaver, Ploceus (Baglafecht Weaver and others) . .

74

Ploceus (Slender-billed Weaver and others) . . . . . .

. .

75

7

Ploceus (golden weavers and others) . .

90

8

Ploceus (masked weavers and others) . .

91

9

Ploceus (Village Weaver and others) . .

138

10

Ploceus (dark-backed weavers and others)

139

11

Ploceus (nuthatch weavers and others), Compact Weaver . . . . . .

154

Queleas, Red-headed and Bob-tailed Weavers, yellow bishops . . . .

155

13

Red bishops

. .

202

14

Red-collared Widowbird and others . .

203

15

Marsh widowbirds, Long-tailed Widowbird

218

6

12

16 17

. .

. .

. .

. .

. .

. .

. .

Plate

. .

. .

Facing Page Twinspots, waxbills (grey, swee and others) . . . . . .

. .

267

19

Masked waxbills, Zebra Waxbill

. .

282

20

Seedcrackers, bluebills, twinspots

. .

283

21

Cordon-bleus, grenadiers, Locust-Finch, quailfinches

. .

. .

330

22

Firefinches

. .

. .

. .

. .

331

23

Firefinches

. .

. .

. .

. .

346

24

Mannikins, silverbills, Cut-throat Finch and others . . . .

. .

347

Steel-blue, Pin-, Straw- and Shaft-tailed Whydahs, Cuckoo-Finch . . . .

394

26

Paradise-whydahs

. .

. .

. .

395

27

Indigobirds

. .

. .

. .

410

28

Citrils, canaries, seedeaters

. .

. .

411

29

Canaries (Brimstone and others)

. .

458

30

Seedeaters, Yellow-crowned and Cape Canaries . . . . . .

. .

459

Serins, siskins, black-headed canaries, Oriole-Finch . . . .

. .

474

32

Grosbeaks, northern finches, linnets . .

475

33

Sarahan and North African finches

522

34

Chaffinch, golden and cinnamon buntings 523

25

31

. .

. .

Thick-billed Weaver, antpeckers, negrofinches . . . . . .

. .

219

35

Rock and northern buntings . .

. .

538

Olivebacks, pytilias, crimsonwings

. .

266

36

Northern buntings

. .

539

. .

. .

Plate 1

Swainson's Sparrow (p. 4) Passer swainsonii P. g. ugandae

Imm. Northern Grey-headed Sparrow (p. 2) Passer griseus Ad.

P. g. laeneni

Swahili Sparrow (p. 7) Passer suahelicus

Imm.

Parrot-billed Sparrow (p. 6) Passer gongonensis

Southern Grey-headed Sparrow (p. 7) Passer diffusus diffusus

Ad.




Chestnut Sparrow (p. 34) Passer eminibey



Sudan Golden Sparrow (p. 36) Passer luteus



Arabian Golden Sparrow (p. 40) Passer euchlorus 


Bush Petronia (p. 42) Petronia dentata

Yellow-spotted Petronia (p. 44) Petronia pyrgita pyrgita

Rock Sparrow (p. 47) Petronia petronia barbara

10

6 in 15 cm

Yellow-throated Petronia (p. 45) Petronia superciliaris superciliaris

Plate 2

 breeding


P. d. indicus  non-breeding

P. d. rufidorsalis  breeding


Spanish Sparrow (p. 24) Passer hispaniolensis hispaniolensis

House Sparrow (p. 20) Passer domesticus

P. d. indicus  breeding P. d. tingitanus  breeding

Eurasian Tree Sparrow (p. 32) Passer montanus montanus






Great Sparrow (p. 9) Passer motitensis motitensis

Shelley’s Rufous Sparrow (p. 14) Passer shelleyi  Kenya Rufous Sparrow (p. 13) Passer rufocinctus

 Kordofan Rufous Sparrow (p. 15) Passer cordofanicus

Socotra Sparrow (p. 16) Passer insularis















Desert Sparrow (p. 29) Passer simplex simplex 

Cape Sparrow (p. 17) Passer melanurus melanurus 

 Somali Sparrow (p. 31) Passer castonopterus castanopterus

6 in 15 cm

11

Plate 3

P. m. pectoralis

 non-breeding Juv.

White-browed Sparrow-Weaver (p. 56) Plocepasser mahali White-billed Buffalo-Weaver (p. 50) Bubalornis albirostris  breeding

P. m. melanorhynchus

Speckle-fronted Weaver (p. 64) Sporopipes frontalis emini

Juv.

D. d. dinemelli

D. d. boehmi

Scaly-fronted Weaver (p. 65) Sporopipes squamifrons



White-headed Buffalo-Weaver (p. 54) Dinemellia dinemelli Red-billed Buffalo-Weaver (p. 52) Bubalornis niger intermedius




Chestnut-crowned Sparrow-Weaver (p. 58) Plocepasser superciliosus

Juv.

Donaldson-Smith’s Sparrow-Weaver (p. 61) Plocepasser donaldsoni

Ad.

Rufous-tailed Weaver (p. 62) Histurgops ruficaudus

Chestnut-backed Sparrow-Weaver (p. 60) Plocepasser rufoscapulatus

26

6 in 15 cm

1

Plate 4 M. m. malimbicus

Ad.



Juv.

M. r. rubricollis



M. m. crassirostris Ad. 

Red-headed Malimbe (p. 95) Malimbus rubricollis

Ad. 

Crested Malimbe (p. 89) Malimbus malimbicus Ad.  Blue-billed Malimbe (p. 93) Malimbus nitens

M. r. bartletti






Juv. Rachel’s Malimbe (p. 80) Malimbus racheliae






Cassin’s Malimbe (p. 82) Malimbus cassini

Juv.  Gola Malimbe (p. 78) Malimbus ballmanni 







Red-crowned Malimbe (p. 87) Malimbus coronatus

Red-vented Malimbe (p. 81) Malimbus scutatus scutatus






Juv.




Red-bellied Malimbe (p. 86) Malimbus erythrogaster erythrogaster

Juv. 

 Ibadan Malimbe (p. 84) Malimbus ibadanensis

6 in 15 cm

1 27

Plate 5 P. a. arnaudi

Juv.

Juv.

Ad.

Ad. Black-capped Social Weaver (p. 70) Pseudonigrita cabanisi

Grey-headed Social Weaver (p. 67) Pseudonigrita arnaudi

Ad. P. a. dorsalis

Juv. P. b. neumanni

Ad. Sociable Weaver (p. 72) Philetairus socius

 breeding

 non-breeding P. b. emini Baglafecht Weaver (p. 106) Ploceus baglafecht

Juv.

 breeding

P. b. reichenowi
breeding

Ad.


P. b. emini

Ad. 

Imm.  Imm. 



P. b. reichenowi

P. b. nyikae  




P. b. stuhlmanni

74

Bannerman’s Weaver (p. 109) Ploceus bannermani

6 in 15 cm

Black-chinned Weaver (p. 108) Ploceus nigrimentus


Plate 6









Bertram’s Weaver (p. 104) Ploceus bertrandi

Bates’s Weaver (p. 110) Ploceus batesi


 breeding






Slender-billed Weaver (p. 112) Ploceus pelzelni

 nonbreeding




Little Weaver (p. 111) Ploceus luteolus 

Loango Weaver (p. 114) Ploceus subpersonatus







 P. n. nigricollis 

P. n. brachypterus P. n. melanoxanthus

Black-necked Weaver (p. 115) Ploceus nigricollis









Spectacled Weaver (p. 117) Ploceus ocularis suahelicus Strange Weaver (p. 121) Ploceus alienus 

Black-billed Weaver (p. 120) Ploceus melanogaster



6 in 15 cm

75

Plate 7

P. c. ‘olivaceus’  breeding

 breeding




 non-breeding

P. c. ‘capensis’ Eastern Golden Weaver (p. 126) Ploceus subaureus

 breeding

 non-breeding

P. s. subaureus

 breeding


P. s. aureoflavus

Cape Weaver (p. 122) Ploceus capensis


 breeding




 non-breeding

Holub’s Golden Weaver (p. 128) Ploceus xanthops 

Bocage’s Weaver (p. 125) Ploceus temporalis












Orange Weaver (p. 130) Ploceus aurantius

Taveta Golden Weaver (p. 133) Ploceus castaneiceps



Golden Palm Weaver (p. 132) Ploceus bojeri





 non-breeding

 breeding Southern Brown-throated Weaver (p. 135) Ploceus xanthopterus

90

6 in 15 cm

Príncipe Golden Weaver (p. 134) Ploceus princeps

Plate 8 Kilombero Weaver (p. 140) Ploceus burnieri 




Northern Brown-throated Weaver (p. 137) Ploceus castanops







Northern Masked Weaver (p. 141) Ploceus taeniopterus 

P. i. intermedius  P. i. cabanisii
Lesser Masked Weaver (p. 142) Ploceus intermedius

Southern Masked Weaver (p. 145) Ploceus velatus velatus  breeding


breeding  breeding

Vitelline Masked Weaver (p. 148) Ploceus vitellinus vitellinus

Lake Lufira Weaver (p. 152) Ploceus ruweti  breeding

Katanga Masked Weaver (p. 151) Ploceus katangae  breeding  nonbreeding

Tanganyika Masked Weaver (p. 150) Ploceus reichardi  breeding


 breeding

  non-breeding

Heuglin’s Masked Weaver (p. 153) Ploceus heuglini

 breeding

Rüppell’s Weaver (p. 157) Ploceus galbula

91 6 in 15 cm

Plate 9 Ɋ ɉ

Ɋ ɉ

Fox’s Weaver (p. 160) Ploceus spekeoides

Speke’s Weaver (p. 158) Ploceus spekei

ɉ P. c. nigriceps

ɉ Ɋ

ɉ dark-backed form

P. c. cucullatus Village Weaver (p. 164) Ploceus cucullatus

Ɋ ɉ P. c. bohndorffi

Giant Weaver (p. 170) Ploceus grandis ɉ ɉ

Ɋ

ɉ non-breeding ɉ breeding

Juba Weaver (p. 173) Ploceus dicrocephalus Yellow-backed Weaver (p. 174) Ploceus melanocephalus

P. m. dimidiatus

ɉ breeding

Ɋ ɉ

Ɋ Golden-backed Weaver (p. 175) Ploceus jacksoni

P. m. melanocephalus

138

6 in 15 cm

Plate 10


P. n. nigerrimus

 P. n. castaneofuscus






Maxwell’s Black Weaver (p. 182) Ploceus albinucha albinucha Vieillot’s Black Weaver (p. 161) Ploceus nigerrimus






Yellow-legged Weaver (p. 104) Ploceus flavipes


Yellow-mantled Weaver (p. 181) Ploceus tricolor 

Golden-naped Weaver (p. 180) Ploceus aureonucha





Weyns’s Weaver (p. 171) Ploceus weynsi

São Tomé Weaver (p. 193) Ploceus sanctithomae



P. b. amaurocephalus

P. b. stictifrons

P. b. kersteni Dark-backed Weaver (p. 183) Ploceus bicolor

P. b. mentalis

P. b. bicolor

6 in 15 cm

139

Plate 11


 Clarke’s Weaver (p. 172) Ploceus golandi




Usambara Weaver (p. 191) Ploceus nicolli

Bar-winged Weaver (p. 192) Ploceus angolensis







Olive-headed Weaver (p. 189) Ploceus olivaceiceps

Preuss’s Weaver (p. 188) Ploceus preussi





Brown-capped Weaver (p. 186) Ploceus insignis

 nonbreeding

Yellow-capped Weaver (p. 187) Ploceus dorsomaculatus






Compact Weaver (p. 196) Pachyphantes superciliosus  breeding











Streaked Weaver (p. 195) Ploceus manyar

Chestnut Weaver (p. 177) Ploceus rubiginosus rubiginosus 

Cinnamon Weaver (p. 179) Ploceus badius

154




6 in 15 cm

Plate 12






A. r. rubriceps 

A. r. leuconotus 



Red-headed Weaver (p. 97) Anaplectes rubriceps

Cardinal Quelea (p. 204) Quelea cardinalis

Q. q. quelea  A. r. jubaensis





  Red-billed Quelea (p. 207) Quelea quelea

Red-headed Quelea (p. 205) Quelea erythrops 


Q. q. lathami 

Bob-tailed Weaver (p. 213) Brachycope anomala

 non-breeding

 breeding E. c. capensis

E. a. afer




E. a. strictus

 breeding

 breeding

Yellow Bishop (p. 233) Euplectes capensis

Yellow-crowned Bishop (p. 228) Euplectes afer  breeding E. c. crassirostris



 breeding Golden-backed Bishop (p. 232) Euplectes aureus

6 in 15 cm

155

Plate 13

E. o. ‘nigrifrons’

E. o. ‘orix’


 breeding

 breeding

 non-breeding Southern Red Bishop (p. 217) Euplectes orix



 breeding

 non-breeding

 breeding

Northern Red Bishop (p. 215) Euplectes franciscanus

Zanzibar Bishop (p. 223) Euplectes nigroventris

 nonbreeding  breeding


 non-breeding

Black Bishop (p. 227) Euplectes gierowii friederichseni

 breeding




Fire-fronted Bishop (p. 230) Euplectes diadematus

 breeding
 non-breeding

 nonbreeding Black-winged Bishop (p. 224) Euplectes hordeaceu

202

6 in 15 cm

Plate 14


E. a. eques  breeding

E. m. macrocerca

 breeding

 breeding


 breeding E. m. macroura

E. a. albonotatus

Yellow-mantled Widowbird (p. 237) Euplectes macroura

White-winged Widowbird (p. 235) Euplectes albonotatus

 non-breeding

 non-breeding




 breeding E. a. ‘concolor’  breeding E. a. suahelica

 breeding E. a. bocagei

 non-breeding

 breeding E. a. ardens Fan-tailed Widowbird (p. 242) Euplectes axillaris


E. a. axillaris Red-collared Widowbird (p. 239) Euplectes ardens

 breeding




 non-breeding

203 6 in 15 cm

Plate 15




Long-tailed Widowbird (p. 246) Euplectes progne

 breeding  breeding  nonbreeding




Mountain Marsh Widowbird (p. 246) Euplectes psammocromius

 breeding

 breeding

E. h. humeralis

 non-breeding E. h. hartlaubi


Hartlaub’s Marsh Widowbird (p. 244) Euplectes hartlaubi




 nonbreeding

218

Jackson’s Widowbird (p. 249) Euplectes jacksoni

 breeding

6 in 15 cm

 nonbreeding

Plate 16

ɉ A. a. montana

A. a. capitalba

A. a. albifrons

Ɋ Thick-billed Weaver (p. 198) Amblyospiza albifrons

ɉ

Ɋ

Ɋ Woodhouse’s Antpecker (p. 261) Parmoptila woodhousei woodhousei

ɉ Red-fronted Antpecker (p. 263) Parmoptila rubrifrons

Juv. Juv.

ɉ Ɋ Juv. Jameson’s Antpecker (p. 264) Parmoptila jamesoni N. b. saturatior Ɋ sub-adult N. f. uropygialis N. b. bicolor White-breasted Negrofinch (p. 259) Nigrita fusconotus

Chestnut-breasted Negrofinch (p. 257) Nigrita bicolor

N. f. fusconotus Ɋ

N. c. schistaceus N. c. emiliae

Pale-fronted Negrofinch (p. 253) Nigrita luteifrons luteifrons ɉ

N. c. canicapillus

Grey-headed Negrofinch (p. 255) Nigrita canicapillus

6 in 15 cm

219

Plate 17






Grey-headed Oliveback (p. 270) Nesocharis capistrata

Shelley’s Oliveback (p. 265) Nesocharis shelleyi shelleyi

White-collared Oliveback (p. 268) Nesocharis ansorgei ansorgei









P. m. citerior 

P. m. melba

Green-winged Pytilia (p. 354) Pytilia melba  P. m. belli







Orange-winged Pytilia (p. 358) Pytilia afra

Yellow-winged Pytilia (p. 350) Pytilia hypogrammica






Red-billed Pytilia (p. 354) Pytilia lineata Red-winged Pytilia (p. 352) Pytilia phoenicoptera



Red-faced Crimsonwing (p. 281) Cryptospiza reichenovii



Dusky Crimsonwing (p. 279) Cryptospiza jacksoni

Shelley’s Crimsonwing (p. 278) Cryptospiza shelleyi






 Abyssinian Crimsonwing (p. 280) Cryptospiza salvadorii salvadorii 266

6 in 15 cm

Plate 18









Brown Twinspot (p. 348) Clytospiza monteiri Dusky Twinspot (p. 340) Euschistospiza cinereovinacea

Dybowski’s Twinspot (p. 338) Euschistospiza dybowskii

Lavender Waxbill (p. 287) Estrilda caerulescens

Grey Waxbill (p. 289) Estrilda perreini

Cinderella Waxbill (p. 290) Estrilda thomensis

Orange-cheeked Waxbill (p. 295) Estrilda melpoda 

Anambra Waxbill (p. 293) Estrilda poliopareia

Swee Waxbill (p. 272) Coccopygia melanotis melanotis


E. p. paludicola

E. p. ochrogaster E. p. roseicrissa Fawn-breasted Waxbill (p. 292) Estrilda paludicola

Yellow-bellied Waxbill (p. 267) Coccopygia quartinia

6 in 15 cm

267

Plate 19 Juv. Crimson-rumped Waxbill (p. 297) Estrilda rhodopyga

Black-rumped Waxbill (p. 298) Estrilda troglodytes

E. a. astrild

Ad.

E. a. minor

E. a. rubriventris

Common Waxbill (p. 300) Estrilda astrild E. a. peasei

E. a. cavendishi

Black-lored Waxbill (p. 304) Estrilda nigriloris

Black-headed Waxbill (p. 307) Estrilda atricapilla Kandt’s Waxbill (p. 308) Estrilda kandti

Black-crowned Waxbill (p. 305) Estrilda nonnula

Black-cheeked Waxbill (p. 311) Estrilda charmosyna E. e. erythronotos


E. e. delamerei


Black-faced Waxbill (p. 309) Estrilda erythronotos

Red Avadavat (p. 398) Amandava amandava amandava 



Zebra Waxbill (p. 392) Sporaeginthus subflavus subflavus

282

6 in 15 cm

Plate 20









Lesser Seedcracker (p. 323) Pyrenestes minor

Crimson Seedcracker (p. 322) Pyrenestes sanguineus


 

Black-bellied Seedcracker (p. 318) Pyrenestes ostrinus

S. h. pustulata









Grant’s Bluebill (p. 313) Spermophaga poliogenys

Western Bluebill (p. 314) Spermophaga haematina




S. h. haematina Red-headed Bluebill (p. 316) Spermophaga ruficapilla ruficapilla













Pink-throated Twinspot (p. 344) Hypargos margaritatus  H. n. macrospilotus







Red-throated Twinspot (p. 342) Hypargos niveoguttatus


H. n. niveoguttatus

M. n. nitidula  M. n. schlegeli Green Twinspot (p. 275) Mandingoa nitidula

6 in 15 cm





283

Plate 21 U. b. bengalus









 U. a. niassensis


U. b. katangae

Blue Waxbill (p. 325) Uraeginthus angolensis

Red-cheeked Cordon-bleu (p. 327) Uraeginthus bengalus





U. b. brunneigularis




U. a. angolensis

Blue-capped Cordon-bleu (p. 332) Uraeginthus cyanocephalus





Purple Grenadier (p. 334) Granatina ianthinogaster



Violet-eared Waxbill (p. 335) Granatina granatina





P. l. uelensis




P. l. locustella

Locust-Finch (p. 390) Paludipasser locustella






Black-faced Quailfinch (p. 384) Ortygospiza atricollis atricollis 

African Quailfinch (p. 388) Ortygospiza fuscocrissa fuscocrissa

Black-chinned Quailfinch (p. 386) Ortygospiza gabonensis gabonensis 



330 6 in 15 cm

Plate 22 Juv.






L. s. senegala

Red-billed Firefinch (p. 360) Lagonosticta senegala

Bar-breasted Firefinch (p. 364) Lagonosticta rufopicta rufopicta

Ad. 




L. s. rendalli

L. s. ruberrima



L. l. nigricollis  

L. l. vinacea Black-faced Firefinch (p. 367) Lagonosticta larvata

 L. l. larvata Juv.


L. r. rara Brown Firefinch (p. 366) Lagonosticta nitidula



Ad.

Black-bellied Firefinch (p. 369) Lagonosticta rara


L. r. forbesi

6 in 15 cm



331

Plate 23







L. r. rubricata  African Firefinch (p. 371) Lagonosticta rubricata L. r. haematocephala
L. r. polionota




Jameson’s Firefinch (p. 373) Lagonosticta rhodopareia

L. r. rhodopareia 


L. r. jamesoni











Mali Firefinch (p. 375) Lagonosticta virata

 Chad Firefinch (p. 378) Lagonosticta umbrinodorsalis Rock Firefinch (p. 376) Lagonosticta sanguinodorsalis 

346

6 in 15 cm

Plate 24 Juv. Ad.

Ad. E. c. orientalis

Juv. Ad. E. c. cantans

Ad.

S. c. cucullatus

African Silverbill (p. 412) Euodice cantans

S. c. scutatus

Bronze Mannikin (p. 401) Spermestes cucullatus

Ad. Juv.

S. b. bicolor West Africa

Juv.

Juv.

Ad. S. b. bicolor Mt Nimba

Grey-headed Silverbill (p. 400) Odontospiza griseicapilla

Ad. Black-and-white Mannikin (p. 405) Spermestes bicolor Juv.

Ad.

Ad.

S. b. ‘stigmatophorus’ S. b. poensis

S. b. nigriceps

Ad. S. f. ‘pica’



Ad.
Juv. S. f. ‘fringilloides’

Red-headed Finch (p. 382) Amadina erythrocephala erythrocephala

Magpie Mannikin (p. 407) Spermestes fringilloides A. f. meridionalis

Ad.



Cut-throat Finch (p. 379) Amadina fasciata

Java Sparrow (p. 415) Padda oryzivora






A. f. fasciata

6 in 15 cm

347

Plate 25

Ad.  breeding

Ad.  nonbreeding

Ad.  breeding

Juv.

Ad.  nonbreeding

Shaft-tailed Whydah (p. 434) Vidua regia Pin-tailed Whydah (p. 420) Vidua macroura

Juv.




Ad.
nonbreeding

Ad.
breeding

Ad.  breeding

Ad.  breeding Juv.

Straw-tailed Whydah (p. 433) Vidua fischeri

Ad.


Juv.

Ad.  nonbreeding

Steel-blue Whydah (p. 431) Vidua hypocherina



Ad.  nonbreeding

Ad.


Imm. 




Cuckoo-Finch (p. 417) Anomalospiza imberbis

394

6 in 15 cm

Plate 26

 breeding

 non-breeding







 breeding Exclamatory Paradise-Whydah (p. 424) Vidua interjecta

 nonbreeding

 non-breeding

Long-tailed Paradise-Whydah (p. 428) Vidua paradisaea

V. o. aucupum

 breeding  breeding Togo ParadiseWhydah (p. 426) Vidua togoensis  breeding Sahel Paradise-Whydah (p. 425) Vidua orientalis

 breeding  nonbreeding

Broad-tailed Paradise-Whydah (p. 427) Vidua obtusa

V. o. orientalis







6 in 15 cm

395

Plate 27

Plateau Indigobird (p. 442) Vidua maryae

Quailfinch Indigobird (p. 441) Vidua nigeriae

Fonio Indigobird (p. 445) Vidua camerunensis

Barka Indigobird (p. 444) Vidua larvaticola

Bar-breasted Firefinch Indigobird (p. 439) Vidua wilsoni Jambandu Indigobird (p. 443) Vidua raricola


Juv.

White-billed form

Red-billed form

Village Indigobird (p. 436) Vidua chalybeata

Ad. . nonbreeding V. c. chalybeata V. c. amauropteryx

V. c. ultramarina

Ad. . breeding

V. f. funerea

Dusky Indigobird (p. 447) Vidua funerea

Purple Indigobird (p. 448) Vidua purpurascens Zambezi Indigobird (p. 450) Vidua codringtoni

410

V. f. nigerrima

6 in 15 cm

Plate 28 





African Citril (p. 460) Serinus citrinelloides kikuyensis


Southern Citril (p. 464) Serinus hypostictus

Western Citril (p. 463) Serinus frontalis











 Black-faced Canary (p. 465) Serinus capistratus

Forest Canary (p. 468) Serinus scotops






Papyrus Canary (p. 467) Serinus koliensis Ad. Imm. Ad. S. l. leucopygius

S. l. riggenbachi S. a. atrogularis White-rumped Seedeater (p. 470) Serinus leucopygius

Imm. Ad. 

Ad. Black-throated Canary (p. 471) Serinus atrogularis

Reichenow’s Seedeater (p. 473) Serinus reichenowi




S. a. lwenarum S. a. deserti S. a. semideserti

6 in 15 cm

Yellow-rumped Seedeater (p. 476) Serinus xanthopygius

411

Plate 29 

Yellow-fronted Canary (p. 479) Serinus mozambicus

S. m. caniceps

S. m. samaliyae







S. m. mozambicus


S. m. barbatus Lemon-breasted Canary (p. 477) Serinus citrinipectus 






Yellow-throated Seedeater (p. 484) Serinus flavigula Salvadori’s Seedeater (p. 486) Serinus xantholaemus

White-bellied Canary (p. 483) Serinus dorsostriatus maculicollis

Ad. Southern Grosbeak-Canary (p. 488) Serinus buchanani


S. s. sulphuratus

S. s. sharpei



Northern Grosbeak-Canary (p. 487) Serinus donaldsoni






Brimstone Canary (p. 489) Serinus sulphuratus

S. f. flaviventris 

Ad. Imm.

Yellow Canary (p. 491) Serinus flaviventris.

S. a. albogularis

S. a. crocopygia

Ad.




S. f. damarensis 

White-throated Canary (p. 494) Serinus albogularis

458

6 in 15 cm

Plate 30

Thick-billed Seedeater (p. 498) Serinus burtoni

Streaky Seedeater (p. 495) Serinus striolatus S. s. graueri

S. b. kilimensis

S. b. burtoni

Tanzania Seedeater (p. 499) Serinus melanochrous

S. s. striolatus

S. r. thomensis Yellow-browed Seedeater (p. 497) Serinus whytii

Príncipe Seedeater (p. 500) Serinus rufobrunneus

Black-eared Seedeater (p. 504) Serinus mennelli

S. r. rufobrunneus




Protea Canary (p. 502) Serinus leucopterus

Streaky-headed Seedeater (p. 506) Serinus gularis S. g. humilis

S. g. benguellensis

S. g. gularis S. g. canicapilla







Yellow-crowned Canary (p. 511) Serinus flavivertex

S. r. reichardi

Juv. Cape Canary (p. 513) Serinus canicollis





Brown-rumped Seedeater (p. 510) Serinus tristriatus

S. r. striatipectus

Reichard’s Seedeater (p. 509) Serinus reichardi Ad.

6 in 15 cm

459

Plate 31


Citril Finch (p. 518) Serinus citrinella citrinella


European Serin (p. 516) Serinus serinus








Syrian Serin (p. 518) Serinus syriacus


 
Red-fronted Serin (p. 520) Serinus pusillus

Ethiopian Siskin (p. 519) Serinus nigriceps



Cape Siskin (p. 521) Pseudochloroptila totta









Damara Canary (p. 529) Alario leucolaemus


Black-headed Canary (p. 527) Alario alario 




Oriole-Finch (p. 531) Linurgus olivaceus


 L. o. olivaceus

Drakensberg Siskin (p. 525) Pseudochloroptila symonsi



L. o. kilimensis  L. o. prigoginei



474

6 in 15 cm

Plate 32 Juv.


São Tomé Grosbeak (p. 530) Neospiza concolor




Somali Golden-winged Grosbeak (p. 536) Rhynchostruthus louisae 

Arabian Golden-winged Grosbeak (p. 534) Rhynchostruthus socotranus

 Juv.

Winter




European Goldfinch (p. 541) Carduelis carduelis parva European Greenfinch (p. 537) Carduelis chloris voousi Summer Juv. 




Eurasian Siskin (p. 543) Carduelis spinus

 Common Redpoll (p. 545) Carduelis flammea



Ankober Serin (p. 549) Carduelis ankoberensis





Juv.

Juv. Common Linnet (p. 545) Carduelis cannabina cannabina



Warsangli Linnet (p. 547) Carduelis johannis



6 in 15 cm

475

Plate 33 Pale Rockfinch (p. 551) Carpospiza brachydactyla







Scarlet Rosefinch (p. 550) Carpodacus erythrinus erythrinus

Crimson-winged Finch (p. 553) Rhodopechys sanguinea alienus 




B. g. crassirostris

Desert Finch (p. 555) Rhodospiza obsoleta



 B. g. zedlitzi




 Trumpeter Finch (p. 556) Bucanetes githaginea Eurasian Bullfinch (p. 559) Pyrrhula pyrrhula iberiae









 non-breeding Hawfinch (p. 559) Coccothraustes coccothraustes buvryi Common Crossbill (p. 562) Loxia curvirostra poliogyna  breeding Juv.

Juv.


522

6 in 15 cm

Plate 34

F. c. africana

 non-breeding







Brambling (p. 454) Fringilla montifringilla

 Chaffinch (p. 542) Fringilla coelebs


 breeding F. c. spodiogenys




Cabanis’s Bunting (p. 565) Emberiza cabanisi cabanisi 
Somali Bunting (p. 571) Emberiza poliopleura




E. f. flaviventris 

E. a. affinis

Golden-breasted Bunting (p. 567) Emberiza flaviventris

Juv.

Ad.

Brown-rumped Bunting (p. 572) Emberiza affinis 

Ad. Ad. E. a. omoensis




E. f. flavigaster

E. a. nigeriae

E. t. tahapisi 





Cinnamon-breasted Bunting (p. 574) Emberiza tahapisi E. t. goslingi

6 in 15 cm

E.t. insularis

523

Plate 35

Juv.





Rock Bunting (p. 587) Emberiza cia cia House Bunting (p. 577) Emberiza striolata

 

E. s. sahari  E. s. jebelmarrae







Socotra Bunting (p. 580) Emberiza socotrana

Ad.  E. c. reidi Juv.



Cape Bunting (p. 582) Emberiza capensis


Vincent’s Bunting (p. 584) Emberiza vincenti

Ad. Ad.

E. c. cinnamomea E. c. capensis

Ad.

Lark-like Bunting (p. 585) Emberiza impetuani


Juv.





Cirl Bunting (p. 589) Emberiza cirlus 

Pine Bunting (p. 588) Emberiza leucocephalos leucocephalos

Yellowhammer (p. 589) Emberiza citrinella citrinella

538

6 in 15 cm

Plate 36 Juv. E. c. semenowi





E. c. cineracea

Cinereous Bunting (p. 591) E. cineracea





Juv.




Juv. Ortolan Bunting (p. 592) Emberiza hortulana

Cretzschmar’s Bunting (p. 594) Emberiza caesia



Rustic Bunting (p. 595) Emberiza rustica rustica  non-breeding

 Little Bunting (p. 596) Emberiza pusilla


Common Reed Bunting (p. 597) Emberiza schoeniclus

Juv.  breeding

E. c. witherbyi  breeding E. c. schoeniclus


Ad. Juv. Corn Bunting (p. 599) Miliaria calandra Snow Bunting (p. 601) Plectrophenax nivalis nivalis

Black-headed Bunting (p. 598) Emberiza melanocephala 

 1st winter

6 in 15 cm

539

AUTHORSHIP

C. H. Fry: Petronia, Malimbus, Parmoptila, Coccopygia, Estrilda except E. thomensis, Euschistospiza, Clytospiza, Pytilia, Amadina, Ortygospiza, Paludipasser, Sporaeginthus, Amandava, Odontospiza, Spermestes, Euodice, Padda, Fringilla, Serinus, Pseudochloroptila, Alario, Neospiza, Linurgus, Rhynchostruthus, Carduelis, Carpodacus, Carpospiza, Rhodopechys, Rhodospiza, Bucanetes, Pyrrhula, Coccothraustes, Loxia, Emberiza cabanisi, E. flaviventris, E. poliopleura, E. affinis, E. tahapisi, E. striolata, E. socotrana, E. capensis, E. vincenti, E. impetuani, E. leucocephalos, Miliaria calandra and Plectrophenax nivalis2 S. Keith: most Field Characters and Voice Sections; and Nigrita, Nesocharis, Mandingoa, Cryptospiza, Spermophaga and Hypargos1 S. Keith and C. J. H. Hines: Estrilda thomensis1 S. Keith and T. B. Smith: Pyrenestes1 N. E. Collias and E. C. Collias: Ploceus cucullatus1,2 A. J. F. K. Craig: Bubalornis, Dinemellia, Plocepasser, Histurgops, Sporopipes, Pseudonigrita, Philetairus, Ploceus (except Ploceus burnieri, P. intermedius, P. velatus, P. katangae, P. ruweti and P. cucullatus), Pachyphantes, Quelea, Brachycope, Euplectes1 A. J. F. K. Craig and D. Oschadleus: D. C. Moyer:

Ploceus reichardi1

Ploceus burnieri2

D. Oschadleus: Ploceus intermedius, P. velatus, P. katangae, P. ruweti, Anaplectes and Amblyospiza1,2 R. B. Payne: Uraeginthus, Granatina, Lagonosticta, Anomalospiza and Vidua1 D. J. Pearson: most Description Sections; and Emberiza (remaining species)1 E. K. Urban: Passer1,2

Edited by 1C. H. Fry, 2S. Keith

vii

ACKNOWLEDGEMENTS Ian Hinze, Philip Hockey, L. C. Holcomb, Mark Hopkins, Kim M. Howell, Kit Hustler, Michael P. S. Irwin, Paul Isenmann, Colin Jackson, C. Jameson, Michael C. Jennings, Roland Klein, Karen Klitz, T. Konigkramer, Peter Lack, Bruno Lamarche, Marc Languy, Peter Leonard, Michel Louette, Lynda A. Lotz, D. N. Mansfield, Ross McGregor, M. B. Markus, R. P. Martins, G. Mauersberger, Bob Medland, W. Meise, Michael Mills, Ge´rard Morel, Marie-Yvonne Morel, Audrey Msimanga, David Moyer, Georgina Mwansat, P. J. Murphy, O. Nasirwa, Fleur Ngweno, M. E. D. Nhlane, Gerhardt Nikolaus, R. Nuttall, Martin Okot, G. M. Ranger, Barbara Passmore, L. L. Payne, Robert Payne, Will Peach, David Pearson, Chris Perrins, ˆs-Jones, Hugo J. Rainey, Richard Porter, Robert Pry J. Van Remsen, Robin Restall, C. Richardson, A. de Roo, Roger Safford, Otto Schmidt, K. Schuchmann, Manu Shiiwua, C. J. Skead, Neville Skinner, M. D. Sorensen, P. Spierenburg, Claire Spottiswoode, Frank Steinheimer, Terry Stephenson, Robert Stillbravy, R. Stjernstedt, J. Denis Summers-Smith, Warwick Tarboton, Michel The´venot, Danelle du Toit, Alan Tye, J. P. Vande weghe, R. F. Verheyen, J. D. R. Vernon, YvesMartin de Vivie`s, Michael Walters, Mrs F. E. (Effie) Warr, Geoff Welch, Hilary Welch, Roger Wilkinson, Jacko Williams, J. H. Williams, Diane Willis, Jared Wilson, Malcolm Wilson, Mike Wilson, Ross Wilson and H. E. Wolters. Our especial thanks go to Neil Baker, Robert Dowsett, Franc¸oise Dowsett-Lemaire and Marc Languy for providing unpublished point-plotted distribution data ˆ i and Cameroon, to Linda for Tanzania, Zambia, Malaw Birch and Mike Wilson of the Alexander Library of the Edward Grey Institute of Field Ornithology, Oxford University, for help with the literature, to Neville Brickell and Ian Hinze for advice about the biology of captive granivorous birds, to Ben Fisher for loaning material from his collection of weaver nests, and to Robert Dowsett and Franc¸oise Dowsett-Lemaire for taking a particular interest in the entire project and advising on numerous issues. For generously providing unpublished tape-recordings to help in the preparation of Voice sections, we wish to thank: National Sound Archive, London; Library of Natural Sounds, Cornell University; Tom Butynski, Peter Jones, and especially Claude Chappuis and Franc¸oise Dowsett-Lemaire. We are particularly grateful to Audrey Msimanga of the National Museum of Zimbabwe, Bulawayo, for providing copies of nest record cards under the most trying circumstances.

The manuscript of the final volume of the series was delivered to Academic Press, publishers of Vols 1–6, not long after that company’s purchase by Elsevier Science. Academic Press’s many natural history titles were put up for sale and The Birds of Africa was traded to Christopher Helm in June 2002. For a quarter century the Editors worked closely with Academic Press and for much the greater part of that time Dr Andrew Richford, Commissioning Editor at Academic, took an inspirational interest in the project. We are immensely indebted to him for his unwavering commitment, professional help and personal friendship. Put simply, The Birds of Africa might well have foundered midseries, but for Andy Richford’s enabling role. We would like to thank our editors at Christopher Helm, Nigel Redman and Mike Unwin, for steering this final volume to completion. We owe a debt of gratitude also to all authors of this volume, and indeed to all collaborators in the entire series. A work of this size could not have been contemplated, and certainly not brought to a timely conclusion, without its many contributors drawing upon the help and expertise of a small army of specialists. It says a great deal about those colleagues, biologists, birders, authors, artists, sound-recordists and other specialists and friends, that they have responded with enthusiasm and goodwill to incessant demands upon their time. For providing field observations, tape-recordings, ringing and nestrecord data, weights, longevities, information on particular species or regions, mapping data, museum specimen data printouts, taxonomic advice, reprints, translations, contacts and hospitality, we are much indebted to Marc Adams, D. Allan, David Allen, Tony Archer, John S. Ash, Graeme Backhurst, G. Balanc¸a, Mindy Baha el Din, Sherif Baha el Din, Liz Baker, Richard Banks, Clive Barlow, the late Luis F. Baptista, C. R. Barlow, P. Barnard, Leon Bennun, Aldo Berruti, Linda Birch, Mark Boorman, Nik Borrow, Neville Brickell, Peter Britton, Joost Brouwer, W. F. Bruce-Miller, Tom Butynski, A. Camara, Paul G. Cardy, Robin Chancellor, Ruth T. Chapin, Claude Chappuis, Patrice Christy, Peter Clement, Peter Colston, Adrian J. F. K. Craig, Will Cresswell, T. Crisler, Sandra Dantu, W. Richard J. Dean, Ron Demey, D. Drinkrow, Robert J. Dowsett, Franc¸oise Dowsett–Lemaire, Steve Dudley, Mike Dyer, Gus Ezealor, John Fanshawe, Ben Fisher, Lincoln Fishpool, Roger Fotso, K. Garrett, Steven Goodman, Robert Gray, Andrew Grieve, K. D. Groschupf, Mrs Dale Hanmer, David Happold, Marc Herremans, C. Herrmann, P. Herroelen, Chris Hines, Janet Hinshaw,

viii

ACKNOWLEDGEMENTS

For access to museum and library holdings, we are most grateful to administrators, scientists, curators and librarians of: Aberdeen University Department of Zoology; American Museum of Natural History Department of Ornithology; Augusta State University Department of Biology; Avian Demography Unit, University of Cape Town; Carnegie Museum of Natural History, Pittsburgh; Durban Natural History Museum; Edward Grey Institute Library, Oxford; Field Museum of Natural History, Chicago; l’Institut pour la Re´cherche Scientifique en Afrique Centrale; Josselyn Van Tyne Library, Wilson Ornithological Society; Liverpool Museum of Natural History; Los Angeles County Museum (Bird Division); Muse´e Nationale d’Histoire Naturelle, Paris; Muse´e Royale de l’Afrique Centrale, Tervuren; Museum of Natural Sciences, Louisiana State University; Namibian Scientific Society; National Museums of Kenya, Nairobi; National Museum of Zimbabwe, Bulawayo; National Museums of Scotland, Edinburgh; Natural History Museum (London) Department of Ornithology; Niven Library, Percy FitzPatrick Institute of African Ornithology; South African Museum, Cape Town; Transvaal Museum (Bird Division), Pretoria; US National Museum of Natural History, Washington; Western Foundation of Vertebrate Zoology; and Zoological Society of London. As in earlier volumes, Martin Woodcock researched, designed and executed the colour plates and painted the jacket portraits; we take particular pleasure in thanking him for his consummate skill, good-natured collaboration and unwavering support. Ian Willis prepared most of the line drawings, again as in previous volumes, and we extend our warm gratitude to him too. We are well aware that the beautiful work of both artists has enhanced this volume and the entire series immeasurably. Other line drawings, by Hilary Fry, are marked (*) below. Drawings are from photographs, specimens, field sketches, and diagrams and drawings in the literature. Principal sources, identified by bold numbers in the following acknowledgements listing, are: 1 Birdlife in Southern Africa (K. Newman, 1979, Macmillan South Africa, Johannesburg); 2 The Birds of the Belgian Congo (Pt 4, J. P. Chapin, 1954, Bull. Amer. Nat. Hist 75B); 3 Birds of Kenya and Northern Tanzania (D. A. Zimmerman, D. A. Turner and D. J. Pearson, 1996, Christopher Helm, London) (drawings by D. A. Zimmerman); 4 The Birds of the Western Palearctic (eds S. Cramp and C. M. Perrins, Vol. 8, 1994, Oxford Univ. Press, Oxford); 5 Birds of the World (ed. J. Gooders, 1971, IPC, London); 6 The Complete Book of Southern African Birds (P. J. Ginn, W. G. McIlleron and P. le S. Milstein, 1989, Struick Winchester, Cape Town); 7 The Encyclopaedia of Birds (eds C. M. Perrins and A. L. A. Middleton, Vol. 3, 1985, Unwin Animal Library, London); 8 Ben Fisher, weaver nest collection; 9 Nesting Birds (P. Steyn, 1996, Fernwood Press, Vlaeberg); 10 R. B. Payne, photos of

estrildid and viduid nestling mouthparts; 11 World of Wildlife (F. R. de la Fuente, 1971–1972, Orbis, London). Passer melanurus, photo by P. Steyn in 9; P. domesticus, drawing by R. Gillmor in 4, photo by D. Zingel in 5; P. hispaniolensis, photo by H. Schrempp in 5; P. luteus, drawings by R. Gillmor in 4; Petronia dentata, drawing by J. Trollope in Avicult Mag. 83, 1977 (*); P. petronia, drawing by R. Gillmor in 4; Bubalornis albirostris, drawing in 3; B. niger, photo by P. J. Ginn in 6; Dinemellia dinemelli, drawing in 3; Plocepasser mahali, photo in 2, photo by P. Steyn in 9; P. superciliosus and P. donaldsoni, drawings in 3; Sporopipes squamifrons, photo by P. Johnson in 5; Pseudonigrita arnaudi, photo by J. Pato in 11, drawings in 3; P. cabanisi, photo by J. Pato in 11, drawing in 3, photos by Ian Willis; Philetairus socius, photo by P. Steyn in 9, drawings by E. C. Collias and N. E. Collias in Ibis 120, 1978, photo by J&B Photographers in Africa Birds and Birding 3 (4), 1998; Malimbus ballmanni, painting by F. Weick in W. Gatter, Birds of Liberia, 1977 (*); M. cassini, photo by A. Devez in A. Brosset, La Vie dans la Foreˆt Equatoriale, 1976, photo by A. Brosset in A. Brosset, Ibis 120, 1978; M. coronatus and M. malimbicus, photos by A. Brosset in A. Brosset, Ibis 120, 1978; M. nitens, photo by A. Devez in A. Brosset, La Vie dans la Foreˆt Equatoriale, 1976, photo by A. Brosset in A. Brosset, Ibis 120, 1978; M. rubricollis, drawing in 3; Anaplectes rubriceps, photo by Jacana in 7, photo in V. G. L. van Someren, Days with Birds, 1956, photo by C. Haagner in P. A. R. Hockey, Birds of Southern Africa, 1991, photo by P. Steyn in 9; P. baglafecht, photo by J. P. Scott in A Souvenir Guide Book to African Birds, Kensta, Nairobi, 1990; P. luteolus and P. pelzelni, drawings by W. E. Almond in J. H. Crook, Ibis 102, 1960 (*); P. nigricollis, photo by A. Devez in A. Brosset, La Vie dans la Foreˆt Equatoriale, 1976, drawing in 3; P. ocularis, photo by D. McCulloch in 1, photo in 2, photo by P. Steyn in 9; P. capensis, photos by P. Steyn in 9; P. temporalis, 8; P. subaureus, photo by F. Hartmann in The Living World of Animals, 1970, Reader’s Digest Association, photo by P. Steyn in 9; P. xanthops, photo by P. J. Ginn in 6; P. aurantius, drawing by W. E. Almond in J. H. Crook, Ibis 102, 1960 (*); P. princeps, both 8; P. xanthopterus, illustration in G. R. McLachlan and R. Liversidge, Roberts’ Birds of South Africa, 1978; P. castanops, drawing in 3; P. taeniopterus, photo in 2; P. intermedius, photo by W. Nichol in 6, photos by P. Steyn in 9; P. velatus, photo by C. Haagner in P. A. R. Hockey, Birds of Southern Africa, 1991; P. vitellinus, drawing by W. E. Almond in J. H. Crook, Ibis 102, 1960 (*); P. heuglini, drawing in 3; P. galbula, photo in H. and J. Eriksen, Birdlife in Oman, 1999, halved nest supplied by C. H. Fry (*); P. spekei, 8; P. nigerrimus, photo by P. Gatter in W. Gatter, Birds of Liberia, 1997; P. cucullatus, A, painting by P. A. Clancey in Ibis 130, 1988 (*), B, photo in C. Stuart and T. Stuart, Birds in Africa from Seabirds to Seed-eaters,

ix

x

ACKNOWLEDGEMENTS

1999, C–F, drawings supplied by N. E. Collias, G, anon. photo in 11; P. grandis, 8; P. jacksoni, photo by S. Trevor in 11, drawing in 3; P. rubiginosus, photo in W. Tarboton, A Guide to Nests and Eggs of Southern African Birds, 2001, drawings in 3; P. bicolor, photo by P. Steyn in 9; P. olivaceiceps, photo by M. Rees in Africa Birds and Birding, 3 (1) 1998, and 8; P. angolensis and P. sanctithomae, 8; P. manyar, photo in 5; Pachyphantes superciliosus, drawing in 3; Amblyospiza albifrons, photos by P. Steyn in 9, photo by G. L. Maclean in 1; Quelea quelea, photo by P. Johnson in As Free as a Bird, 1976, photo by P. Steyn in 9; Euplectes orix, photo by W. G. McIlleron in 6, photo by G. J. Broekhuysen in B. Campbell, The Dictionary of Birds in Colour, 1974, photo by P. Steyn in 9, photo by K. B. Newman in 5; E. hordeaceus, photo in 2; E. afer, photo by R. M. Bloomfield in 5, photo by W. G. McIlleron in 6; E. capensis, photo by P. Steyn in 9; E. macrourus, photo in 2; E. ardens, photo by W. T. Miller in 1, photo by V. G. L. van Someren in V. G. L. van Someren, Days with Birds, 1956; E. progne, photo by J. S. Wightman in 5, photo by P. G. Johnson in 1, photo by W. R. Tarboton in 6; E. jacksoni, photo by J. F. Reynolds in B. Campbell and E. Lack (eds) A Dictionary of Birds, 1985, photo by V. G. L. van Someren in H. Friedmann, Smithson. Rep. (1949), 1950 (*), photo by W. T. Miller in 3; full page figures of mouth markings of young estrildids and viduids, all 10 except Nigrita canicapilla, field sketch by C. H. Fry (*), and Sporaeginthus subflavus, drawing in B. D. Colahan, Ostrich 53, 1982 (*); Parmoptila woodhousei, painting by P. Burton in 5; E. astrild, photo by Cape Bird Club in 6, photo by P. Steyn in 9; Pyrenestes ostrinus, photos by T. B. Smith in Auk 107, 1990 (*), photo supplied by M. de Vivie`s (*); Uraeginthus angolensis, drawing by D. Goodwin in Ibis 107, 1965 (*); U. bengalus, photo in 2; U. cyanocephalus, drawing in D. Goodwin, Ibis 107, 1965 (*); Granatina ianthinogaster, photo in J. Nicolai, J. Orn. 110, 1969 (*); G. granatina, photos by J. Nicolai in Z. Tierpsychol. 21, 1964 (*), painting in J. Nicolai, Z. Tierpsychol. 25, 1968 (*); Euschistospiza cinereovinacea, photo by W. Baars in Gefiederte Welt, 1967, 81 (*); Hypargos niveoguttatus, both 10 ; H. margaritatus, photo by C. Scholtz in Cage & Aviary Birds, Jan. 1998 (*); Clytospiza monteiri, photo by R. Neff in Gefiederte Welt 1977, 181 (*), and 10; Pytilia hypogrammica and P. melba, 10; Lagonosticta senegala, drawing in 3, photo by M. -Y. Morel in Me´m. Mus. Nat. Hist. Nat. Se´r. A, 78, 1964 (*), and 10; L. rubricata, both 10; Ortygospiza atricollis, photo by P. Steyn in 9; O. fuscocrissa, drawing in R. J. Nuttall, Ostrich 63, 1992 (*); Sporaeginthus subflavus, 10, photo by V. G. L. van Someren, Days with Birds 1956; Anomalospiza imberbis, photo by P. Steyn in 9; Vidua macroura, photo by J. Nicolai, Z. Tierpsychol. 21, 1964 (*), drawings in P. Shaw, Ibis 126, 1984 (*); V. obtusa, drawing in J. Nicolai, Z. Tierpsychol. 21, 1964 (*); V. paradisaea, paintings by M. A. Castan ˜ os in 11,

photo by R. A. C. Jensen in 6, drawings in J. Nicolai, J. Orn. 110, 1969 (*), photo in J. Nicolai, Z. Tierpsychol. 21, 1964 (*); V. chalybeata, 10; V. wilsoni, photo by H. Kacher in J. Orn. 113, 1972 (*); Serinus citrinelloides, (*), V. G. L. van Someren in V. G. L. van Someren Days with Birds, 1956; S. capistratus, (*); S. citrinipectus, photo in B. Mayer, Die Voliere 15, 1992 (*), photo by H. Classen in R. van den Elzen, H. Classen and R. Schon, Trochilus 11, 1990 (*); S. mozambicus, S. donaldsoni and S. buchanani, (*); S. sulphuratus, (*), photo in C. Stuart and T. Stuart, Birds in Africa from Seabirds to Seed-eaters, 1999, photo by J. van Jaarsveld in P. A. R. Hockey, Birds of Southern Africa, 1991; S. flaviventris, (*), photo by P. Steyn in 9; S. albogularis, (*), S. a. crocopygia, (*), photo by J. Frandsen, Birds of the South Western Cape, 1982 (*); S. burtoni, skin (*), photo by H. Classen in R. van den Elzen, H. Nemeschkal and H. Classen, Bonn. Zool. Beitr. 38, 1987 (*); S. rufobrunneus, (*); S. leucopterus, photo by J. Frandsen, Birds of the South Western Cape, 1982; S. gularis, photo by C. J. Skead, The Canaries, Seedeaters and Buntings of Southern Africa, 1960 (*), photo by P. Steyn in 9; S. canicollis, photo by H. Classen in R. van den Elzen, H. Nemeschkal and H. Classen, Bonn. Zool. Beitr. 38, 1987 (*), photo by P. Steyn in 9, photo by J. Frandsen, Birds of the South Western Cape, 1982; S. nigriceps, (*), photo by H. Shirihai in J. Francis and H. Shirihai, Ethiopia In Search of Endemic Birds, 1999 (*); Pseudochloroptila totta, photo by P. Steyn in 9; Alario alario, photo by R. E. Viljoen in 6, photo in J. Frandsen Birds of the South Western Cape, 1982; A. leucolaemus, (*); Neospiza concolor, (*); Rhynchostruthus socotranus, (*); Carduelis chloris, drawings by D. Nurney in 4 (*); C. ankoberensis, (*); Carpospiza brachydactyla, photo by H. & J. Eriksen and D. Robinson in G. Kirwan, Sandgrouse 20, 1998 (*); Rhodopechys sanguinea, sketchbook drawings by Ian Willis; Coccothraustes coccothraustes, (*); Loxia curvirostra, drawing by D. Nurney in 4; Emberiza cabanisi, (*); E. flaviventris, (*), anon. photo in 11, photo by P. Steyn in 9; E. poliopleura, (*); E. affinis, (*); E. striolata, photo by J. Eriksen in C. H. Fry and J. Eriksen, Brit. Birds 87, 1994 (*); E. socotrana, photo by C. Barrau in M. Clouet, J.-L. Goar and C. Barrau, Alauda 66, 1998 (*); E. capensis, photo by P. Steyn in 9; E. impetuani, photo by P. Steyn in 9; E. cia, photo by K. J. Carlson in 5; E. schoeniclus, drawing by D. Nurney in S. Cramp and C. M. Perrins (eds) The Birds of the Western Palearctic 9, 1994; and Miliaria calandra, anon. photo in 11. We have benefitted greatly from the wealth of information in recently published books. We would like to mention in particular the southern African atlas (Harrison et al. 1997), books on the birds of Algeria (Isenmann and Moali 2000), Mauritania and Mali (Lamarche 1993–1994), The Gambia (Barlow et al. 1997), Liberia (Gatter 1997), Togo (Cheke and Walsh 1996), Gabon (Christy and Clarke 1994), Sa˜o Tome´ and

ACKNOWLEDGEMENTS

Prı´ncipe islands (Christy and Clarke 1998), western Africa (Borrow and Demey 2001), Somalia (Ash and Miskell 1998), eastern Africa (Zimmerman et al. 1996, Stevenson and Fanshawe 2002), Angola (Dean 2000) and southern Mozambique (Parker 1999); and on nesting birds (Steyn 1996, Tarboton 2001), cage and aviary birds (Brickell and Konigkramer 1997) and important bird areas (Byaruhanga et al. 2001, Fishpool and Evans 2001). Some monographs are not so recent yet have been no less valuable: on sparrows (SummersSmith 1988), mannikins (Restall 1996), estrildids (Goodwin 1982), estrildids and finches (Clement et al. 1993) and buntings (Byers et al. 1995). In related fields, we take common and scientific names of mammals from Kingdon (1997) and plant scientific names from Mabberley (1993). B. Lamarche’s erudite bird atlas for Mauritania and Mali (1993–1994), in 8 volumes totalling 2350 pages, seems to have been overlooked by much of the bird world until a set found its way to the Edward Grey Library, Oxford, early in 2001. We wish that we had been able to benefit from its wealth of detail in our

July 2002

By rotation, responsibility for this volume should have fallen principally to Stuart Keith, but for reasons of health he asked Hilary Fry to shoulder the task. We wish to express here our gratitude to and appreciation of Hilary for the great amount of time and effort that he has put into the entire project, especially into the final

January 2003

earlier volumes, but in this one we have drawn heavily upon it and are deeply indebted to its author. The Birds of Morocco (M. The´venot, R. Vernon and P. Bergier, B.O.U., 2003) appeared too late for us to be able to incorporate its wealth of information. We are more than ever conscious of reliance on the works of the great naturalists of old, G. L. Bates, J. P. Chapin, R. E. Moreau, C. J. Skead, W. Serle, V. G. L. van Someren, A. W. Vincent, J. Vincent, J. M. Winterbottom and others. There are those today in Africa whose fieldcraft matches that of the old-timers, but they are few. Hilary Fry wishes to record here his sincere thanks to his patient wife Kathie for her continual support, understanding and encouragement as well as for much practical assistance. Sallyann Keith, Lois Urban, Diane Willis, Barbara Woodcock, and doubtless the wives and families of other contributors, have had their lives variously affected by the demands of this project, and we thank them all for their forbearance.

C. Hilary Fry Stuart Keith

two volumes, in which he has undertaken the lion’s share of authorship, has edited hundreds of species accounts through multiple drafts, prepared nearly all of the maps, integrated the line artwork, and seen the manuscript through production.

Stuart Keith Emil K. Urban

xi

The Birds of Africa I–VII Vol. I (1982). L. H. Brown, E. K. Urban and K. Newman. Pp xiii þ 521; 32 plates by M. Woodcock and P. Hayman. Academic Press, London. 269 species: Ostrich; albatrosses, shearwaters, petrels; penguins; divers; grebes; tropicbirds, boobies, cormorants, darters, pelicans, frigatebirds; herons, Hamerkop, storks, Shoebill, ibises; flamingoes; ducks; hawks, Secretarybird, falcons. Vol. II (1986). E. K. Urban, C. H. Fry and S. Keith (eds). C. W. Benson, F. M. Benson, P. L. Britton, A. Brosset, L. H. Brown, N. J. Collar, T. M. Crowe, C. H. Fry, P. D. Goriup, P. A. R. Hockey, T. R. Howell, S. Keith, G. L. Maclean, G. J. Morel, M.-Y. Morel, D. J. Pearson, P. E. Osborne, P. B. Taylor, E. K. Urban. Pp. xvi þ 552; 32 plates by Martin Woodcock. Academic Press, London. 298 species: francolins; button-quail, rails, cranes, African Finfoot, bustards, jacanas, Painted Snipe, Crab Plover, oystercatchers, stilts, thick-knees, Egyptian Plover, coursers, plovers, lapwings, sandpipers, phalaropes, skuas, gulls, terns, African Skimmer, auks; sandgrouse; doves. Vol. III (1988). C. H. Fry, S. Keith and E. K. Urban (eds). C. W. Benson, F. M. Benson, A. Brosset, G. R. Cunningham-van Someren, N. C. Davidson, J. P. Decoux, C. H. Fry, R. M. Harwin, J. F. M. Horne, M. P. S. Irwin, A. C. Kemp, D. Ligon, L. L. Short, S. N. Stuart, M. P. Wilson, R. T. Wilson. Pp. xvi þ 611; 32 plates by Martin Woodcock. Academic Press, London. 312 species: parrots; turacos; cuckoos; owls, barn owls; nightjars; swifts; colies; trogons, kingfishers, bee-eaters, rollers, wood-hoopoes, Hoopoe, hornbills; barbets, honeyguides, woodpeckers. Vol. IV (1992). S. Keith, E. K. Urban and C. H. Fry (eds). W. R. J. Dean, C. Erard, C. H. Fry, V. Haas, R. A. C. Jensen, S. Keith, P. C. Lack, T. B. Oatley, D. J. Pearson, S. N. Stuart, P. B. Taylor, A. Tye, E. K. Urban. Pp. xv þ 609; 32 plates by Martin Woodcock. Academic Press, London. 303 species: broadbills; pittas; larks; swallows, martins; wagtails, pipits, longclaws; cuckoo-shrikes; bulbuls; Bohemian Waxwing, Grey Hypocolius; White-throated Dipper; Winter Wren; accentors; robins, alethes, robin-chats, wheatears and relatives. Vol. V (1997). E. K. Urban, C. H. Fry and S. Keith (eds). C. Erard, C. H. Fry, L. G. Grimes, M. P. S. Irwin, S. Keith, P. C. Lack, R. de Naurois, D. J. Pearson, A. Prigogine, A. Tye, E. K. Urban. Pp. xix þ 669; 32 plates by Martin Woodcock. Academic Press, London. 309 species: thrushes; warblers, cisticolas, prinias, apalises, eremomelas and relatives; flycatchers; paradise-flycatchers and monarchs; shrike-flycatchers, wattle-eyes and batises. Vol. VI (2000). C. H. Fry and S. Keith (C. H. Fry, S. Keith and E. K. Urban, series eds). A. J. F. K. Craig, C. H. Fry, L. Grimes, M. P. S. Irwin, S. Keith, D. J. Pearson, E. K. Urban, D. Wiggins, R. Wilkinson. Pp. xvii þ 724; 36 plates by Martin Woodcock. Academic Press, London. 334 species: picatharteses; babblers; Long-tailed Tit; tits; penduline tits; nuthatches; Spotted Creeper; tree-creepers; sunbirds; white-eyes; sugarbirds; shrikes; bush-shrikes; Black-collared Bulbul, nicators; helmet-shrikes; orioles; drongos; crows; starlings and oxpeckers. Vol. VII (2004). C. H. Fry and S. Keith (C. H. Fry, S. Keith and E. K. Urban, series eds). E. C. Collias, N. E. Collias, A. J. F. K. Craig, C. H. Fry, C. J. H. Hines, S. Keith, D. Moyer, D. Oschadleus, R. B. Payne, D. J. Pearson, T. B. Smith, E. K. Urban. Pp. xxii þ 666; 36 plates by Martin Woodcock. Christopher Helm, London. 309 species: sparrows; weavers, malimbes, bishops, widowbirds; waxbills, firefinches, pytilias, mannikins and relatives; whydahs, indigobirds; finches, canaries and relatives; buntings.

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INTRODUCTION The project had its origins in the mid-1960s when Leslie Brown, Kenya’s Chief Agriculturist and a field ornithologist with numerous books and papers already to his credit, thought of writing a definitive handbook on Africa’s avifauna. It would deal with the resident, visiting and vagrant birds of the continent and its offshore islands, and be the modern successor to Reichenow’s Die Vo¨gel Afrikas (1900–1903), Sclater’s Systema Avium Aethiopicarum (1928–1930) and Mackworth–Praed and Grant’s African Handbook of Birds (1957–1973). Emil Urban was invited to co-author the work in 1967, and the two developed ideas on field trips and reciprocal visits between Addis Ababa, Ethiopia, and Karen, Kenya. Finding a suitable publisher took a decade until Academic Press came to the rescue, proposing in 1976 to publish the work in two volumes. Negotiations continued into 1978 when a handbook of four volumes was agreed upon, and writing began in earnest late in September that year. Meanwhile, Leslie Brown’s health had been deteriorating, and he died in 1980. It necessitated some re-organization of Volume I, published in 1982, and restructuring of the remainder as an edited, multi-author, series. Hilary Fry and Stuart Keith, members of the first volume’s Advisory Editorial Board, were appointed editors jointly with Emil Urban. The four volumes were to have comprised two nonpasserine and two passerine ones, but the editors realized soon after starting Volume II that, since Volume I had got only as far as the falcons, it would not be practicable to compress the rest of the non-passerines into a single volume. They persuaded Academic Press to expand the work to six volumes, three for non-passerines and three for passerines. Although there are far more passerine than non-passerine species, it was thought that more was known about non-passerines, mostly large and conspicuous birds, so the two parts should be of about the same length. Soon after researches for Volume IV began, however, it transpired that far more is in fact known about the relatively small, inconspicuous passerines than had been anticipated. Once again Academic Press agreed to an increase, and with the present seventh volume the series is at last completed. Our objectives are to integrate salient facts from field, literature and museum and to produce a compendium characterising the lives and biology of every species. Originally, we tried also to identify topics and taxa in need of further research and indeed we still do so, but have been somewhat overtaken by events. These are the accelerating pace of change across the face of Africa and the consequent imperative for nature conservation, now led by BirdLife International; increasing tourism and the resulting improvement in knowledge about bird identification and distribution; the growth in ornithological institutions and birding activities, the latter spectacularly in South Africa; and an exponential increase in publications. This 25-year

project has spanned an era of dramatic developments in information and communication technology. Globally, there has been great progress in molecular biology, which has brought about a new understanding of systematics and taxonomy (relationships between species, and classification). The very nature of species is being constantly re-evaluated. In trying to keep abreast of such changes and discoveries and ever-improving knowledge about Africa’s birds, progressive material and presentational changes have been introduced with each volume, in such features as species account length, text figures, plate keys, colour maps, mapping precision, voice transcriptions, reference systems and indexing. Contractual deadlines and cajoling notwithstanding, it became increasingly evident years ago that production was as slow as the slowest contributor, and successive interims between volumes from II to V were of 2, 4 and 5 years. To speed matters up, procedural reforms were introduced after V appeared. In VI and VII sectional or ‘horizontal’ as well as systematic or ‘vertical’ research and authorship have been employed, with Stuart Keith responsible for most of the Voice and Field Characters sections and David Pearson for most Descriptions. With the appointment of a single Executive Editor, Volume VI was able to appear only 3 years after V and the present volume was delivered to the publishers a mere 21 months after the publication of VI. About 2130 bird species have been covered – between one-fifth and one-quarter of the world’s bird fauna. The three non-passerine volumes have on average about 1.6 text pages per species and the four passerine ones nearly 2.0. It results from the lifting, half way through the series, of the Publisher’s constraint on average lengths of species accounts and from burgeoning knowledge. Were the volumes to be written anew, they would have to deal with probably well over 2250 species, some new birds having been found and the contemporary taxonomic trend being to upgrade many subspecies to the status of full species. In this volume the literature has been reviewed fully up to August 2001. Much additional information from the current literature was added up to the time of manuscript delivery in March 2002. The Publishers kindly allowed several species accounts to be updated to November 2002, when production began, and a few to March 2003. In outline, the publishing history, contributors, and contents of the complete series of The Birds of Africa are shown in the box opposite.

Vegetation and Topography Africa’s vegetation was discussed in Vol. I, pp 4 –11 (Brown et al. 1982), with accompanying essays placing it in the context of the geological past, climate, topography, and bird habitats. Figure 1 reproduces in colour the map in

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INTRODUCTION

Vol. I, after Keay (1959), and shows the principal lowland vegetation types. The subdesert southern fringe of the Sahara is termed the Sahel, with sahelian savannas; in West and N-central Africa, dry wooded savannas comprise the soudanian zone and, further south, the northern guinean zone, and moist savanna woodlands comprise southern guinean and derived savanna zones, the latter bordering coastal rain forest; we use these terms commonly. The maps are the same size as species-distribution maps and superspecies maps throughout this book so that, by comparison, the degree to which species have ranges determined by vegetation can readily be appreciated. Figure 2 shows the principal montane regions and rivers. Montane forest and other mountain vegetation types above the treeline provide wildlife habitats as distinctive as the main lowland types; also important are aquatic habitats – lakesides, rivers, marshes, estuaries, mangroves – and manmade habitats like towns and gardens. At this scale each vegetation zone appears to be a vast, continuous swath.

Boundaries in Figure 1 are more theoretical than real; the inset shows the actual extent of closed-canopy rain forest in West Africa in the mid-1990s, and even that may be optimistic (Gatter 1997).

Metres 500–1000 1000–1500 Over 1500

Fig. 2.

Topographic features.

Distribution Maps

Macchia Desert and subdesert Dry wooded savannas

Moist wooded savannas Lowland tropical rain forest Montane

Fig. 1. Notional boundaries of major African vegetation zones. An enlargement of West Africa shows recent fragmentation of rain forest (adapted from Gatter 1997).

Agricultural and industrial development and burgeoning human populations are affecting much of Africa. Woodland, wetland and other habitats have shrunk and been degraded, diminishing birds’ ranges and putting some montane forest endemics at risk of extinction. Conversely, many farmland and garden species have expanded their ranges. We generally remark such range shifts in the RANGE and STATUS sections, but have made little attempt to map them; rather, maps purport to show contemporary ranges. However, ranges remain sketchy in some parts where famine or civil unrest have precluded field studies, and for such areas our maps are often based on little more than the point-plotted distributions of Hall and Moreau (1970). Conversely, this book benefits greatly from bird atlases prepared in recent years for many countries, the most detailed being for Mauritania, Mali, Tanzania, ˆ i and the whole of southern Africa. Zambia, Malaw The Mauritania/Mali atlases (Lamarche 1993–1994) extend the ranges of many lowland rain forest species hundreds of kilometres northward from central Guinea into southern Mali. Several of the localities concerned were revisited by R. Dowsett and F. Dowsett-Lemaire in February 2002; they investigated several gallery forests (where they discovered Dybowski’s Twinspot and Black-

INTRODUCTION

bellied Firefinch, new to Mali) but were unable to find any trace of rain forest; indeed, there seems to be no knowledge that rain forest has ever existed there. Rain forest birds are thought to be sedentary and their occurrence in Mali, even as wind-blown vagrants, is unexpected, so it will be pleasing if the records can be corroborated. The species concerned in this volume are malimbes, negrofinches and some other estrildids.

? ?

? ?

Fig. 3.

Breeding or resident Migration Dense non-breeding Sparse non-breeding Occurrence uncertain Breeding uncertain Isolated records Isolated breeding Isolated occurrence

?

Shading and symbols on species maps.

Figure 3 shows the colour and shading conventions used in species maps. Superspecies maps are simplified representations of the ranges of their constituent species and are intended to show areas of marginal overlap or sympatry, parapatric boundaries, and the extent of allopatry. The political map of Africa is in Figure 5 on p. xxi

Sedentary, Resident and Migrant Birds A sedentary species, population or pair of birds is one that remains in the same locality for years on end, individuals roaming no more than a few kilometres at the most. A species or population is resident when it occurs year-round in the same area and breeds there, even though the same individuals may not be in the same area all year round. A species resident throughout most of its range is a partial migrant if it occurs regularly towards the edges of its range only at given seasons. If that part of its range where it is resident is small in comparison with adjacent parts where it appears only seasonally, the species can be described as an intra-African migrant rather than a partial migrant; a species that breeds in one part of Africa and ‘winters’ in quite a different one is unquestionably an intra-African migrant. Species that breed in Europe or Palearctic Asia and winter in Africa are Palearctic migrants, whether or not they also breed in Palearctic North Africa.

Systematics and Taxonomy Hitherto, the traditional evolutionary taxonomy with its ‘biological species concept’ or BSC, championed by Mayr (1942), has been used widely by ornithologists. During the life of The Birds of Africa, evolutionary taxonomy has been challenged by alternative classification methodologies and the BSC by alternative species concepts (such as evolutionary, recognition and phylogenetic ones). In particular, the phylogenetic species concept or PSC has received considerable advocacy; it makes the species the smallest irreducible unit, without subspecies, and in effect upgrades well-defined subspecies, or most kinds of them, to the status of full species. There are theoretical and practical difficulties with the PSC, however, and it has been argued persuasively that the BSC appeals better to common sense, serves ornithology best, and is likely to remain the most valuable and popular species concept among field ornithologists for a good many years to come. However, as birds become better and better known, proposals for multiplying species are becoming ever more pressing, particularly by raising island and ‘habitat-island’ populations to species level. Amongst European birds, recent taxonomic changes have led to the formal recognition of no less than 18 additional species (Sangster et al. 2002), and the trend is accelerating in Africa too. Continuing debate about the nature of species has led to detailed guidelines for taxonomic treatment (Collinson 2002; Helbig et al. 2002 and references therein). Systematic treatment is based largely upon the works of White (1963), Hall and Moreau (1970), Sibley and Monroe (1990) and Dowsett and Forbes-Watson (1993), all of whom use the BSC. The treatment of Estrildidae and Viduidae stems from the mitochondrial DNA studies of M. Sorenson and R. B. Payne (pers. comm.). We continue to use subspecies (or races) and species in the conventional manner, but have seen fit to raise several populations that have usually been regarded as regional subspecies, to the rank of species. A group of evidently very closely allied populations, with geographical ranges that are separate or abutting, composes a superspecies. An example of pitfalls in inferring proximity of relationship from overt similarities of plumage and voice is given in the Chaffinch TAXONOMIC NOTE on p. 453, and the systematics of brood-parasitic viduines, as well as relationships with their hosts, are particularly intriguing (Payne et al. 2002). The concept of superspecies, discussed in Vol. II, xi–xii, is invaluable in treating continental avifaunas but, like other taxonomic ranks, it is not without its difficulties, as noted in Vols III, xiii and IV, x–xi. A rule-of-thumb definition of a superspecies is that its members are regarded by some taxonomists as subspecies of a single species and by others as full species. Ideally its members should not overlap geographically, but a limited amount of marginal overlap or sympatry may be allowed in some instances. In practice, whether overlap is real, involving breeding birds rather than vagrants, is often uncertain; and how abutting (parapatric) populations (paraspecies) interact where they meet or just overlap has almost never been adequately studied: are they separated altitudinally

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INTRODUCTION

1 2

Passer griseus superspecies 1 2 3 4 5

4

P. griseus P. swainsonii P. gongonensis P. suahelicus P. diffusus 1, 4 or 5, not identified

3

1

Range overlaps: griseus/diffusus griseus/swainsonii, griseus/suahelicus gongonensis/swainsonii, gongonensis/suahelicus griseus/gongonensis/suahelicus (small region in NE Tanzania)

5

Fig 4. Ranges of grey-headed sparrows: see text.

or by habitat, are they interspecifically territorial, what is the rate of hybridisation and what is the demography of gene flow? In Vol. VI, xiv–xv, the biogeography and putative differentiation of Telophorus bush-shrikes was presented in relation to taxonomy. Here we illustrate the application of the superspecies concept – as well as difficulties frequently encountered with mapping range boundaries – by reference to sparrows. 19 species are resident in Africa, 14 being endemic. Of the 14, the five rufous sparrows of the Passer motitensis complex have separate, relatively small ranges (mapped on p. 2). They were formerly thought to be subspecies of a single species; but improving knowledge emphasizes their small differences rather than their similarities and they are currently widely regarded as five separate species composing a superspecies (a sixth being P. iagoensis of the Cape Verde Islands). Figure 4 shows the distributions of another group of five, the grey-headed sparrows. They too were formerly treated as a single species, P. griseus, until it became known that there is limited overlap near the breeding range boundaries of the various populations. These zones of sympatry are detailed in the Passer diagnosis on pp. 1–2. In some areas, two or even three different grey-headed sparrow taxa occur together without interbreeding, but in others the same taxa apparently hybridize. For instance, griseus and diffusus are

widely sympatric in Zambia north and west of L. Kariba, without hybridizing, but along the Zambezi, east of L. Kariba and well into Mozambique, they have been held to hybridize commonly (Hall and Moreau 1970), although an alternate explanation is that diffusus there is part of the same colour/size/bill-size trend that produced the population luangwae in Zambia’s Luangwa Valley (Irwin 2002). In areas of overlap they behave as distinct species but, if they do intergrade on the Zambezi, they would fail to meet one criterion of good species (Helbig et al. 2002). The taxa all meet another species-criterion of diagnosability in the museum, but delineating their ranges is not made any easier by problems with field identification, as in Tanzania (Figure 4). M. P. S. Irwin is studying the provenance and spread of Northern and Southern Grey-headed Sparrows in Zambia and Zimbabwe (Irwin 2002 and pers. comm.). He concludes that the northern P. griseus originated in arid, northern sahelian and soudanian savannas and started to spread about 7,000 years ago, in close association with agriculture, villages and later towns, as far south now as Harare, Zimbabwe. 100 years ago in Zambia the southern P. diffusus was confined to the Zambezi and Luangwa valleys, but 70 or 80 years ago it moved into southern plateau areas, only recently becoming sympatric with P. griseus. P. diffusus is not an urban bird and Irwin speculates that it may not be quite as closely allied to P. griseus as appear-

INTRODUCTION

ances suggest. It is quite possible that one or both of these sparrows are evolving as their ranges extend, like the House Sparrow P. domesticus in newer parts of its world range. Irwin is convinced that swainsonii and suahelicus are conspecific, so he counts four species of grey-headed sparrows. In treating the complex as a superspecies, whether of four or five species, our intention is not to take the easy way out of taxonomic controversy so much as to point the way to further investigation. It may prove that vocal, behavioural, social and even genetic/morphological differences are greater than they presently seem and that the species have differentiated too far even for acceptable accommodation in a single superspecies – or contrarily that genetic merging will occur as P. griseus penetrates the ranges of other taxa and swamps them; grey-headed sparrows would then constitute only a single, polytypic species after all.

Vernacular and Scientific Names; Species Sequence

Age classes and terms for the various feather generations have been defined by Jenni and Winkler (1994) and others. Nomenclature is complicated by different species having different moult regimes and plumage sequences; by complete, partial and interrupted moults; and by some birds breeding in immature plumage. Our descriptions of age classes are often greatly hampered by the poor state of knowledge about immature moult schedules and plumage sequences in numerous species of African birds. We retain IMMATURE as a descriptive heading, and distinguish juveniles from immatures where we can. In some cases where IMMATURE and NESTLING stages have never been described we simply omit the headings. For plumage and bare parts terminology the reader is referred to Topography of African Passerine Birds in Vol. IV, pp xiv–xv. By flight feathers we mean primaries and secondaries (some authorities include tail feathers too), which we often refer to also as remiges (sing. remex), in distinction from tail feathers or rectrices (sing. rectrix).

Place Names With few exceptions, English names are those recommended by the Nomenclatural Committee of the International Ornithological Congress. French names are from Devillers and Ouellet (1993). Scientific names are from J. L. Peters’s Check-list of Birds of the World, XIII, XIV and XV (Mayr and Greenway 1962, Paynter 1968, 1970), in some cases modified by subsequent taxonomic rearrangement. A few entrenched scientific names have been revised in respect of gender (David and Gosselin 2002a, b). Species sequence is somewhat modified from that of Dowsett and Forbes-Watson (1993). Within a family or genus, we order the species from those thought to be closest to ancestral stock, to those considered to be historically and geographically derivative from them. We tend to see equatorial forest as the ancestral biome in Africa, savanna birds as derived ‘from’ their forest relatives, desert birds from savanna ones, high-latitude birds, including many Palearctic species, from low-latitude ones, and habitatspecialist, diet-specialist and socially complex ones from more generalized kinds. Constructing a sensible linear, evolutionary sequence from the three-dimensional tree of life, however, is bound to remain a subjective matter. Within each polytypic species, the subspecies described first is generally the best-known one, and the sequence thereafter is determined partly by ease of comparison and partly by ranges (broadly, from west to east and north to south).

Young Birds Having replaced ‘Downy Young’ with NESTLING with the start of the passerine volumes, we retain the latter term as a subheading in Description sections, although what is actually meant is hatchling or pre-feathered nestling. In its first plumage of true feathers the bird is a juvenile, and in subsequent plumages it is immature for months or years until moulting into adult plumage at sexual maturity. A bird moulting from immature to adult plumages is sometimes called a subadult.

350 names of national parks, faunal reserves, forests and highlands were listed with regional maps in Vol. V pp. xiv– xix, and complementary information can be had in Fishpool and Evans (2001). For place names, we generally follow The Times Atlas of the World (1990) but depart from it in several cases where prevalence of current local usage seems so to demand. The Democratic Republic of Congo has officially changed its name several times and for brevity, as well as to distinguish it from Congo (Brazzaville), we continue to call it Zaı¨re. Its south-eastern province Shaba has reverted to its old name, Katanga. South African provincial boundaries and names have changed; the former Cape Province is now Northern Cape, Western Cape and Eastern Cape; Transvaal has become North West Province (incorporating a part of the former northern Cape), Limpopo, Mpumalanga and Gauteng; Orange Free State is Free State and Natal is KwaZulu-Natal. The new provinces are shown in Figure 5b on p. xxi. However, the boundaries shown in the Africa map here and throughout the book are the former provincial ones, and the former names are in general use in the text in order to facilitate cross-reference to the five standard South African bird atlases. Kabalega Falls of The Times Atlas is generally called Murchison Falls again; Winam Gulf reverts to Kavirondo Gulf; Lake Mobutu Sese Seko for L. Albert proved to be quite transient; Fernando Po ´o and Rio Muni are nowadays Bioko and Mbini, together comprising Equatorial Guinea; Annobo ´n Island should be so called once more, Pagulu being its administrative area. Other popularly used alternative names or spellings are legion, for instance Impenetrable/Bwindi Forest, Murchison/Kabalega Falls, Muodisho/Mogadishu, Zanzibar/Unguja, Yabelo/Yavelo/ Yabello (Sanctuary and Hills in S Ethiopia). Transliterations between English and Arabic have always been problematic. Errachidia (Morocco) is properly Er Rachidia, and the Arabic for rocky hill is locally spellt jebel, jbel, gebel, djebel, chabal, chappal and gibral (as in Gibraltar). The Arabic for a dry river bed has wide currency as wadi in

xvii

xviii

INTRODUCTION

English and oued in French. Gebel Elba is within the Sudan Government Administrative Area but lies astride the Political Boundary between Egypt and Sudan, and we may not always have attributed it correctly.

Abbreviations Linear measurements are in millimetres (mm) and weights are in grams (g) unless otherwise specified. Under LAYING DATES, A, B, C, D and E refer to East African climatic zones, shown in Figure 6; months not in parentheses are those in which eggs have been found, and they are followed parenthetically by times of other breeding indications. Mainly for the benefit of readers for whom English is not the first language, other abbreviations and symbols are: Y male, X female, ad. adult, ads adults, a.s.l. above sea level, av. average, c. approximately, cm centimetre, E east, h hour, I. Island, imm. immature, IBA Important Bird

Area, Is islands, juv. juvenile, km kilometre, L. Lake, max. maximum, m metre, min. minute, Mt Mount, mts mountains, N north, n sample size, Nat. National (Park), P primary (feather) (e.g. P10 = outermost primary, usually small), Pre´f. Pre´fecture, R. River, Res. Reserve, Rs Rivers, S south, s or sec. second, sp. species (sing.), spp. species (plural), T tail feather, W west.

Reference System A short list of key references is given at the end of most species accounts. They have all been used in compiling accounts, although they are not necessarily cited therein. All references cited in the main text, together with some that may not have been cited, are given in the complementary GENERAL AND REGIONAL REFERENCES, SYSTEMATIC REFERENCES and ACOUSTIC REFERENCES at the end of the book.

References Ash, J. S. and Miskell, J. E. (1998). ‘Birds of Somalia’. Pica Press, Robertsbridge. Barlow, C., Wacher, T. and Disley, T. (1997). ‘A Field Guide to Birds of Gambia and Senegal’. Pica Press, Robertsbridge. Borrow, N. and Demey, R. (2001). ‘Birds of Western Africa’. Christopher Helm, London. Brickell, N. and Konigkramer, T. (1997). ‘African Birds in Cage and Aviary’. African Bird Book Publ., Westville, South Africa. Brown, L. H., Urban, E. K. and Newman, K. (1982). ‘The Birds of Africa’, Vol. I. Academic Press, London. Byaruhanga, A., Kasoma, P. and Pomeroy, D. (2001). ‘Important Bird Areas in Uganda’. Nature/Uganda/EANHS, Kampala. Byers, C., Olsson, U. and Curson, J. (1995). ‘Buntings and Sparrows’. Pica Press, Sussex. Cheke, R. A. and Walsh, J. F. (1996). ‘The Birds of Togo’. B.O.U. Check-list No. 14. British Ornithologists’ Union, Tring. Christy, P. and Clarke, W. V. (1994). ‘Guide des Oiseaux de la Re´serve de la Lope´’. ECOFAC, Libreville, Gabon. Christy, P. and Clarke, W. V. (1998). ‘Guide des Oiseaux de Sa˜o Tome´ et Prı´ncipe’. ECOFAC, Libreville, Gabon. Clement, P., Harris, A. and Davis, J. (1993). ‘Finches and Sparrows’. Christopher Helm/A & C Black, London. Collinson, M. (2002). Comment – a written constitution for species-level taxonomy. British Birds 95, 370–371. David, N. and Gosselin, M. (2002a). Gender agreement of avian species names. Bull. Br. Orn. Club 122, 14–49. David, N. and Gosselin, M. (2002b). The grammatical gender of avian genera. Bull. Br. Orn. Club 122, 257–282. Dean, W. R. J. (2000). ‘The Birds of Angola’. B.O.U. Check-list No. 18. British Ornithologists’ Union, Tring. Devillers, P. and Ouellet, H. (1993). ‘Noms Franc¸ais des Oiseaux du Monde’. MultiMondes, St-Foy, Quebec. Dowsett, R. J. and Forbes-Watson, A. D. (1993). ‘Checklist of Birds of the Afrotropical and Malagasy Regions’. Tauraco Press, Lie`ge.

Fishpool, L. D. C. and Evans, M. I. (2001). ‘Important Bird Areas in Africa and Related Islands’. Pisces Publ. and BirdLife International, Newbury and Cambridge. Fry, C. H., Keith, S. and Urban, E. K. (1988). ‘The Birds of Africa’, Vol. III. Academic Press, London. Fry, C. H., Keith, S. and Urban, E. K. (2000). ‘The Birds of Africa’, Vol. VI. Academic Press, London. Gatter, W. (1997). ‘Birds of Liberia’. Pica Press, Robertsbridge. Goodwin, D. (1982). ‘Estrildid Finches of the World’. British Museum (Natural History), London. Hall, B. P. and Moreau, R. E. (1970). ‘An Atlas of Speciation in African Passerine Birds’. British Museum (Natural History), London. Harrison, J. A., Allan, D. G., Underhill, L. G., Herremans, M., Tree, A. J., Parker, V. and Brown, C. J. (1997). ‘The Atlas of Southern African Birds’, Vol. 2: Passerines. BirdLife South Africa, Johannesburg. Helbig, A. J., Knox, A. G., Parkin, D. T., Sangster, G. and Collinson, M. (2002). Guidelines for assigning species rank. Ibis 144, 518–525. Irwin, M. P. S. (2002). The origins and spread of the Northern and Southern Grey-headed Sparrows in Zambia. Honeyguide 48, 41–46. Isenmann, P. and Moali, A. (2000). ‘Oiseaux d’Alge´rie’. SEOF/ MNHN, Paris. Jenni, L. and Winkler, R. (1994). ‘Moult and Ageing of European Passerines’. Academic Press, London. Keay, R. W. J. (1959). ‘Vegetation Map of Africa South of the Tropic of Cancer. Explanatory Notes’. Oxford Univ. Press, Oxford. Keith, S., Urban, E. K. and Fry, C. H. (1992). ‘The Birds of Africa’. Vol. IV. Academic Press, London. Kingdon, J. (1997). ‘The Kingdon Field Guide to African Mammals’. Academic Press, London. Lamarche, B. (1993–1994). ‘Atlas de Re´partition des Oiseaux du Mali et de Mauritanie’, Fasc. I–VIII. Etudes Sahariennes et Ouest-Africaines, Assoc. Nat. Sahariens et OuestAfricains, Nouakchott/Paris.

INTRODUCTION

Laurent, A. (1990). ‘Catalogue commente´ des oiseaux de Djibouti’. Ministe`re du Commerce, des Transports et du Tourisme, Djibouti. Mabberley, D. J. (1993). ‘The Plant-Book. A Portable Dictionary of the Higher Plants’. Cambridge Univ. Press, Cambridge. Mayr, E. (1942). ‘Systematics and the Origin of Species’. Columbia Univ. Press, New York. Mayr, E. and Greenway, J. C. (Eds) (1962). ‘Check-list of Birds of the World’, Vol. X. Museum of Comparative Zoology, Cambridge, MA. Parker, V. (1999). ‘The Atlas of the Birds of Sul do Save, Southern Mozambique’. Avian Demography Unit and Endangered Wildlife Trust, Johannesburg. Payne, R. B., Hustler, K., Stjernstedt, R., Sefc, K. M. and Sorenson, M. D. (2002). Behavioural and genetic evidence of a recent population switch to a novel host species in brood-parasitic indigobirds Vidua chalybeata. Ibis 144, 373–383. Paynter, R. A. (Ed.) (1968, 1970). ‘Check-list of Birds of the World’, Vol. XIII (1970), XIV (1968). Museum of Comparative Zoology, Cambridge, MA. Restall, R. (1996). ‘Munias and Mannikins’. Pica Press, Sussex. Sangster, G., Knox, A. G., Helbig, A. J. and Parkin, D. T. (2002). Taxonomic recommendations for European birds. Ibis 144, 153–159.

Sibley, C. G. and Monroe, B. L. (1990). ‘Distribution and Taxonomy of Birds of the World’. Yale University Press, New Haven, CT and London. Stevenson, T. and Fanshawe, J. (2002). ‘Field Guide to the Birds of East Africa’. T. & A. D. Poyser, London. Steyn, P. (1996). ‘Nesting Birds’. Fernwood Press, Vlaeberg, South Africa. Summers-Smith, J. D. (1988). ‘The Sparrows’. T. & A. D. Poyser, Calton, UK. Tarboton, W. (2001). ‘A Guide to the Nests and Eggs of Southern African Birds’. Struik Publ., Cape Town. ‘The Times Atlas of the World’ Comprehensive Edition (1990) (ed. J. C. Bartholomew et al.). Times Books, London. Urban, E. K., Fry, C. H. and Keith, S. (1986). ‘The Birds of Africa’. Vol. II. Academic Press, London. Urban, E. K., Fry, C. H. and Keith, S. (1997). ‘The Birds of Africa’. Vol. V. Academic Press, London. Zimmerman, D. A., Turner, D. A. and Pearson, D. J. (1996). ‘Birds of Kenya and Northern Tanzania’. Princeton University Press, Princeton, NJ. White, C. M. N. (1963). ‘A Revised Check List of African Flycatchers, Tits, Tree Creepers, Sunbirds, White-eyes, Honey Eaters, Buntings, Finches, Weavers and Waxbills’. Government Printer, Lusaka.

xix

xxi

INTRODUCTION

10 W

0

10 E

Madeira

20

30

40

50

Tunisia Morocco Algeria

Libya Eygpt

il e

Mali

n

W

oo Ca

o ng Co

Democratic Republic of Congo ¨ (Zaire)

Mbini (Equatorial Guinea)

Burundi Angola

10 N Ethiopia

lia

ma

So Kenya Rwanda Tanzania Malawˆ i

Pemba Zanzibar Mafia

q

am

oz

M

Ma

Botswana

0

20

da

Zimbabwe

10 S

ue

bi

a R. Z

500

0

Comoros Zambia bezi m

Namibia

0

den l f of A

r

´ (Pagulu) Annobon

Gu

Uganda ngo Co River

Togo Bioko Benin ´ Principe Gabon ~ Tome´ Sao

hi

Socotra

Djibouti

Central African Republic

m er

Gh

Sudan

Nigeria

an

a

Burkina Faso

e Nil

Ivory Coast

20

sca

er

Chad

g

e Blu

Ni ver Ri

Guinea Bissau Guinea Sierra Leone LIberia

Eritrea

Niger t e Nile

Mauritania Senegal Gambia

ver N Ri

Western Sahara

30

Suez Canal

ga

Canary Is.

Swaziland

1000 miles

South Africa

30

Lesotho

500 1000 1500 km

Fig. 5(a) Political map.

Ethiopia Ju b a

Riv e r Nile

Sudan

L. Turkana

Rive r

A

L. Albert

L. Edward

B Victoria

L. Kivu

D

E

a

Limpopo

T an ver Ri

Lake

Mpumalanga Gauteng

Somalia

Kenya

Uganda

¨ Zaire

Rwanda C Burundi

North West

Pemba

Tanzania L. Tanganyika

Zanzibar

Free State R iv e

Northern Cape

Eastern Cape

L. Malawˆ i

Zambia

Malaw Êi

KwaZuluNatal

r Rufiji

Western Cape

Mozambique

Fig. 5(b) South African Provinces.

Fig. 6 East African climatic zones (see p. xviii).

Mafia

Passer

Family PASSERIDAE: true sparrows, petronias and snow finches

Small (typically 10–20 cm), chunky birds with large head and seed-eating bill. Short, stout and conical bill tapers to a point; bill of some black only in breeding season, horn-coloured at other times. Nostrils large, covered by short bristles and some forehead feathers; rictal bristles poorly developed. Wing short to fairly long, broad and bluntly pointed; 10 primaries with much reduced P10 (dorsal to P9 and extending only slightly from under outermost upper primary coverts). Legs and feet short and stout, with large scutes covering front of legs. Tail short, square-tipped or slightly notched; 12 feathers. Plumage usually brown, grey or rufous, sometimes with black and in a few species with some yellow or white; upperparts often with dark streaks. Sexes alike in some, markedly different in others. Seasonal variation in some, with feather tips concealing bright colours until abraded. Juvenile like ad. X; nestling naked or covered with dense down. Single annual ad. moult complete, starting after last brood; post-juvenile moult within 1 month of fledging, complete in most, partial in 1. Inhabit open, dry or semi-arid country, including savanna, bush and desert; also woodland, cultivation and human habitation; some species highly urbanized. Most nest in loose colonies, a few solitarily; usually monogamous, some pairing for life. Nest typically a bulky unwoven domed structure with side entrance, placed among tree branches, in cavities, under eaves of houses or in thatched roofs; sometimes a shallow cup placed in a hole; built by both sexes. Eggs speckled. Both sexes incubate and feed young. Flight fairly swift, sometimes undulating; hop or hop and walk or run. Bathe and dust frequently. Form large communal roosts and foraging flocks outside breeding season. Feed largely on seeds on or near ground but also on other plant food, insects (especially for young) and scraps from humans. Some give simple harsh chirps, with chirps sometimes running together to form a song; others produce a variety of songs including 1 that is finch-like. Mainly sedentary with some local or seasonal movements. At one time placed in Fringillidae because of similar skeletal characters (Chapin 1917, Bates 1934), then placed in Ploceidae because of similar appendicular myology, palate structure, nesting habits and feeding behaviour (Mayr and Amadon 1951, Moreau and Greenway 1962, Bentz 1979). Currently considered a separate family because of preglossal muscle, much reduced and dorsally positioned P10, and distinct electrophoretic pattern of egg-white proteins (Sibley 1970, Voous 1977, Bock and Morony 1978, Summers-Smith 1988, Cramp and Perrins 1994). DNA work by Sibley and Ahlquist (1990) suggests that passerids and ploceids should be subfamilies in an enlarged ‘Passeridae’ with wagtails, pipits, accentors and waxbills. Occurs naturally in Eurasia and Africa with centre of distribution in Africa; 2 Passer species introduced elsewhere including the New World, Australia, New Zealand and scattered oceanic islands. 3 genera, 37 species: Passer (true sparrows) 25 species, Petronia (petronias and rock sparrows) 5 species, and Montifringilla (snow finches) 7 species. In Africa, 19 Passer and 4 Petronia species.

Genus Passer Brisson

True sparrows. Sexes alike in P. griseus superspecies and P. montanus, other species sexually dimorphic, Y brightly coloured and often with prominent black throat patch. Unlike Petronia, no yellow spot on throat. Bill of Y black only in breeding season, horn-coloured at other times; sometimes black all year. Nestling naked. Annual post-juvenile moult complete. All species have similar courtship and pair formation displays, including pursuit of single X by several YY; do not perform display flights. Taxonomic treatment has varied considerably (Moreau and Greenway 1962, White 1963, Hall and Moreau 1970, Summers-Smith 1988, Sibley and Monroe 1990, Clement et al. 1993, Dowsett and Forbes-Watson 1993, Cramp and Perrins 1994). Rufous sparrows of the P. iagoensis group have been considered 1, 2, 4, or 6 species. There is a broad measure of agreement that all forms are long-isolated relicts of a formerly widespread single polytypic species, and that they compose a superspecies. P. iagoensis is endemic to Cape Verde Is; the other 5 occur in Africa. All are allopatric, and the 2 that come closest (shelleyi and rufocinctus) are also the most different. Grey-headed sparrows (P. griseus group) have been considered 1, 2, or 5 species. We follow Hall and Moreau (1970) and Summers-Smith (1988) in treating them as a superspecies, despite limited hybridization in zones of overlap. Zones of overlap include: (1) swainsonii and griseus in W Eritrea and W Ethiopia (Assosa and Gambela); (2) swainsonii and gongonensis in SW Ethiopia (Kaffa, Gamo-Gofa, Sidamo provinces) and N central Kenya (Turkana and Marsabit districts); (3) gongonensis and griseus in extreme SE Sudan, E Uganda, W Kenya, NE Tanzania; (4) gongonensis and suahelicus

1

Passer

Family PASSERIDAE: true sparrows, petronias and snow finches

Small (typically 10–20 cm), chunky birds with large head and seed-eating bill. Short, stout and conical bill tapers to a point; bill of some black only in breeding season, horn-coloured at other times. Nostrils large, covered by short bristles and some forehead feathers; rictal bristles poorly developed. Wing short to fairly long, broad and bluntly pointed; 10 primaries with much reduced P10 (dorsal to P9 and extending only slightly from under outermost upper primary coverts). Legs and feet short and stout, with large scutes covering front of legs. Tail short, square-tipped or slightly notched; 12 feathers. Plumage usually brown, grey or rufous, sometimes with black and in a few species with some yellow or white; upperparts often with dark streaks. Sexes alike in some, markedly different in others. Seasonal variation in some, with feather tips concealing bright colours until abraded. Juvenile like ad. X; nestling naked or covered with dense down. Single annual ad. moult complete, starting after last brood; post-juvenile moult within 1 month of fledging, complete in most, partial in 1. Inhabit open, dry or semi-arid country, including savanna, bush and desert; also woodland, cultivation and human habitation; some species highly urbanized. Most nest in loose colonies, a few solitarily; usually monogamous, some pairing for life. Nest typically a bulky unwoven domed structure with side entrance, placed among tree branches, in cavities, under eaves of houses or in thatched roofs; sometimes a shallow cup placed in a hole; built by both sexes. Eggs speckled. Both sexes incubate and feed young. Flight fairly swift, sometimes undulating; hop or hop and walk or run. Bathe and dust frequently. Form large communal roosts and foraging flocks outside breeding season. Feed largely on seeds on or near ground but also on other plant food, insects (especially for young) and scraps from humans. Some give simple harsh chirps, with chirps sometimes running together to form a song; others produce a variety of songs including 1 that is finch-like. Mainly sedentary with some local or seasonal movements. At one time placed in Fringillidae because of similar skeletal characters (Chapin 1917, Bates 1934), then placed in Ploceidae because of similar appendicular myology, palate structure, nesting habits and feeding behaviour (Mayr and Amadon 1951, Moreau and Greenway 1962, Bentz 1979). Currently considered a separate family because of preglossal muscle, much reduced and dorsally positioned P10, and distinct electrophoretic pattern of egg-white proteins (Sibley 1970, Voous 1977, Bock and Morony 1978, Summers-Smith 1988, Cramp and Perrins 1994). DNA work by Sibley and Ahlquist (1990) suggests that passerids and ploceids should be subfamilies in an enlarged ‘Passeridae’ with wagtails, pipits, accentors and waxbills. Occurs naturally in Eurasia and Africa with centre of distribution in Africa; 2 Passer species introduced elsewhere including the New World, Australia, New Zealand and scattered oceanic islands. 3 genera, 37 species: Passer (true sparrows) 25 species, Petronia (petronias and rock sparrows) 5 species, and Montifringilla (snow finches) 7 species. In Africa, 19 Passer and 4 Petronia species.

Genus Passer Brisson

True sparrows. Sexes alike in P. griseus superspecies and P. montanus, other species sexually dimorphic, Y brightly coloured and often with prominent black throat patch. Unlike Petronia, no yellow spot on throat. Bill of Y black only in breeding season, horn-coloured at other times; sometimes black all year. Nestling naked. Annual post-juvenile moult complete. All species have similar courtship and pair formation displays, including pursuit of single X by several YY; do not perform display flights. Taxonomic treatment has varied considerably (Moreau and Greenway 1962, White 1963, Hall and Moreau 1970, Summers-Smith 1988, Sibley and Monroe 1990, Clement et al. 1993, Dowsett and Forbes-Watson 1993, Cramp and Perrins 1994). Rufous sparrows of the P. iagoensis group have been considered 1, 2, 4, or 6 species. There is a broad measure of agreement that all forms are long-isolated relicts of a formerly widespread single polytypic species, and that they compose a superspecies. P. iagoensis is endemic to Cape Verde Is; the other 5 occur in Africa. All are allopatric, and the 2 that come closest (shelleyi and rufocinctus) are also the most different. Grey-headed sparrows (P. griseus group) have been considered 1, 2, or 5 species. We follow Hall and Moreau (1970) and Summers-Smith (1988) in treating them as a superspecies, despite limited hybridization in zones of overlap. Zones of overlap include: (1) swainsonii and griseus in W Eritrea and W Ethiopia (Assosa and Gambela); (2) swainsonii and gongonensis in SW Ethiopia (Kaffa, Gamo-Gofa, Sidamo provinces) and N central Kenya (Turkana and Marsabit districts); (3) gongonensis and griseus in extreme SE Sudan, E Uganda, W Kenya, NE Tanzania; (4) gongonensis and suahelicus

1

2

PASSERIDAE

in NE Tanzania and SW Kenya to about L. Elmenteita; (5) suahelicus and griseus in SW Kenya (E side of L. Victoria to about Nairobi) and N and central Tanzania; and (6) diffusus and griseus in central coastal Angola, SE Tanzania, W, S and E ^ P. griseus breeds alongside Zambia in Luangwa valley, Namibia (Caprivi Strip), NW Zimbabwe, NE Botswana, S Malawi. gongonensis and suahelicus in Kenya (L. Elmenteita) and Tanzania (Mkomazi Game Res. and Serengeti Nat. Park: N. E. Baker, pers. comm.) without hybridizing. Limited hybridization is known in some zones of sympatry as follows: (1) swainsonii and gongonensis in Ethiopia (50 km east of Yavello) and Kenya (Turkana, Marsabit and Mandera); (2) griseus and gongonensis in Kenya (Rift Valley and Kisumu); (3) griseus and suahelicus in W and central Tanzania; and (4) griseus and ^ In addition, the last 2 species hybridize widely and commonly around diffusus in NW, central and E Zambia and S Malawi. L. Kariba and along the Zambezi east to c. 33
300 E in Mozambique (12 hybrid specimens mapped by Hall and Moreau 1970). In these zones of overlap, griseus mainly associates with human habitation and open country, nests in holes and openly in trees, and has small bill and body; swainsonii usually occurs in highlands around human habitation, nests in holes and openly in trees, and has small bill and medium-sized body; gongonensis prefers arid thornbush and grassland, often away from human habitation, nests openly in trees and occasionally in holes, and has the largest bill and body size; suahelicus usually occurs in grassland, bush and around human habitation, nests openly in trees and not in holes, and has fairly large bill but small body; diffusus prefers open acacia and broad-leaved woodland, nests only in holes, and has small bill and body. Evolution and systematics of Passer have been discussed at length by Summers-Smith (1988); our classification and species sequence, from primitive to derived groups, are based on his scheme. 25 species in Africa and Eurasia; 19 in Africa, 13 endemic to mainland, 1 to Socotra, 1 to Africa and Arabia and 4 Palearctic species resident in N Africa (with P. domesticus widely introduced elsewhere). 3 superspecies: P. griseus/ P. swainsonii/P. gongonensis/P. suahelicus/P. diffusus; P. motitensis/P. rufocinctus/P. shelleyi/P. cordofanicus/P. insularis; and P. luteus/P. euchlorus.

Passer griseus superspecies – see Introduction. Passer motitensis superspecies

Passer luteus superspecies

1

4 3

3

2

5

2 1 P. motitensis 2 P. rufocinctus 3 P. shelleyi 4 P. cordofanicus 5 P. insularis

1 P. luteus 2 P. euchlorus

1 1

Plate 1

Passer griseus (Vieillot). Northern Grey-headed Sparrow. Moineau gris.

(Opp. p. 10)

Fringilla grisea Vieillot, 1817. Nouv. Dict. d’Hist. Nat., nouv. ´ed., 12, p. 198, United States [error ¼ Senegal (Lafresnaye, 1839, Rev. Zool., p. 95)]. Forms a superspecies with P. swainsonii, P. gongonensis, P. suahelicus and P. diffusus – see map in Introduction. Range and Status. Endemic resident south of Sahara from Mauritania to Eritrea and south to Angola, N Zimbabwe ^ common to abundant around human habitand Malawi; ation; mainly frequent to uncommon in natural habitats.

Mauritania south of c. 17
N, north along coast to Nouakchott and in east to Kidal. Mali north to c. 19
N, especially abundant between 13
and 16
N. Niger north to Aı¨r (c. 18
N). Chad north to Ouadi Rime´-Ouadi Achim

2

PASSERIDAE

in NE Tanzania and SW Kenya to about L. Elmenteita; (5) suahelicus and griseus in SW Kenya (E side of L. Victoria to about Nairobi) and N and central Tanzania; and (6) diffusus and griseus in central coastal Angola, SE Tanzania, W, S and E ^ P. griseus breeds alongside Zambia in Luangwa valley, Namibia (Caprivi Strip), NW Zimbabwe, NE Botswana, S Malawi. gongonensis and suahelicus in Kenya (L. Elmenteita) and Tanzania (Mkomazi Game Res. and Serengeti Nat. Park: N. E. Baker, pers. comm.) without hybridizing. Limited hybridization is known in some zones of sympatry as follows: (1) swainsonii and gongonensis in Ethiopia (50 km east of Yavello) and Kenya (Turkana, Marsabit and Mandera); (2) griseus and gongonensis in Kenya (Rift Valley and Kisumu); (3) griseus and suahelicus in W and central Tanzania; and (4) griseus and ^ In addition, the last 2 species hybridize widely and commonly around diffusus in NW, central and E Zambia and S Malawi. L. Kariba and along the Zambezi east to c. 33
300 E in Mozambique (12 hybrid specimens mapped by Hall and Moreau 1970). In these zones of overlap, griseus mainly associates with human habitation and open country, nests in holes and openly in trees, and has small bill and body; swainsonii usually occurs in highlands around human habitation, nests in holes and openly in trees, and has small bill and medium-sized body; gongonensis prefers arid thornbush and grassland, often away from human habitation, nests openly in trees and occasionally in holes, and has the largest bill and body size; suahelicus usually occurs in grassland, bush and around human habitation, nests openly in trees and not in holes, and has fairly large bill but small body; diffusus prefers open acacia and broad-leaved woodland, nests only in holes, and has small bill and body. Evolution and systematics of Passer have been discussed at length by Summers-Smith (1988); our classification and species sequence, from primitive to derived groups, are based on his scheme. 25 species in Africa and Eurasia; 19 in Africa, 13 endemic to mainland, 1 to Socotra, 1 to Africa and Arabia and 4 Palearctic species resident in N Africa (with P. domesticus widely introduced elsewhere). 3 superspecies: P. griseus/ P. swainsonii/P. gongonensis/P. suahelicus/P. diffusus; P. motitensis/P. rufocinctus/P. shelleyi/P. cordofanicus/P. insularis; and P. luteus/P. euchlorus.

Passer griseus superspecies – see Introduction. Passer motitensis superspecies

Passer luteus superspecies

1

4 3

3

2

5

2 1 P. motitensis 2 P. rufocinctus 3 P. shelleyi 4 P. cordofanicus 5 P. insularis

1 P. luteus 2 P. euchlorus

1 1

Plate 1

Passer griseus (Vieillot). Northern Grey-headed Sparrow. Moineau gris.

(Opp. p. 10)

Fringilla grisea Vieillot, 1817. Nouv. Dict. d’Hist. Nat., nouv. ´ed., 12, p. 198, United States [error ¼ Senegal (Lafresnaye, 1839, Rev. Zool., p. 95)]. Forms a superspecies with P. swainsonii, P. gongonensis, P. suahelicus and P. diffusus – see map in Introduction. Range and Status. Endemic resident south of Sahara from Mauritania to Eritrea and south to Angola, N Zimbabwe ^ common to abundant around human habitand Malawi; ation; mainly frequent to uncommon in natural habitats.

Mauritania south of c. 17
N, north along coast to Nouakchott and in east to Kidal. Mali north to c. 19
N, especially abundant between 13
and 16
N. Niger north to Aı¨r (c. 18
N). Chad north to Ouadi Rime´-Ouadi Achim

Passer griseus

Passer griseus

?

? ?

? ? ? ? ??? ? ?

?

Faunal Reserve and Fada. Sierra Leone, common Freetown in 1870, not recorded since although common elsewhere except NW (Outamba area), where uncommon. Liberia, has spread since 1960s, now occurs throughout. Ghana, throughout, having become widespread in N savannas in recent years. Sudan north to c. 18
N. Eritrea, in W below 1200 m. Ethiopia, in W at Assosa and Gambela. Uganda except NE. Kenya, most of Rift Valley from Mt Elgon, Kapenguria and Kitale south to Nairobi, L. Magadi and South Nyanza. Rwanda, Burundi. Zaı¨re, probably throughout but status in SW not known. Tanzania, as mapped, but probably far more widespread and may occur in towns and villages throughout (N. E. Baker, pers. comm.). Angola, from Cabinda south through Zaı¨re, Uı´ge, Bengo and Luanda to Cuanze Norte, lower Cuanza R. and NE Lunda Norte; also along coast and escarpment south to Benguela. ^ south to c. Zambia throughout except parts of SW; Malawi 16
S. NW Zimbabwe at Victoria Falls, Hwange Nat. Park, Kariba; also Harare suburbs (P. Leonard, pers. comm.). Namibia, E tip of Caprivi Strip at Impalila I., Katima Mulilio, Bukalo, Ngoma; also NW along Kunene R. at Ruacana (C. J. H. Hines, pers. comm.). N Botswana, at Kazungula border area. NW Mozambique (Tete Province). Density in S Aı¨r (Nigeria) 5 pairs per ha (P. Giraudoux in Summers-Smith 1988). Description. P. g. griseus (Vieillot). W Africa south of laeneni from Mauritania and Senegal to Cameroon, N Gabon, W Central African Republic and Chad. ADULT Y: forehead to nape ashy grey, merging with grey-brown hindneck, sides of neck and upper mantle; lower mantle tawny-brown, grading to rufous-brown on scapulars; back, rump and uppertail-coverts rufous. Tail feathers dark brown, finely edged buff-brown when fresh. Lores and around eye dark grey-brown, merging with paler grey-brown

ear-coverts. Cheeks and sides of throat and breast pale greybrown, enclosing buffish white chin and centre of throat; sometimes a diffuse darker greyish malar stripe. Rest of underparts white, tinged buff, flanks washed grey-brown. Flight feathers dark brown, primaries narrowly edged buff-brown, secondaries edged rufous-brown; tertials dark brown, outer edge and tip broadly fringed rufous-brown. Primary coverts and alula dark brown, tinged rufous; greater coverts rufous-brown; median coverts rufous, inner feathers with broad white tips, forming small, variable white patch; lesser coverts rufous. Underwingcoverts and axillaries buffish white; inner borders of undersides of flight feathers pale grey-buff. Bill black, blackish horn or dark horn; eyes light brown; legs brownish flesh. Sexes alike. SIZE (10 YY, 10 XX, Nigeria and Sudan): wing, Y 82–90 (860), X 82–90 (845); tail, Y 62–73 (657), X 61–68 (639); bill, Y 145–165 (157), X 145–16 (153); tarsus, Y 19–225 (208), X 195–21 (206). Birds in W smaller: (n ¼ 10, Gambia and Sierra Leone) wing, Y, 78–81 (794). WEIGHT: unsexed (n ¼ 53, Nigeria) 242– 330 (281) (Fry 1971). IMMATURE: juvenile similar to adult, but head and neck browner, some dark streaks on back, rump tawny-brown; wing coverts tawny-brown, inner medians with narrower, buff (not white) tips. Bill horn. NESTLING: naked at hatching. P. g. laeneni Niethammer: edge of Sahara from E Mali (in Azzawakh area, c. 15
300 N, 03
180 E), N Burkina Faso, Niger, N Nigeria (south to Kano), N Cameroon and Chad (north of L. Chad) to W Sudan (Darfur Province). Paler than nominate race, top of head light grey, mantle fawn, underparts whiter, bib patch less differentiated. Wing, Y (n ¼ 6, Darfur) 86–89 (872). P. g. ugandae Reichenow: S Gabon, Angola, Congo, E Central African Republic, Zaı¨re, Sudan (except Darfur Province) and ^ Like nominate race but slightly Eritrea to Zimbabwe and Malawi. browner grey on crown, more tawny on mantle, more extensively grey-brown below, with white more restricted to belly; whitish ^ throat patch less distinct. Wing, Y (n ¼ 10, Uganda and Malawi) 82–88 (851). WEIGHT: unsexed (n ¼ 34, W Kenya) 34–43 (392). TAXONOMIC NOTE: we follow Summers-Smith (1988) and Clement et al. (1993) in recognizing ugandae, contra Moreau and Greenway (1962) and White (1963). Boundaries between the 3 races remain unclear.

Field Characters. Length 15 cm. Common and widespread in villages and bush; the only grey-headed sparrow in W Africa. Smallest member of this group, with relatively small bill. Paler and redder than others, with light grey head, reddish chestnut back, rump, tail and wing shoulders; white chin and throat contrast with grey breast; flanks grey, belly paler grey; white bar across chestnut shoulder. Race ugandae has deeper grey head, reddish areas darker and browner. In all members of the group, bill black when breeding, otherwise horn-brown. Immature browner with some dark streaks on back. Meets all 4 other species; for further comparisons, see those accounts. Voice. Tape-recorded (86, 104). Calls, ‘chip’, ‘cherp’, ‘chyo’, ‘tyup’, and more liquid, ‘twee’, ‘tyooy’, rising ‘weet’ and descending ‘peeu’. These are strung together in disjointed series to form song. Alarm scold, dry churring ‘cherdidit’ or ‘cheedadadadat’. General Habits. Inhabits open plains with bushes, savanna, light woodland, coastal scrub and edges of marshes and salt pans from sea level to over 2500 m; absent from dense forest except in clearings. Some fly over primary forest, searching for clearings and occupying them within

3

4

PASSERIDAE

5 or 6 years of their formation (Brosset and Erard 1986). Prefers humid areas, although occurs in arid country S fringes of Sahara. In recent decades, as natural habitat has been cleared, commonest around cultivation, villages, and towns. Occurs in pairs, small groups or flocks of 20–50 or even 100; in arid areas large flocks sometimes damage crops. Noisy, bold and active around settlements, where it fills the role of House Sparrow P. domesticus. Forages on ground, often with other sparrows, firefinches Lagonosticta, indigobirds Vidua, cordon-bleus Uraeginthus, weavers Ploceus and queleas Quelea; also feeds in trees. Scuffles with other sparrows and wagtails around houses for insects attracted at night by lights, which litter the ground at daybreak. Anting reported (Little 1961). Roosts communally in bushes and trees and around houses. At Lovanium, Kinshasa Province, Zaı¨re, moults after nesting in early Mar, young from latest brood do not finish moulting primaries until mid-June (De Bont 1970); in Parc National de l’Upemba, Zaı¨re, moults in dry season (July–Aug) and at beginning of rainy season (Oct–Dec) (Verheyen 1953); in Sudan, breeds Sept–Oct, moults in Mar. Mainly sedentary, but south of Sahara birds move north away from breeding range in dry season, e.g. in Mauritania to Nouakchott, in Sudan to Nuba Prov., in Ghana to Mole Nat. Park. Food. Mainly seeds of grasses and cultivated cereals; also flowers, berries, insects (including caterpillars, flying ants, termites, beetles and locusts); bread, fruit and other household scraps. Breeding Habits. Solitary or loosely colonial nester; in arid N Senegal solitary and territorial, elsewhere in Senegal colonial (G. J. Morel in Summers-Smith 1988). In NE Gabon solitary pair defends territory of several hundred m2 around house containing nest. Early in breeding season Y chases X; when she lands, he lands nearby, postures in front of her and chirps excitedly, stretching neck and head upwards and elevating tail (Summers-Smith 1988). Birds engage in fierce battles, rolling over on ground and striking opponent with bill and claws. Triple brooded, fourth brood possible (Lovanium, Zaı¨re: De Bont 1970). NEST: untidy accumulation of grasses, lined with feathers; domed, with side entrance; placed either openly on branch of tree, or in bunch of plantains or in hole in tree (often old nest of barbet or woodpecker); often uses old nest of bee-eater, kingfisher, swift, or swallow; also on

beam of house, in thatch or under roof, in hollow pipe on electric pylon. Both sexes visit nest during building but only 1 actually carries nest material (Serle 1940). EGGS: 3–4 (occasionally 2). Oval, smooth and slightly glossy; white to yellowish white, heavily blotched and spotted with greys and browns. SIZE (Nigeria, n ¼ 10): 178–214  14–156 (203  150). LAYING DATES: Mauritania, Apr–Oct; Senegal, June–Nov; Gambia, Mar–Dec; Mali, Apr–Nov; Burkina Faso, June, Sept–Nov; Niger, Apr–June; Chad, July–Sept; Sudan, Aug–Nov; Eritrea (juvenile, July); Liberia (courtship Feb–May, nestlings being fed Mar–Apr); Ghana, May– July, (fledglings Jan, Aug); Togo, Sept (entering nest June, feeding young at nest Oct); Nigeria, Jan–Apr, July–Dec; Cameroon, Jan, June–Oct; Gabon, Jan–June, Aug–Dec; Mali, Apr–Nov; Bioko (nest-building Mar, feeding juvenile Sept); Zaı¨re, all year; Uganda, Apr–Aug, Nov; Kenya, Oct–May; Tanzania, Jan (8), Feb (15), Mar (15), Apr (12), May (21), June (14), July (6), Aug (2), Oct (2), Nov (1), Dec ^ (5); Zambia, Jan–May; Malawi, Jan–Apr, July–Sept; Angola (feeding chicks, Feb; breeding condition Apr; young under parental care, July). Lays during rains, but in cultivated areas where presence of water ensures adequate food for young, may nest at any time of year. INCUBATION: by X only, at least in Zaı ¨re. Period: 16 days (Zambia). DEVELOPMENT AND CARE OF YOUNG: both parents feed young equally (Ghana: Lamm 1959), starting at first light and continuing until after dark. Feeding rate: Senegal, c. 20 visits per h in early morning and late afternoon, c. 10 visits per h in middle of day; Zambia, 6–8 visits per h, increasing to 25 visits per h when winged termites available. One parent (presumably X) roosts overnight in nest cavity with young. Young fed mainly insects including locusts, flying termites and Hymenoptera, particularly ants (SummersSmith 1988). Fledging period: 19 days. BREEDING SUCCESS/SURVIVAL: parents usually accompanied by single young, very rarely 2, although once 3 nestlings in 1 nest (Zaı¨re: De Bont 1970). Adult ringed Zaı¨re recaptured more than 7 years later; two birds in captivity lived for more than 11 years. Parasitized by African Cuckoo Cuculus gularis; preyed on by domestic cats, tree snakes and men using slingshots (Zaı¨re: De Bont 1970).

Key References Bannerman D. A. (1948), De Bont, A. F. (1970), Dowsett, R. J. and Dowsett-Lemaire, F. (1980), Lamm, D. W. (1959), Summers-Smith, J. D. (1988).

Plate 1

Passer swainsonii (Ru ¨ ppell). Swainson’s Sparrow. Moineau de Swainson.

(Opp. p. 10)

Pyrgita swainsonii Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 94, pl. 33; Abyssinia. Forms a superspecies with P. griseus, P. gongonensis, P. suahelicus and P. diffusus. Range and Status. Endemic resident in NE Africa, common except in Sudan. Sudan, uncommon Port Sudan, extreme SE,

and 1 record on Eritrean border at about 15
300 N, 36
300 E0 , another on Ethiopian border at about 8
300 N, 33
300 E.

4

PASSERIDAE

5 or 6 years of their formation (Brosset and Erard 1986). Prefers humid areas, although occurs in arid country S fringes of Sahara. In recent decades, as natural habitat has been cleared, commonest around cultivation, villages, and towns. Occurs in pairs, small groups or flocks of 20–50 or even 100; in arid areas large flocks sometimes damage crops. Noisy, bold and active around settlements, where it fills the role of House Sparrow P. domesticus. Forages on ground, often with other sparrows, firefinches Lagonosticta, indigobirds Vidua, cordon-bleus Uraeginthus, weavers Ploceus and queleas Quelea; also feeds in trees. Scuffles with other sparrows and wagtails around houses for insects attracted at night by lights, which litter the ground at daybreak. Anting reported (Little 1961). Roosts communally in bushes and trees and around houses. At Lovanium, Kinshasa Province, Zaı¨re, moults after nesting in early Mar, young from latest brood do not finish moulting primaries until mid-June (De Bont 1970); in Parc National de l’Upemba, Zaı¨re, moults in dry season (July–Aug) and at beginning of rainy season (Oct–Dec) (Verheyen 1953); in Sudan, breeds Sept–Oct, moults in Mar. Mainly sedentary, but south of Sahara birds move north away from breeding range in dry season, e.g. in Mauritania to Nouakchott, in Sudan to Nuba Prov., in Ghana to Mole Nat. Park. Food. Mainly seeds of grasses and cultivated cereals; also flowers, berries, insects (including caterpillars, flying ants, termites, beetles and locusts); bread, fruit and other household scraps. Breeding Habits. Solitary or loosely colonial nester; in arid N Senegal solitary and territorial, elsewhere in Senegal colonial (G. J. Morel in Summers-Smith 1988). In NE Gabon solitary pair defends territory of several hundred m2 around house containing nest. Early in breeding season Y chases X; when she lands, he lands nearby, postures in front of her and chirps excitedly, stretching neck and head upwards and elevating tail (Summers-Smith 1988). Birds engage in fierce battles, rolling over on ground and striking opponent with bill and claws. Triple brooded, fourth brood possible (Lovanium, Zaı¨re: De Bont 1970). NEST: untidy accumulation of grasses, lined with feathers; domed, with side entrance; placed either openly on branch of tree, or in bunch of plantains or in hole in tree (often old nest of barbet or woodpecker); often uses old nest of bee-eater, kingfisher, swift, or swallow; also on

beam of house, in thatch or under roof, in hollow pipe on electric pylon. Both sexes visit nest during building but only 1 actually carries nest material (Serle 1940). EGGS: 3–4 (occasionally 2). Oval, smooth and slightly glossy; white to yellowish white, heavily blotched and spotted with greys and browns. SIZE (Nigeria, n ¼ 10): 178–214  14–156 (203  150). LAYING DATES: Mauritania, Apr–Oct; Senegal, June–Nov; Gambia, Mar–Dec; Mali, Apr–Nov; Burkina Faso, June, Sept–Nov; Niger, Apr–June; Chad, July–Sept; Sudan, Aug–Nov; Eritrea (juvenile, July); Liberia (courtship Feb–May, nestlings being fed Mar–Apr); Ghana, May– July, (fledglings Jan, Aug); Togo, Sept (entering nest June, feeding young at nest Oct); Nigeria, Jan–Apr, July–Dec; Cameroon, Jan, June–Oct; Gabon, Jan–June, Aug–Dec; Mali, Apr–Nov; Bioko (nest-building Mar, feeding juvenile Sept); Zaı¨re, all year; Uganda, Apr–Aug, Nov; Kenya, Oct–May; Tanzania, Jan (8), Feb (15), Mar (15), Apr (12), May (21), June (14), July (6), Aug (2), Oct (2), Nov (1), Dec ^ (5); Zambia, Jan–May; Malawi, Jan–Apr, July–Sept; Angola (feeding chicks, Feb; breeding condition Apr; young under parental care, July). Lays during rains, but in cultivated areas where presence of water ensures adequate food for young, may nest at any time of year. INCUBATION: by X only, at least in Zaı ¨re. Period: 16 days (Zambia). DEVELOPMENT AND CARE OF YOUNG: both parents feed young equally (Ghana: Lamm 1959), starting at first light and continuing until after dark. Feeding rate: Senegal, c. 20 visits per h in early morning and late afternoon, c. 10 visits per h in middle of day; Zambia, 6–8 visits per h, increasing to 25 visits per h when winged termites available. One parent (presumably X) roosts overnight in nest cavity with young. Young fed mainly insects including locusts, flying termites and Hymenoptera, particularly ants (SummersSmith 1988). Fledging period: 19 days. BREEDING SUCCESS/SURVIVAL: parents usually accompanied by single young, very rarely 2, although once 3 nestlings in 1 nest (Zaı¨re: De Bont 1970). Adult ringed Zaı¨re recaptured more than 7 years later; two birds in captivity lived for more than 11 years. Parasitized by African Cuckoo Cuculus gularis; preyed on by domestic cats, tree snakes and men using slingshots (Zaı¨re: De Bont 1970).

Key References Bannerman D. A. (1948), De Bont, A. F. (1970), Dowsett, R. J. and Dowsett-Lemaire, F. (1980), Lamm, D. W. (1959), Summers-Smith, J. D. (1988).

Plate 1

Passer swainsonii (Ru ¨ ppell). Swainson’s Sparrow. Moineau de Swainson.

(Opp. p. 10)

Pyrgita swainsonii Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 94, pl. 33; Abyssinia. Forms a superspecies with P. griseus, P. gongonensis, P. suahelicus and P. diffusus. Range and Status. Endemic resident in NE Africa, common except in Sudan. Sudan, uncommon Port Sudan, extreme SE,

and 1 record on Eritrean border at about 15
300 N, 36
300 E0 , another on Ethiopian border at about 8
300 N, 33
300 E.

Passer swainsonii

Passer swainsonii

Field Characters. Length 16 cm. The grey-headed sparrow of NE Africa. Darker than Northern Grey-headed Sparrow P. griseus and Parrot-billed Sparrow P. gongonensis, especially on head; underparts solid grey (no white throat) except for touch of white on belly; dull brown mantle and back contrast with rufous-chestnut rump; smaller than Parrot-billed Sparrow, with slimmer bill, no brownish wash on head or underparts.

Voice. Not tape-recorded. Chirping calls similar to Northern Grey-headed Sparrow but slightly more musical or liquid, ‘chirrip’ or ‘chirri-up’ (Clement et al. 1993).

General Habits. Inhabits alpine areas, moorland, light woodland, savanna and grassland with bushes, but commonest in and around cultivation, villages and towns. In Eritrea chiefly in open highlands above 1200 m, but in NE descends to coast, where confined to villages and towns. In Ethiopia occurs at c. 1200–4500 m, in all habitats except dense forest. Outside breeding season occurs in flocks, sometimes numbering several hundred; does some damage to crops and gardens. Behaviour like Northern Grey-headed Sparrow P. griseus around settlements. Searches for insects under eaves of houses. Sedentary, with some seasonal wandering. Present at Harar (Ethiopia) Mar–May and Dec, absent Oct. Eritrea, highlands and E escarpment to coast, where largely confined to towns and villages. Ethiopia, throughout except SE (Ogaden), and not in dense forest. Djibouti, throughout. NW and N Somalia, north of 9
N and west of 49
E. N-central Kenya from Moyale south to Marsabit; also NW shore of L. Turkana (Lokitaung). Description. ADULT Y: forehead to nape brownish grey, merging with grey-brown hindneck, sides of neck and mantle, and grading to dull rufous-brown on scapulars; back, rump and short uppertail-coverts deep rufous-chestnut; long uppertail-coverts dark brown, fringed rufous. Tail feathers dark brown, finely edged buff-brown. Lores and around eye dark grey-brown, merging with paler grey-brown ear-coverts. Cheeks, throat, breast, flanks and thighs light grey-brown, enclosing paler area on centre of chin and upper throat, and grading to pale grey-buff and some white on centre of belly; undertail-coverts light greybrown, fringed buffy white. Remiges dark brown, primaries narrowly edged buff-brown, secondaries edged rufous-brown, tertials broadly edged and tipped deep rufous-brown. Primary coverts and alula dark brown, tinged rufous; greater coverts rufous-brown; median coverts rufous, inner feathers with broad white tips; lesser coverts deep rufous-chestnut. Underwingcoverts and axillaries grey-buff. Bill black, blackish horn or dark horn, eyes dark brown; legs pale brown. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 81–90 (860), X 79–88 (846); tail, Y 63–68 (662), X 61–67 (645); bill, Y 15–16 (154), X 15–16 (154); tarsus, Y 19–21 (204), X 195–205 (201). WEIGHT: Ethiopia (Addis Ababa), unsexed (n ¼ 20) 273–352 (316). IMMATURE: juvenile like ad. but browner, without white wingbar. Bill sepia above, horn brown below. NESTLING: naked at hatching.

Food. Seeds of grasses and cereals; also insects.

Breeding Habits. 1 bird (sex?) postured in front of another, holding its wings out and slightly drooped to expose chestnut lower back and rump; upper back feathers were ruffled, giving it a hunched appearance (Summers-Smith 1988). NEST: untidy bundle of grass, lined with feathers; domed, with side entrance; placed on branch of tree, in crown of palm, or in hole in building or tree; also relines and uses nests of other birds including Ethiopian Swallow Hirundo aethiopica and Plain Martin (Brown-throated Sand Martin) Riparia paludicola. Twice (L. Abiata, Ethiopia) usurped Plain Martin nest-hole in sand bank, ejecting nestlings (Pierce and Rayment 1998). EGGS: 3–6; Somalia (n ¼ 12) 4–5 (3). White, marked with brown and grey. SIZE (Eritrea, Ethiopia and Somalia, n ¼ 60): 185–220  145–160 (202  156). LAYING DATES: Eritrea, Jan–Mar, May–Nov; Ethiopia, Apr–Dec; Djibouti, Mar; Somalia, Mar (3), June (9), Nov (1).

Key Reference Summers-Smith, J. D. (1988).

5

6

PASSERIDAE

Plate 1

Passer gongonensis (Oustalet). Parrot-billed Sparrow. Moineau perroquet.

(Opp. p. 10)

Pseudostruthus gongonensis Oustalet, 1890. Naturaliste, p. 274; Gongoni, near Mombasa. Forms a superspecies with P. griseus, P. swainsonii, P. suahelicus and P. diffusus. Range and Status. Endemic resident; common except in Sudan. Sudan in extreme SE; E Uganda west to c. 33
300 E; Ethiopia south of c. 7
N; Somalia south of 6
N; Kenya, as mapped but probably throughout except SW Nyanza Prov.; Tanzania, in dry NE lowlands from Same District and Mkomazi Game Res. west across Pangani R. valley to N and NE Masai steppe. Description. ADULT Y: forehead to nape brownish grey, merging with grey-brown hindneck, sides of neck and mantle, and grading to rufous-brown on scapulars; feathers from crown to mantle tipped greyish when fresh; back, rump and uppertail-coverts rufous. Tail feathers dark brown, finely edged buff-brown when fresh. Lores and around eye dark grey-brown, merging with paler grey-brown ear-coverts. Cheeks, throat, breast, flanks and thighs light grey-brown, with paler grey-buff area on centre of chin and upper throat, grading to pale grey-buff on centre of belly; undertail-coverts light grey-brown, fringed buffy white. Remiges dark brown, primaries narrowly edged buff-brown, secondaries edged rufous-brown, tertials broadly edged and tipped rufousbrown. Primary coverts and alula dark brown, tinged rufous; greater coverts rufous-brown; median coverts rufous, inner feathers with broad white tips, forming conspicuous white patch; lesser coverts rufous. Underwing-coverts and axillaries grey-buff; inner borders of undersides of flight feathers pale grey-buff. Bill blackish horn; eyes brown; legs dusky flesh. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 93–100 (972), X89–96 (930); tail, Y 66–71 (679), X 60–66 (614); bill, Y 16–185 (175), X 165–17 (168); tarsus, Y 205–23 (217), X 205–22 (213). WEIGHT: central and SE Kenya, unsexed (n ¼ 23) 335–48 (399). IMMATURE: juvenile resembles adult, but head and mantle browner, rump and upperwing-coverts more tawny brown; white patch vestigial or absent; bill sepia above, horn brown below. NESTLING: naked at hatching.

Field Characters. Length 18 cm. Larger than other greyheaded sparrows, with thicker, rounded, ‘parrot’ bill; head and underparts medium grey with slight brownish cast, no white on throat but a touch on undertail-coverts; mantle and upper back dull brown contrasting with rufous lower back and rump (some rufous extending to lower flanks); conspicuous white patch on reddish wing-shoulder. Immature has brown wing-shoulder, white patch vestigial or absent, rump paler, white at sides. Voice. Tape-recorded (B). Calls louder, deeper and more full than other grey-headed sparrows, produced in series as song, ‘choop, chyop, choop, chup, chop, choop, tsut, tyop . . . ’. Also said to give churring ‘chrrrrrrryek’ or ‘wurrrrrrk’, and a longer ‘tserkisee-chirr’ (Zimmerman et al. 1996). General Habits. Occurs in open, often arid thorn bush, savanna, and grassland with bushes and scrub; also around human settlements, but less than other grey-headed sparrows, although behaves like House Sparrow P. domesticus in Mogadishu, Somalia (J. S. Ash in Summers-Smith 1988). Occurs in all habitats in Tsavo Nat. Park, Kenya,

Passer gongonensis

commonest in riverine vegetation, rare in grassland and woodland (Lack 1985). Sea level to c. 1800–2000 m. Occurs in pairs or small groups of 4–6, sometimes flocks of up to 20–30, rarely more. Shyer and less approachable than other grey-headed sparrows; feeds around outskirts of villages but seldom in village streets. Feeds on ground, mainly in early morning and late afternoon; has recently taken to feeding on scraps at lodges and picnic sites in game reserves. Food. Seeds of grasses and doubtless insects; human scraps. Breeding Habits. In courtship display, 1 bird chases another in headlong flight; pursuer (presumably Y) lands and hops in front of the other, holding wings out and slightly drooped (Summers-Smith 1988). NEST: untidy domed structure of grass with side entrance, lined with feathers; placed in open in trees and bushes and occasionally in holes in baobob trees (N. E. Baker, pers. comm.); uses old nests of other birds. EGGS: 2 (Summers-Smith 1988); Somalia (n ¼ 12) 4–5. Greyish white, with numerous grey and rufous-brown markings. SIZE (Tsavo, Kenya, n ¼ 2): c. 22  15. LAYING DATES: Ethiopia, Dec–Mar, May; Somalia, June, Sept, Nov, (building, Apr); Kenya, Jan, Mar–July, Oct– Dec. Breeds usually during or following rains, also in dry season near irrigated fields. Key Reference Summers-Smith, J. D. (1988).

Passer diffusus

7

Passer suahelicus (Reichenow). Swahili Sparrow. Moineau swahili.

Plate 1

Passer griseus suahelicus Reichenow, 1904. Vo ¨g. Africa, 3, p. 231; Bussissi [Mwanza district, Tanganyika Territory].

(Opp. p. 10)

Forms a superspecies with P. griseus, P. swainsonii, P. gongonensis and P. diffusus. Range and Status. Endemic resident, locally common. SW Kenya from Karungu Bay, L. Elmenteita and Simba south through Narok, Mara Game Res. and Loita Plains to Taveta and Tanzania border; Tanzania from south end of L. Victoria, Serengeti and Crater Highlands south to Rukwa and Usangu Flats.

Passer suahelicus

Description. ADULT Y: forehead to hindneck, sides of neck and upper mantle greyish brown, merging with drab brown lower mantle and scapulars; back, rump and short uppertail-coverts rufous-cinnamon; long uppertail-coverts dark grey-brown, edges tinged cinnamon. Tail feathers dark brown, finely edged buffish. Lores and around eye dark grey-brown, merging with paler greybrown ear-coverts. Cheeks and underparts grey-buff, slightly paler on throat, belly and undertail-coverts. Remiges dark brown, primaries narrowly edged buff-brown, secondaries edged tawnybrown, tertials broadly edged and tipped tawny-brown. Primary coverts and alula dark brown; greater coverts tawny-brown; median coverts rufous-cinnamon, inner feathers with broad white tips; lesser coverts rufous-cinnamon. Underwing-coverts and axillaries pale grey-buff. Bill dark or blackish horn; eyes light brown to cinnamon-grey; legs brownish flesh. Sexes alike. SIZE (5 YY, 5 XX): wing, Y 84–91 (874), X 85–89 (862); tail, Y 64–66 (652), X 61–67 (642); bill, Y 15–155 (151), X 15–155 (152); tarsus, Y 19–205 (200), X 195–205 (198). IMMATURE: juvenile resembles adult. NESTLING: naked at hatching.

Field Characters. Length 16 cm. The grey-headed sparrow of SW Kenya and central Tanzania. Smaller and thinnerbilled than Parrot-billed Sparrow P. gongonensis, which it meets in N of range. In SW Kenya and Tanzania overlaps Northern Grey-headed Sparrow P. griseus ugandae; same size but back grey-brown concolorous with head and neck, contrasting sharply with red-brown rump (back of Northern Grey-headed darker and redder, not contrasting with rump); underparts greyer, throat and belly pale grey but not white. Voice. Tape-recorded (CHA). Varied 1- and 2-syllable calls given in conversational medley (song?): ‘chup, chu-weew, chop, wor, chu-weet, chip, choy, chaw-chip, chaw, worchip, chyoop . . . ’. General Habits. Inhabits mainly grassland with scattered scrub and bushes, also light woodland, edges of cultivation and villages, at 1000–2000 m; associates with human

habitation more than does Parrot-billed Sparrow P. gongonensis. Sedentary, with local movements during rains. Food. A variety of seeds; crumbs and scraps at game lodges. Breeding Habits. NEST: untidy domed structure with tunnel side entrance; built in holes in trees and houses; regularly uses old swallow nests. Once pair dragged 2 almost-fledged Lesser Striped Swallows Hirundo abyssinica from nest to ground and stabbed them to death with their bills; they then enlarged side entrance and relined nest, but eventually abandoned it (Kenya: Wilson and Wilson 1994). EGGS: 1, 220  160. LAYING DATES: Kenya, Jan, Apr–May; Tanzania, Jan– Mar, May.

Key Reference Summers-Smith, J. D. (1988).

Passer diffusus (Smith). Southern Grey-headed Sparrow. Moineau sud-africain.

Plate 1

Pyrgita diffusus Smith, 1836. Report Exped. Centr. Africa, p. 50; between the Orange River and the tropic; restricted to ‘near Kuruman’, by Macdonald and Hall (1957, Ann. Transvaal Mus., 23, p.35).

(Opp. p. 10)

Forms a superspecies with P. griseus, P. swainsonii, P. gongonensis and P. suahelicus. Range and Status. Endemic resident in southern Africa, common and widespread in most of range. Angola, in

Namibe, Huı´la, Cunene and Cuando Cubango, also along coastal areas from Benguela north to Bengo (Ambriz and

Passer diffusus

7

Passer suahelicus (Reichenow). Swahili Sparrow. Moineau swahili.

Plate 1

Passer griseus suahelicus Reichenow, 1904. Vo ¨g. Africa, 3, p. 231; Bussissi [Mwanza district, Tanganyika Territory].

(Opp. p. 10)

Forms a superspecies with P. griseus, P. swainsonii, P. gongonensis and P. diffusus. Range and Status. Endemic resident, locally common. SW Kenya from Karungu Bay, L. Elmenteita and Simba south through Narok, Mara Game Res. and Loita Plains to Taveta and Tanzania border; Tanzania from south end of L. Victoria, Serengeti and Crater Highlands south to Rukwa and Usangu Flats.

Passer suahelicus

Description. ADULT Y: forehead to hindneck, sides of neck and upper mantle greyish brown, merging with drab brown lower mantle and scapulars; back, rump and short uppertail-coverts rufous-cinnamon; long uppertail-coverts dark grey-brown, edges tinged cinnamon. Tail feathers dark brown, finely edged buffish. Lores and around eye dark grey-brown, merging with paler greybrown ear-coverts. Cheeks and underparts grey-buff, slightly paler on throat, belly and undertail-coverts. Remiges dark brown, primaries narrowly edged buff-brown, secondaries edged tawnybrown, tertials broadly edged and tipped tawny-brown. Primary coverts and alula dark brown; greater coverts tawny-brown; median coverts rufous-cinnamon, inner feathers with broad white tips; lesser coverts rufous-cinnamon. Underwing-coverts and axillaries pale grey-buff. Bill dark or blackish horn; eyes light brown to cinnamon-grey; legs brownish flesh. Sexes alike. SIZE (5 YY, 5 XX): wing, Y 84–91 (874), X 85–89 (862); tail, Y 64–66 (652), X 61–67 (642); bill, Y 15–155 (151), X 15–155 (152); tarsus, Y 19–205 (200), X 195–205 (198). IMMATURE: juvenile resembles adult. NESTLING: naked at hatching.

Field Characters. Length 16 cm. The grey-headed sparrow of SW Kenya and central Tanzania. Smaller and thinnerbilled than Parrot-billed Sparrow P. gongonensis, which it meets in N of range. In SW Kenya and Tanzania overlaps Northern Grey-headed Sparrow P. griseus ugandae; same size but back grey-brown concolorous with head and neck, contrasting sharply with red-brown rump (back of Northern Grey-headed darker and redder, not contrasting with rump); underparts greyer, throat and belly pale grey but not white. Voice. Tape-recorded (CHA). Varied 1- and 2-syllable calls given in conversational medley (song?): ‘chup, chu-weew, chop, wor, chu-weet, chip, choy, chaw-chip, chaw, worchip, chyoop . . . ’. General Habits. Inhabits mainly grassland with scattered scrub and bushes, also light woodland, edges of cultivation and villages, at 1000–2000 m; associates with human

habitation more than does Parrot-billed Sparrow P. gongonensis. Sedentary, with local movements during rains. Food. A variety of seeds; crumbs and scraps at game lodges. Breeding Habits. NEST: untidy domed structure with tunnel side entrance; built in holes in trees and houses; regularly uses old swallow nests. Once pair dragged 2 almost-fledged Lesser Striped Swallows Hirundo abyssinica from nest to ground and stabbed them to death with their bills; they then enlarged side entrance and relined nest, but eventually abandoned it (Kenya: Wilson and Wilson 1994). EGGS: 1, 220  160. LAYING DATES: Kenya, Jan, Apr–May; Tanzania, Jan– Mar, May.

Key Reference Summers-Smith, J. D. (1988).

Passer diffusus (Smith). Southern Grey-headed Sparrow. Moineau sud-africain.

Plate 1

Pyrgita diffusus Smith, 1836. Report Exped. Centr. Africa, p. 50; between the Orange River and the tropic; restricted to ‘near Kuruman’, by Macdonald and Hall (1957, Ann. Transvaal Mus., 23, p.35).

(Opp. p. 10)

Forms a superspecies with P. griseus, P. swainsonii, P. gongonensis and P. suahelicus. Range and Status. Endemic resident in southern Africa, common and widespread in most of range. Angola, in

Namibe, Huı´la, Cunene and Cuando Cubango, also along coastal areas from Benguela north to Bengo (Ambriz and

8

PASSERIDAE

lower Cuanza valley). Zambia, south of c. 14
S; also ^ south of S Luangwa valley north to near Isoka. Malawi, ^ Coastal Tanzania southward from 7
250 S end of L. Malawi. (Kisiju); also Pemba, Zanzibar and Mafia Is. Mozambique, throughout except N Niassa and NW Tete Provinces. Zimbabwe, throughout, spreading since 1950s and 1960s over all Mashonaland Plateau and parts of E (e.g. Vumba Highlands) where previously absent (Irwin 2001). Namibia, absent only from coast and Namib Desert. Lowlands of Lesotho, spreading into mountains along rivers. Swaziland, resident in lowveld, S middleveld and S highveld; uncommon visitor elsewhere. South Africa, throughout in KwaZulu-Natal, Transvaal, Free State and E Cape; in 1940s and 1950s spread from KwaZulu-Natal into Transkei, then westward across Cape south of c. 32
S, reaching SW Cape at c. 19
E (Elgin) and in S to c. 34
S (Bradasdorp). Density of breeding pairs in Acacia karroodominated habitats in Free State: Modder R., 06 pairs per 10 km, Rhenoster Spruit 08 pairs per 10 km (Kopij 2000). Population in Sul do Save, S Mozambique, >50,000 birds (Parker 1999). Density, Acacia savanna, Swaziland, 51 birds per 100 ha (Monadjem 2002). Description. P. d. diffusus (Smith) (including ‘georgicus’): Angola, Namibia, SW Zambia, Botswana, arid W Zimbabwe, and W South Africa (W Transvaal, W Free State and N Cape). ADULT Y: forehead to hindneck and upper sides of neck rather pale ashgrey; mantle and scapulars brown, fresh feather tips greyish, outer scapulars tinged rufous; back to short uppertail-coverts rufous-cinnamon; long uppertail-coverts brown. Tail feathers dark brown, finely edged buff-brown when fresh. Lores to earcoverts grey-brown. Cheeks, sides of throat, lower sides of neck, breast and flanks pale brownish grey; whitish patch on chin and centre of throat; belly and undertail-coverts whitish. Flight feathers dark brown, primaries narrowly edged buff, secondaries edged cinnamon-buff; tertials dark brown, outer edge and tip broadly fringed cinnamon-buff. Primary coverts and alula dark brown; greater coverts cinnamon-buff; median coverts rufouscinnamon, inner feathers broadly tipped white; lesser coverts rufous-chestnut. Underwing-coverts and axillaries buffish white; inner borders of undersides of flight feathers pale grey-buff. Bill horn, lower mandible pinkish, becoming all-black in breeding season; eyes dark brown or greyish brown; legs dark horn to flesh-brown. Sexes alike. SIZE (8 YY, 10 XX): wing, Y 78–88 (829), X 73–83 (785); tail, Y 57–68 (625), X 55–62 (592); bill, Y 13–155 (139), X 13–15 (137), depth at base c. 7; tarsus, Y 185– 20 (191), X 175–195 (185). WEIGHT: unsexed (n ¼ 130, W Transvaal) 20–296 (242). IMMATURE: juvenile like adult, but head rather browner, upperparts faintly mottled, small wing-coverts less rufous. NESTLING: naked at hatching. ^ S Mozambique, P. d. stygiceps Clancey: SE Zambia, SW Malawi, Zimbabwe (except W Matabeleland), E South Africa (Transvaal except arid W, E Free State, Natal, E Cape), Lesotho and Swaziland. Slightly darker, browner on head and upperparts; suffused more grey-brown (less whitish) below. P. d. luangwae Benson: Luangwa valley, Zambia. Like mosambicus, but smaller, with relatively small bill. SIZE (4 YY, 2 XX): wing, Y 75–80 (777), X 79, 82; bill, Y 11–12 (115), X 115, 125, depth at base c. 6 mm. ^ N Mozambique and P. d. mosambicus van Someren: SE Malawi, SE Tanzania. Like stygiceps but crown greyer, contrasting with slightly warmer brown back. Wing length as nominate race: (10 YY) 80–86 (830); bill slightly deeper, c. 8 mm at base. TAXONOMIC NOTE: we follow Clancey (1980b) in recognizing P. d. stygiceps, but it may be of hybrid origin, possibly diffusus 

Passer diffusus

? ?

ugandae (R. J. Dowsett, pers. comm.) or it may represent a W-to-E cline of P. d. diffusus from lighter to darker (¼ stygiceps) plumage (Summers-Smith 1988). We also follow Hall and Moreau (1970), Summers-Smith (1988), Short et al. (1990) and Zimmerman et al. (1996) in treating mosambicus as a race of P. diffusus, although Benson (1956) and R. J. Dowsett (pers. comm.) believe it may be a race of P. griseus.

Field Characters. Length 15–16 cm. The grey-headed sparrow of southern Africa, meeting Northern Grey^ headed Sparrow P. griseus in W Angola, Zambia, Malawi and coastal Tanzania. Slightly slimmer, shorter-tailed and thinner-billed, grey head paler, back paler and browner, white wing-bar more pronounced; more uniform pale grey below, whitish throat contrasting less with breast. In Zambia at any rate, bill black only during breeding season (early Sept–late Mar), otherwise pale yellow, whereas black all year in Northern Grey-headed (Dowsett and DowsettLemaire 1980). In E coastal race mosambicus head darker and back richer brown, closer to Northern Grey-headed. Ecologically separated in Zambia, where Southern Greyheaded inhabits mopane and other woodland while Northern lives around buildings.

Voice. Tape-recorded (7, 58, 72, 88, 99, C, F). Chirping calls somewhat more liquid and less scratchy than House Sparrow P. domesticus; when strung together form loose song, ‘chyup, chet, chyew, chyurp, chip, chyit, chooy . . . ’; pace irregular, with some slightly faster spurts, gaps between notes varying in length, overall rate 1 note in c. 08 s. Tinny quality cited by Maclean (1993) not apparent on either of the tapes cited above. Also gives a dry churr ending with several distinct notes, ‘churrrrrdididit’, like Great Tit Parus major.

Passer motitensis General Habits. Inhabits primarily open acacia and broadleaved woodland including mopane and miombo; occurrence in wide range of other habitats (open plains, grassland, savanna) due to association with man; occupies farmyards, cattle camps, plantations of exotics, gardens and villages. Common in thorn tree savanna (but replaced in broad-leaved savanna by Yellow-throated Sparrow Petronia superciliaris: Ginn et al. 1989). Rare in fynbos and succulent Karoo habitats; avoids forest and dense woodland. Where sympatric with House Sparrow in urban environments, favours suburban habitats; around farm houses often displaced by House Sparrow, returning if House Sparrow removed (Ginn et al. 1989). Where sympatric with Northern Grey-headed Sparrow, prefers mopane and other dry woodland away from man (Northern more in cultivation and around settlements). Rarely competes with Cape Sparrow P. melanurus which is much more adapted to man; when in same area usually loses nest-disputes with it (Summers-Smith 1988). Occurs in pairs, outside breeding season also in small flocks (probably family groups), sometimes flocks of 50–60 or even 500. Forages for seeds on bare ground, including cement paving, road verges and inside open sheds and barns (Ginn et al. 1989), often with Cape, House and Great Sparrows and other seedeaters. Walks as well as hops (very unusual in Passer: Summers-Smith 1988) which distinguishes it in mixed flocks from House Sparrows (A. J. F. K. Craig in Summers-Smith 1988). When not feeding, usually in trees not far from water. Foliage-bathes after rain (Tyler 1999). Moult follows breeding season; post-juvenile moult complete. Mainly sedentary, with some local movements. 5 recoveries of ringed birds in Zimbabwe less than 10 km from where ringed. Less frequent in winter in NW and extreme S areas (Zones 1–3, 8) (Harrison et al. 1997); birds common June but absent Nov (Botswana: Brewster 1991). Population at Mwaku Pan, NW Botswana, nomadic (Tree 1972). Food. Seeds and insects, including termite alates (latter also fed to young); also buds, berries, aloe nectar and food scraps. Breeding Habits. Solitary nester, occasionally in loose colonies; monogamous. Early in breeding season Y and X fly together to investigate nest-sites, and engage in display chases. At nest-site, calls repeatedly; when excited ruffles lower back and rump feathers, and flicks tail from side to

9

side in rotary motion. Threatens competing conspecifics and congeners (e.g. House Sparrow) by holding wings away from body and raising and spreading tail. Fights occur, bird grasping opponent with bill or claws and both falling to ground; when pairs fight, birds attack opponent of same sex (Summers-Smith 1988). Double-brooded. NEST: untidy bunch of grass, sometimes small twigs, lined with feathers, wool and hair; placed in hole in tree, including old nest hole of woodpecker or barbet, also under eaves or in thatched roof of building, hole in wall, fencepost or telegraph pole, hollow pipe, nest box and in abandoned farm or mine machinery; also uses old nests of Greater Striped Swallow Hirundo cucullata, Little Swift Apus affinis, Red-billed Buffalo-Weaver Bubalornis niger and other weavers; once sited only 40 cm above ground in hollow towbar of caravan (Steyn 2000). Both sexes build nest; in one case 1 bird brought nest material while other perched nearby (Uys and Underhill 1992). EGGS: (southem Africa, n ¼ 33) 2–5 (33), usually 3–4. Bluish to greenish white, heavily spotted and blotched with grey or brown; sometimes almost uniform chocolate brown (Maclean 1993). SIZE (southern Africa, n ¼ 103): 17–213  131–152 (192  143). ^ Dec (1), LAYING DATES: Tanzania (Pemba), July; Malawi, Jan (8), Feb (8), Mar (3), Apr (3), May (5), June (1); Zambia, Jan–Apr; Zimbabwe, Sept (8), Oct (5), Nov (10), Dec (23), Jan (34), Feb (32), Mar (23), Apr (8), May (n ¼ ?), June (1), July (n ¼ ?); Angola (probably July); Namibia, Dec–May; Mozambique, Jan–Mar, June, Sept–Dec; Botswana, Oct (2), Nov (1), Dec (5), Jan (4), Feb (7), Mar (9), Apr (1); South Africa, Oct–Mar. In southern Africa, breeding peaks in S of range in early summer, late summer in N, particularly arid W (Harrison et al. 1997). INCUBATION: by both sexes. DEVELOPMENT AND CARE OF YOUNG: X (and Y?) broods young a few days old for periods of 3–10 min; both feed young; young a few days old fed 28 times in 150 min. Fledging period 16–25 days. Fledglings fed by both parents and roost in nest with them; once they returned to nest nightly for 34 days before being turned away by parents (Cook and Cook 1998). BREEDING SUCCESS/SURVIVAL: southern Africa (n ¼ 5) 2–4 (26) fledged young per nest. Parasitized by African ^ lived 50–55 years. Cuckoo Cuculus gularis. 1 bird (Malawi) Key References Benson, C. W. (1956), Dowsett, R. J. and Dowsett-Lemaire, F. (1980), Craig, A. J. F. K. et al. (1987), Harrison, J. A. et al. (1997), Summers-Smith, J. D. (1988).

Passer motitensis A. Smith. Great Sparrow. Grand Moineau.

Plate 2

Passer motitensis A. Smith, 1848. Ill. Zool. S. Afr., Aves, pl. 114; 60 miles south of Orange River; restricted to near Hopetown, northern Cape Prov., by Macdonald (1957, Contr. Orn. W. South Africa, p. 157).

(Opp. p. 11)

Forms a superspecies with P. rufocinctus, P. shelleyi, P. cordofanicus, P. insularis and P. iagoensis. Range and Status. Endemic resident, southern Africa. SW Angola from SW Namibe north to coastal plain of Benguela;

Namibia, throughout except Namib desert; rare or absent in broad-leaved woodland in NE and Caprivi Strip.

Passer motitensis General Habits. Inhabits primarily open acacia and broadleaved woodland including mopane and miombo; occurrence in wide range of other habitats (open plains, grassland, savanna) due to association with man; occupies farmyards, cattle camps, plantations of exotics, gardens and villages. Common in thorn tree savanna (but replaced in broad-leaved savanna by Yellow-throated Sparrow Petronia superciliaris: Ginn et al. 1989). Rare in fynbos and succulent Karoo habitats; avoids forest and dense woodland. Where sympatric with House Sparrow in urban environments, favours suburban habitats; around farm houses often displaced by House Sparrow, returning if House Sparrow removed (Ginn et al. 1989). Where sympatric with Northern Grey-headed Sparrow, prefers mopane and other dry woodland away from man (Northern more in cultivation and around settlements). Rarely competes with Cape Sparrow P. melanurus which is much more adapted to man; when in same area usually loses nest-disputes with it (Summers-Smith 1988). Occurs in pairs, outside breeding season also in small flocks (probably family groups), sometimes flocks of 50–60 or even 500. Forages for seeds on bare ground, including cement paving, road verges and inside open sheds and barns (Ginn et al. 1989), often with Cape, House and Great Sparrows and other seedeaters. Walks as well as hops (very unusual in Passer: Summers-Smith 1988) which distinguishes it in mixed flocks from House Sparrows (A. J. F. K. Craig in Summers-Smith 1988). When not feeding, usually in trees not far from water. Foliage-bathes after rain (Tyler 1999). Moult follows breeding season; post-juvenile moult complete. Mainly sedentary, with some local movements. 5 recoveries of ringed birds in Zimbabwe less than 10 km from where ringed. Less frequent in winter in NW and extreme S areas (Zones 1–3, 8) (Harrison et al. 1997); birds common June but absent Nov (Botswana: Brewster 1991). Population at Mwaku Pan, NW Botswana, nomadic (Tree 1972). Food. Seeds and insects, including termite alates (latter also fed to young); also buds, berries, aloe nectar and food scraps. Breeding Habits. Solitary nester, occasionally in loose colonies; monogamous. Early in breeding season Y and X fly together to investigate nest-sites, and engage in display chases. At nest-site, calls repeatedly; when excited ruffles lower back and rump feathers, and flicks tail from side to

9

side in rotary motion. Threatens competing conspecifics and congeners (e.g. House Sparrow) by holding wings away from body and raising and spreading tail. Fights occur, bird grasping opponent with bill or claws and both falling to ground; when pairs fight, birds attack opponent of same sex (Summers-Smith 1988). Double-brooded. NEST: untidy bunch of grass, sometimes small twigs, lined with feathers, wool and hair; placed in hole in tree, including old nest hole of woodpecker or barbet, also under eaves or in thatched roof of building, hole in wall, fencepost or telegraph pole, hollow pipe, nest box and in abandoned farm or mine machinery; also uses old nests of Greater Striped Swallow Hirundo cucullata, Little Swift Apus affinis, Red-billed Buffalo-Weaver Bubalornis niger and other weavers; once sited only 40 cm above ground in hollow towbar of caravan (Steyn 2000). Both sexes build nest; in one case 1 bird brought nest material while other perched nearby (Uys and Underhill 1992). EGGS: (southem Africa, n ¼ 33) 2–5 (33), usually 3–4. Bluish to greenish white, heavily spotted and blotched with grey or brown; sometimes almost uniform chocolate brown (Maclean 1993). SIZE (southern Africa, n ¼ 103): 17–213  131–152 (192  143). ^ Dec (1), LAYING DATES: Tanzania (Pemba), July; Malawi, Jan (8), Feb (8), Mar (3), Apr (3), May (5), June (1); Zambia, Jan–Apr; Zimbabwe, Sept (8), Oct (5), Nov (10), Dec (23), Jan (34), Feb (32), Mar (23), Apr (8), May (n ¼ ?), June (1), July (n ¼ ?); Angola (probably July); Namibia, Dec–May; Mozambique, Jan–Mar, June, Sept–Dec; Botswana, Oct (2), Nov (1), Dec (5), Jan (4), Feb (7), Mar (9), Apr (1); South Africa, Oct–Mar. In southern Africa, breeding peaks in S of range in early summer, late summer in N, particularly arid W (Harrison et al. 1997). INCUBATION: by both sexes. DEVELOPMENT AND CARE OF YOUNG: X (and Y?) broods young a few days old for periods of 3–10 min; both feed young; young a few days old fed 28 times in 150 min. Fledging period 16–25 days. Fledglings fed by both parents and roost in nest with them; once they returned to nest nightly for 34 days before being turned away by parents (Cook and Cook 1998). BREEDING SUCCESS/SURVIVAL: southern Africa (n ¼ 5) 2–4 (26) fledged young per nest. Parasitized by African ^ lived 50–55 years. Cuckoo Cuculus gularis. 1 bird (Malawi) Key References Benson, C. W. (1956), Dowsett, R. J. and Dowsett-Lemaire, F. (1980), Craig, A. J. F. K. et al. (1987), Harrison, J. A. et al. (1997), Summers-Smith, J. D. (1988).

Passer motitensis A. Smith. Great Sparrow. Grand Moineau.

Plate 2

Passer motitensis A. Smith, 1848. Ill. Zool. S. Afr., Aves, pl. 114; 60 miles south of Orange River; restricted to near Hopetown, northern Cape Prov., by Macdonald (1957, Contr. Orn. W. South Africa, p. 157).

(Opp. p. 11)

Forms a superspecies with P. rufocinctus, P. shelleyi, P. cordofanicus, P. insularis and P. iagoensis. Range and Status. Endemic resident, southern Africa. SW Angola from SW Namibe north to coastal plain of Benguela;

Namibia, throughout except Namib desert; rare or absent in broad-leaved woodland in NE and Caprivi Strip.

12

PASSERIDAE

Botswana, local throughout, except extreme NE; SW and S Zimbabwe, uncommon to frequent south of c. 19
S in W, 21
S in E; South Africa, locally common to uncommon, N Cape Prov. south to c. 27
S; W and N Free State, some records south to c. 30
S, SE; N, central and W Transvaal east to c. 30
E, also SW Kalahari thornveld, absent SE; once Swaziland (Clancey 1964). Density, 1 pair per 100– 500 ha (Botswana, Transvaal: Summers-Smith 1988). Range in arid areas may be expanding with sinking of boreholes, yet contracting locally elsewhere, possible due to woodland clearance (Harrison et al. 1997). Description. P. m. motitensis A. Smith: Botswana and N Cape. ADULT Y: forehead to hindneck, upper mantle and upper side of neck grey, tinged brownish when fresh; lower mantle and scapulars dull cinnamon with short, rather narrow black streaks, well demarcated from top of head; outer scapulars and back to short uppertail-coverts plain cinnamon, more rufous on rump; long uppertail-coverts grey-brown. Tail feathers dark grey-brown, both webs of T1 and outer webs of T2–T6 narrowly fringed buff. Supraloral stripe greyish white, merging above eye with broad cinnamon crescent along side of crown and nape and behind earcoverts. Lores, below eye and broad stripe behind eye black, scaled with cinnamon. Lower ear-coverts, cheeks and lower side of neck greyish white. Chin and throat to border of upper breast black; upper breast light ashy grey, grading to greyish white on flanks, belly, thighs and undertail-coverts. Flight feathers, primary coverts and alula dark grey-brown, narrowly edged dull cinnamon-buff; tertials and greater coverts dark grey-brown, broadly edged and tipped dull cinnamon; median coverts blackish brown, broadly tipped greyish white; lesser coverts rufous-cinnamon. Underwing-coverts and axillaries pale greybuff. Bill black; eyes dark brown; legs grey or slate. ADULT X: top of head to upper mantle drab brown, faintly streaked darker brown, rest of upperparts cinnamon-brown, more rufous on rump. Broad buff superciliary stripe from above lores to side of nape; below this a broad dusky brown band through lores, below eye and through upper ear-coverts; upper side of neck drab brown; lower ear-coverts, cheeks, sides of chin and throat and lower side of neck pale buff. Centre of chin and throat dusky grey; rest of underparts grey-buff, whiter from lower breast to undertail-coverts. Tail, wing and bare parts as in Y. SIZE (6 YY, 5 XX): wing, Y 85–89 (868), X 83–86 (852); tail, Y 56–60 (587), X 57–60 (582); bill, Y 16–17 (165), X 155–165 (160); tarsus, Y 21–215 (212), X 21–22 (214). WEIGHT: (this race? and subsolanus) Y (n ¼ 3) 34–358 (347), X (n ¼ 4) 306–32 (315), unsexed (n ¼ 8) 29–326 (31). IMMATURE: like X. NESTLING: naked at hatching. P. m. subsolanus Clancey: Zimbabwe, Transvaal, Free State and Swaziland. Like nominate race but deeper rufous-cinnamon above, streaking rather heavier; slightly greyer below; Y has grey of head darker. P. m. benguellensis (Lynes): Angola and Namibia. Like nominate race, but lighter cinnamon (less rufous) above, slightly paler below. Y has face patch whiter; X has top of head greyer brown than nominate race, superciliary stripe whiter. Smaller: wing, Y (n ¼ 10) 79–84 (808).

Field Characters. Length 15–16 cm. A large sparrow, superficially similar to House Sparrow P. domesticus, with black bib and eye-line and grey crown, but not found in towns. Chestnut areas redder and paler than House Sparrow, rump bright chestnut, not grey; bib narrower, not reaching sides of neck; narrow white line above black lores. X much more colourful than X House Sparrow, with pale chestnut

Passer motitensis

sides of head, back, rump and shoulders, black line through eye and around ear-coverts. Immature like X. Voice. Tape-recorded (88, 99, F). Calls fuller and deeper than House Sparrow, ‘cheerr’ or ‘wer-cheerr’, often downslurred, ‘chyurr’, ‘tseeurr’. Dry rattle slower and deeper than House Sparrow, ‘bakabakabak . . . ’; also has a fast ‘churrrrr’ and nasal calls, ‘jwer’ or ‘jwoyer’. Song, ‘cheeurr’ and other call-notes repeated at uneven tempo. Pair at nest greet each other with soft, conversational ‘chee-ti-cheet’, ‘ ti-cheet-it’, ‘ti-chee-tit-tit’ or ‘chee-wee’ (Summers-Smith 1988). General Habits. Inhabits dry savanna and scrub, favouring open, rather arid acacia bushveld. Generally not associated with man although sometimes nests close to buildings in parks and rest camps (Ginn et al. 1989); frequents acacia-shaded kraals and fallow land under tribal agriculture. Less sociable or gregarious than other Passer spp. usually in pairs throughout year; forms flocks at water holes, sometimes with ‘immensely large’ concentrations (May and June 1970, Namibia: Tree 1972). Occasionally associates with flocks of Cape P. melanurus, Southern Grey-headed P. diffusus and House Sparrows. Shy and easily overlooked, although occasionally confiding, permitting close approach. Regularly feeds on ground, sometimes in trees; drinks often. In Namibia begins post-breeding moult in Apr. Mainly sedentary; some post-breeding season wandering in Botswana, Zimbabwe and Transvaal. Food. Grain, small seeds, some insects; caterpillars fed to young. Breeding Habits. Solitary nester. Monogamous but not known if it pairs for life. Courtship behaviour not recorded.

Passer rufocinctus NEST: untidy domed structure with side entrance through tunnel leading to egg chamber; built mainly of dried grass, green leaves (e.g. Asparagus) and pieces of string; egg chamber lined with feathers, fine grass and plant down. SIZE: about the size of a rugby football (SummersSmith 1988). Placed in branches of a bush or thorn tree, 15–65 m above ground. Built by both sexes. EGGS: southern Africa (n ¼ 9) 2–4 (33), sometimes up to 6. Oval, slightly glossy; greyish white, blotched with slate-grey and lavender. SIZE: southern Africa (n ¼ 82) 185–222  14–161 (205  150). LAYING DATES: Angola (juvenile Oct); Namibia, May– Aug, Dec–Feb; Botswana, Sept (1), Oct (2), Nov (10), Dec (16), Jan (3), Mar (2), Apr (?), May (?); Zimbabwe, Sept (1), Oct (4), Nov (3), Dec (1), Feb (2); South Africa, Oct–Feb;

13

Transvaal, Sept (1), Oct (1), Nov (4), Dec (5), Jan (3), Feb (1), Mar (4). Breeds mainly in wet summer season, possibly with peak in Jan–Apr in NW (Namibia, W Botswana, W Free State) and elsewhere a peak in Oct–Jan (Harrison et al. 1997). INCUBATION: by both sexes; period 12–14 days. DEVELOPMENT AND CARE OF YOUNG: probably cared for by both parents which feed insects to young until it is fledged; nestling period 15–18 days. BREEDING SUCCESS/SURVIVAL: regularly parasitized by Diederik Cuckoo Chrysococcyx caprius. Key References Harrison, J. A. et al. (1997), Summers-Smith, J. D. (1983a, b, 1988).

Passer rufocinctus Finsch and Reichenow. Kenya Rufous Sparrow. Moineau roux de Kenya.

Plate 2

Passer rufocinctus Finsch and Reichenow, 1884. J. Orn., 32, p. 55; Lake Naivasha [Kenya].

(Opp. p. 11)

Forms a superspecies with P. motitensis, P. shelleyi, P. cordofanicus, P. insularis and P. iagoensis. Range and Status. Endemic resident, common, Kenya and Tanzania. W, central and S Kenya Rift Valley highlands and peripheral plateau, from Eldoret, Kerio valley, Maralal, Laikipia Plateau and N Uaso Nyiro R. south through Masai Mara Game Res., Narok, Kajiado, Thika, Machakos and Taita to Tanzania border; N Tanzania south to c. 5
S and from c. 33
E east through Serengeti Nat. Park and Crater Highlands to Arusha District and Mt Kilimanjaro. Description. ADULT Y: forehead to hindneck, upper mantle and upper side of neck grey, tinged brownish when fresh; lower mantle and inner scapulars dull rufous-cinnamon with broad black streaks formed by inner webs of feathers; outer scapulars uniform rufous-cinnamon; back, rump and shorter uppertail-coverts brighter rufous-cinnamon; long uppertail-coverts grey-brown. Tail feathers blackish, both webs of T1 and outer webs of T2–T6 narrowly fringed pale buff. Supraloral line greyish white, merging above eye with broad rufous-cinnamon crescent which extends along side of crown and nape and around ear-coverts to foreside of neck. Lores, narrowly below eye and short mark behind eye dusky grey. Ear-coverts dusky brownish grey, grading to greyish white on lower cheek and lower side of neck. Chin and throat black, forming short bib which barely extends to centre of upper breast; rest of underparts pale brownish grey, grading to whitish on centre of lower breast and belly. Flight feathers, primary coverts and alula blackish brown, narrowly edged pale buff; tertials and greater coverts blackish brown with cinnamon-buff fringes, narrower, more whitish on greater covert tips; median coverts blackish with broad greyish white tips; lesser coverts rufous. Underwing-coverts whitish with dusky bases; axillaries pale grey-brown. Bill slate; eyes creamy or creamy tan; legs dull brown or grey-brown. ADULTX: similar to ad. Y but lacks bright head pattern. Forehead to hindneck and mantle grey-brown; broad superciliary stripe light buff; lores and stripe behind eye dusky brown; ear-coverts and cheeks light grey-brown; small bib on chin and throat dull grey, bordered by buffy white at sides. Breast and flanks browner than Y. SIZE (10 YY, 10 XX): wing, Y 74–80 (776), X 73–81 (767); tail, Y 48–55 (512), X 49–54 (515); bill, Y 135–15 (144), X 135–15 (141); tarsus, Y 19–21 (200), X 195–205 (202). WEIGHT: Y (n ¼ 27) 25–32 (275), X (n ¼ 37) 25– 32 (281).

Passer rufocinctus

IMMATURE: juvenile like ad. X but lacks grey ‘bib’, chin to breast light brownish grey; top of head to mantle browner, back and rump tawny-brown; eyes dark brown. NESTLING: naked at hatching; gape and edges of mandibles yellow; bill dark olive-brown with yellow edges; interior of mouth flesh pink; eyes grey-brown; legs grey.

Field Characters. Length 13–14 cm. A colourful sparrow of Kenya and N Tanzania highlands, with pale eye (both sexes). Y has pale rufous line over eye and around ear-coverts onto sides of neck, dusky grey stripe from lores to behind eye, rufous shoulder patch and rump and

Passer rufocinctus NEST: untidy domed structure with side entrance through tunnel leading to egg chamber; built mainly of dried grass, green leaves (e.g. Asparagus) and pieces of string; egg chamber lined with feathers, fine grass and plant down. SIZE: about the size of a rugby football (SummersSmith 1988). Placed in branches of a bush or thorn tree, 15–65 m above ground. Built by both sexes. EGGS: southern Africa (n ¼ 9) 2–4 (33), sometimes up to 6. Oval, slightly glossy; greyish white, blotched with slate-grey and lavender. SIZE: southern Africa (n ¼ 82) 185–222  14–161 (205  150). LAYING DATES: Angola (juvenile Oct); Namibia, May– Aug, Dec–Feb; Botswana, Sept (1), Oct (2), Nov (10), Dec (16), Jan (3), Mar (2), Apr (?), May (?); Zimbabwe, Sept (1), Oct (4), Nov (3), Dec (1), Feb (2); South Africa, Oct–Feb;

13

Transvaal, Sept (1), Oct (1), Nov (4), Dec (5), Jan (3), Feb (1), Mar (4). Breeds mainly in wet summer season, possibly with peak in Jan–Apr in NW (Namibia, W Botswana, W Free State) and elsewhere a peak in Oct–Jan (Harrison et al. 1997). INCUBATION: by both sexes; period 12–14 days. DEVELOPMENT AND CARE OF YOUNG: probably cared for by both parents which feed insects to young until it is fledged; nestling period 15–18 days. BREEDING SUCCESS/SURVIVAL: regularly parasitized by Diederik Cuckoo Chrysococcyx caprius. Key References Harrison, J. A. et al. (1997), Summers-Smith, J. D. (1983a, b, 1988).

Passer rufocinctus Finsch and Reichenow. Kenya Rufous Sparrow. Moineau roux de Kenya.

Plate 2

Passer rufocinctus Finsch and Reichenow, 1884. J. Orn., 32, p. 55; Lake Naivasha [Kenya].

(Opp. p. 11)

Forms a superspecies with P. motitensis, P. shelleyi, P. cordofanicus, P. insularis and P. iagoensis. Range and Status. Endemic resident, common, Kenya and Tanzania. W, central and S Kenya Rift Valley highlands and peripheral plateau, from Eldoret, Kerio valley, Maralal, Laikipia Plateau and N Uaso Nyiro R. south through Masai Mara Game Res., Narok, Kajiado, Thika, Machakos and Taita to Tanzania border; N Tanzania south to c. 5
S and from c. 33
E east through Serengeti Nat. Park and Crater Highlands to Arusha District and Mt Kilimanjaro. Description. ADULT Y: forehead to hindneck, upper mantle and upper side of neck grey, tinged brownish when fresh; lower mantle and inner scapulars dull rufous-cinnamon with broad black streaks formed by inner webs of feathers; outer scapulars uniform rufous-cinnamon; back, rump and shorter uppertail-coverts brighter rufous-cinnamon; long uppertail-coverts grey-brown. Tail feathers blackish, both webs of T1 and outer webs of T2–T6 narrowly fringed pale buff. Supraloral line greyish white, merging above eye with broad rufous-cinnamon crescent which extends along side of crown and nape and around ear-coverts to foreside of neck. Lores, narrowly below eye and short mark behind eye dusky grey. Ear-coverts dusky brownish grey, grading to greyish white on lower cheek and lower side of neck. Chin and throat black, forming short bib which barely extends to centre of upper breast; rest of underparts pale brownish grey, grading to whitish on centre of lower breast and belly. Flight feathers, primary coverts and alula blackish brown, narrowly edged pale buff; tertials and greater coverts blackish brown with cinnamon-buff fringes, narrower, more whitish on greater covert tips; median coverts blackish with broad greyish white tips; lesser coverts rufous. Underwing-coverts whitish with dusky bases; axillaries pale grey-brown. Bill slate; eyes creamy or creamy tan; legs dull brown or grey-brown. ADULTX: similar to ad. Y but lacks bright head pattern. Forehead to hindneck and mantle grey-brown; broad superciliary stripe light buff; lores and stripe behind eye dusky brown; ear-coverts and cheeks light grey-brown; small bib on chin and throat dull grey, bordered by buffy white at sides. Breast and flanks browner than Y. SIZE (10 YY, 10 XX): wing, Y 74–80 (776), X 73–81 (767); tail, Y 48–55 (512), X 49–54 (515); bill, Y 135–15 (144), X 135–15 (141); tarsus, Y 19–21 (200), X 195–205 (202). WEIGHT: Y (n ¼ 27) 25–32 (275), X (n ¼ 37) 25– 32 (281).

Passer rufocinctus

IMMATURE: juvenile like ad. X but lacks grey ‘bib’, chin to breast light brownish grey; top of head to mantle browner, back and rump tawny-brown; eyes dark brown. NESTLING: naked at hatching; gape and edges of mandibles yellow; bill dark olive-brown with yellow edges; interior of mouth flesh pink; eyes grey-brown; legs grey.

Field Characters. Length 13–14 cm. A colourful sparrow of Kenya and N Tanzania highlands, with pale eye (both sexes). Y has pale rufous line over eye and around ear-coverts onto sides of neck, dusky grey stripe from lores to behind eye, rufous shoulder patch and rump and

14

PASSERIDAE

black-and-rufous striped back; black bib smaller than House Sparrow P. domesticus. X like Y but rufous areas less bright, tawny-buff rather than red, throat patch grey. Immatures like respective adults but duller, eye brown. Comes close to, but does not meet, Shelley’s Rufous Sparrow P. shelleyi in Kenya/Uganda border area; for differences, see that species. Voice. Tape recorded (104, GREG, McVIC). Calls include loud ‘chreep’, ‘chuweep’ or upward ‘cher-weep’, nasal ‘jewy’ or ‘jerwy’ and short ‘jer-ju’; also sharp ‘tsui’ or ‘tseu pee’, and high-pitched, squeaky ‘tsweet’. Song, thin, sharp metallic notes often alternating with a few lower chirps, ‘tseup chree, tseup chree, tsweet shreep . . . ’, or a repeated ‘chireet’ (Zimmerman et al. 1996). General Habits. Occurs mainly in dry acacia savanna, open wooded country, edges of forests; also grazed land, cultivated areas and around towns and villages. Altitudinal range 1000–3000 m, mainly 1400–2000; nests largely above 1500 m (Kenya: Lewis and Pomeroy 1989). Usually solitary or in pairs throughout year, sometimes in small, probably family flocks of up to 10 in non-breeding season. Behaviour much like House Sparrow; confiding and easily approached in built-up areas. Feeds more on ground than Great Sparrow P. motitensis (Summers-Smith 1988). Once 2 birds joined dancing ring of 10 Anteater Chats Myrmecocichla aethiops and 3 Superb Starlings Lamprotornis superbus, took turns leaping vertically into air, hovering above group for c. 10 s, then dropping back to the ground (Hayes 1982); reason for this behaviour unknown. Moult in Nakuru (Kenya) begins June–July and is completed by Mar–Apr. Mainly sedentary; some non-breeding birds disperse to lower altitudes in Kenya. Food. Grain, small seeds and insects, also household scraps; insects fed to young until they fledge.

Breeding Habits. Solitary nester; presumably monogamous but not known if it pairs for life. Early in courtship Y approaches X, chirruping; then puffs up rufous rump feathers, lowers back, cocks tail up at an angle of c. 45
and points head and bill nearly vertically upward, exposing black throat and chin. Once 2 YY fought on ground with feet in each other’s faces, pecking at each other’s throats; a few other sparrows crowded around them twittering. After 6–7 min the 2 flew to a tree, fought there, fell to ground and fought again. After they disengaged, 1 pursued the other for quite some time in the tree and a nearby bush; dispute appeared to be territorial (Rudnai 1991). Possibly doublebroaded (Betts 1966). NEST: large ovoid mass, rather flat on top, with single side entrance; loosely constructed of grasses, mainly runners of Cynodon dactylon, also Digitaria scalarum, Aristida adoensis and Setaria sp.; lined with fine rootlets, grasses, feathers and hair. Ext. diam. 250, depth 200; entrance opens to 1 side of roof and extends diagonally 100–130 mm under it to egg chamber. Placed among branches of small tree, 1–65 m above ground; sometimes in ant-gall acacia Acacia drepanolobium, or in covered site such as hole in tree, under eaves of house, cavity in concrete electricity pylon or deserted weaver nest, also among fruits of banana plants. Built by both sexes. EGGS: Kenya (n ¼ 6) 3–4, (n ¼ 5) 3–5 (38); sometimes 6. Oval, but shape varies; slightly glossy; greyish to bluish white with blotches of tawny brown, umber dark grey and pale lilac. SIZE: Kenya (n ¼ 13) 209–219  142–155 (208  150). LAYING DATES: E Africa: Kenya, Feb–July, Oct; Tanzania Feb, Apr, June, Dec; Region C, Mar; Region D, Jan (1), Feb (2), Mar (3), Apr (11), May (13), June (6), July (4), Sept (1), Oct (1), Nov (6), Dec (3), with peaks Apr–July in long rains and Nov–Dec in short rains.

Key References Collias, N. E. and Collias, E. C. (1964), Summers-Smith, J. D. (1988).

Plate 2

Passer shelleyi Sharpe. Shelley’s Rufous Sparrow. Moineau roux de Shelley.

(Opp. p. 11)

Passer shelleyi Sharpe, 1891, Ibis, p. 256; Lado [southern Sudan]. Forms a superspecies with P. motitensis, P. rufocinctus, P. cordofanicus, P. insularis and P. iagoensis. Range and Status. Endemic resident, uncommon to rare except in Ethiopia. SE Sudan east of c. 31
E and south of c. 6
N. S and E Ethiopia in 2 areas, (1) between 3
300 –5
300 N and 37
300 –40
300 E south to Kenya border; and (2) 9–10
N and 43–44
E along Ethiopia/Somalia border where more numerous on Ethiopian side; also 1 record (1971) at 7
450 N, 43
150 E (J. S. Ash, pers. comm., Ash and Miskell 1998). On Somalia side of border, 7 records in 1898–1958, none recently (Ash and Miskell 1998). NE Uganda in Teso, Bugisu, Sebei and Karamoja provinces, west to Kidepo Nat. Park; also once Murchison Falls (Rossouw 2001).

Kenya along Uganda border around Kunyao (1
470 N, 35
030 E) and at the base of the Kongelai Escarpment (1
250 N, 35
030 E); old single record from N Kerio valley. Description. ADULT Y: forehead to hindneck, upper mantle and rear side of neck brownish grey, lower mantle and inner scapulars pale cinnamon-brown, broadly streaked with black; outer scapulars uniform cinnamon-brown; back, rump and shorter uppertail-coverts cinnamon; long uppertail-coverts grey-brown. Tail feathers dark grey-brown, both webs of T1 and outer webs of T2–T6 narrowly fringed pale buff. Supraloral stripe buffy white, merging above eye with broad pale cinnamon crescent which

14

PASSERIDAE

black-and-rufous striped back; black bib smaller than House Sparrow P. domesticus. X like Y but rufous areas less bright, tawny-buff rather than red, throat patch grey. Immatures like respective adults but duller, eye brown. Comes close to, but does not meet, Shelley’s Rufous Sparrow P. shelleyi in Kenya/Uganda border area; for differences, see that species. Voice. Tape recorded (104, GREG, McVIC). Calls include loud ‘chreep’, ‘chuweep’ or upward ‘cher-weep’, nasal ‘jewy’ or ‘jerwy’ and short ‘jer-ju’; also sharp ‘tsui’ or ‘tseu pee’, and high-pitched, squeaky ‘tsweet’. Song, thin, sharp metallic notes often alternating with a few lower chirps, ‘tseup chree, tseup chree, tsweet shreep . . . ’, or a repeated ‘chireet’ (Zimmerman et al. 1996). General Habits. Occurs mainly in dry acacia savanna, open wooded country, edges of forests; also grazed land, cultivated areas and around towns and villages. Altitudinal range 1000–3000 m, mainly 1400–2000; nests largely above 1500 m (Kenya: Lewis and Pomeroy 1989). Usually solitary or in pairs throughout year, sometimes in small, probably family flocks of up to 10 in non-breeding season. Behaviour much like House Sparrow; confiding and easily approached in built-up areas. Feeds more on ground than Great Sparrow P. motitensis (Summers-Smith 1988). Once 2 birds joined dancing ring of 10 Anteater Chats Myrmecocichla aethiops and 3 Superb Starlings Lamprotornis superbus, took turns leaping vertically into air, hovering above group for c. 10 s, then dropping back to the ground (Hayes 1982); reason for this behaviour unknown. Moult in Nakuru (Kenya) begins June–July and is completed by Mar–Apr. Mainly sedentary; some non-breeding birds disperse to lower altitudes in Kenya. Food. Grain, small seeds and insects, also household scraps; insects fed to young until they fledge.

Breeding Habits. Solitary nester; presumably monogamous but not known if it pairs for life. Early in courtship Y approaches X, chirruping; then puffs up rufous rump feathers, lowers back, cocks tail up at an angle of c. 45
and points head and bill nearly vertically upward, exposing black throat and chin. Once 2 YY fought on ground with feet in each other’s faces, pecking at each other’s throats; a few other sparrows crowded around them twittering. After 6–7 min the 2 flew to a tree, fought there, fell to ground and fought again. After they disengaged, 1 pursued the other for quite some time in the tree and a nearby bush; dispute appeared to be territorial (Rudnai 1991). Possibly doublebroaded (Betts 1966). NEST: large ovoid mass, rather flat on top, with single side entrance; loosely constructed of grasses, mainly runners of Cynodon dactylon, also Digitaria scalarum, Aristida adoensis and Setaria sp.; lined with fine rootlets, grasses, feathers and hair. Ext. diam. 250, depth 200; entrance opens to 1 side of roof and extends diagonally 100–130 mm under it to egg chamber. Placed among branches of small tree, 1–65 m above ground; sometimes in ant-gall acacia Acacia drepanolobium, or in covered site such as hole in tree, under eaves of house, cavity in concrete electricity pylon or deserted weaver nest, also among fruits of banana plants. Built by both sexes. EGGS: Kenya (n ¼ 6) 3–4, (n ¼ 5) 3–5 (38); sometimes 6. Oval, but shape varies; slightly glossy; greyish to bluish white with blotches of tawny brown, umber dark grey and pale lilac. SIZE: Kenya (n ¼ 13) 209–219  142–155 (208  150). LAYING DATES: E Africa: Kenya, Feb–July, Oct; Tanzania Feb, Apr, June, Dec; Region C, Mar; Region D, Jan (1), Feb (2), Mar (3), Apr (11), May (13), June (6), July (4), Sept (1), Oct (1), Nov (6), Dec (3), with peaks Apr–July in long rains and Nov–Dec in short rains.

Key References Collias, N. E. and Collias, E. C. (1964), Summers-Smith, J. D. (1988).

Plate 2

Passer shelleyi Sharpe. Shelley’s Rufous Sparrow. Moineau roux de Shelley.

(Opp. p. 11)

Passer shelleyi Sharpe, 1891, Ibis, p. 256; Lado [southern Sudan]. Forms a superspecies with P. motitensis, P. rufocinctus, P. cordofanicus, P. insularis and P. iagoensis. Range and Status. Endemic resident, uncommon to rare except in Ethiopia. SE Sudan east of c. 31
E and south of c. 6
N. S and E Ethiopia in 2 areas, (1) between 3
300 –5
300 N and 37
300 –40
300 E south to Kenya border; and (2) 9–10
N and 43–44
E along Ethiopia/Somalia border where more numerous on Ethiopian side; also 1 record (1971) at 7
450 N, 43
150 E (J. S. Ash, pers. comm., Ash and Miskell 1998). On Somalia side of border, 7 records in 1898–1958, none recently (Ash and Miskell 1998). NE Uganda in Teso, Bugisu, Sebei and Karamoja provinces, west to Kidepo Nat. Park; also once Murchison Falls (Rossouw 2001).

Kenya along Uganda border around Kunyao (1
470 N, 35
030 E) and at the base of the Kongelai Escarpment (1
250 N, 35
030 E); old single record from N Kerio valley. Description. ADULT Y: forehead to hindneck, upper mantle and rear side of neck brownish grey, lower mantle and inner scapulars pale cinnamon-brown, broadly streaked with black; outer scapulars uniform cinnamon-brown; back, rump and shorter uppertail-coverts cinnamon; long uppertail-coverts grey-brown. Tail feathers dark grey-brown, both webs of T1 and outer webs of T2–T6 narrowly fringed pale buff. Supraloral stripe buffy white, merging above eye with broad pale cinnamon crescent which

Passer cordofanicus extends along side of crown and nape and around ear-coverts to foreside of neck. Lores and narrowly below eye black, continuous with black stripe through upper ear-coverts, which turns down behind ear-coverts and borders cinnamon crescent. Lower earcoverts ashy grey, merging into buffy white cheeks and side of throat. Chin and centre of throat black, forming short bib which just extends to centre of upper breast; rest of underparts pale buffy brown, grading to buffy white on centre of lower breast, belly and undertail-coverts. Flight feathers, primary coverts and alula blackish brown, narrowly edged pale buff; tertials and greater coverts blackish brown with pale buff fringes, more whitish on greater covert tips; median coverts blackish with broad white tips, forming well-developed wing-bar; lesser coverts cinnamon. Underwing-coverts and axillaries buffy white. Bill slate; eyes brown; legs horn brown, grey-brown or slate. ADULT X: differs from ad. Y in having head pattern less bright; crown to upper mantle slightly browner, broad crescent behind eye cinnamon-buff, lores and stripe through and around upper earcoverts dusky grey-brown, cheeks buff; bib on chin and throat slightly smaller, dull grey, bordered by buffy white at sides; underparts slightly browner. SIZE (10 YY, 10 XX): wing, Y 69–76 (732), X 68–75 (718); tail, Y 42–48 (454), X 44–49 (458); bill, Y 13–145 (136), X 13–145 (137); tarsus Y 185–205 (194), X 185–195 (191). IMMATURE and NESTLING: not described.

Field Characters. Length 13–14 cm. The rufous sparrow of SE Sudan, Uganda, Ethiopia and (just) NW Kenya. Told from Kenya Rufous Sparrow P. rufocinctus by dark eye, black stripe through and behind eye curving down behind ear-coverts, below broad bright crescent which is cinnamon rather than rufous (both sexes). Grey of crown extends onto mantle, back grey-brown striped black, contrasting with bright rufous rump (mantle and back red in Kordofan Rufous Sparrow P. cordofanicus). Voice. Not known. General Habits. Inhabits mostly open, semi-arid grassy savanna and plains; especially partial to whistling thorns (Somalia); also occurs in open woodland and cultivated areas but not in and around villages; usually below 2000 m, in Ethiopia mainly at 900–1430 m (J. S. Ash, pers. comm.). Feeds regularly on ground; less sociable or gregarious than other Passer sparrows. Food. No data; presumably seeds, grain, and insects.

15

Passer shelleyi

Breeding Habits. Nest in small colonies (Somalia: Ash and Miskell 1998). NEST: untidy, ovoid, domed structure with side entrance; built of grasses, lined with feathers. Ext. diam. 250, depth 200; wedged in branches of small thorny acacia 1–2 m above ground. Built by both sexes. EGGS: 4–6, usually 4. White, streaked and blotched with rich umber-brown and violet-grey. SIZE: Somalia (n ¼ 8) av. 198  136. LAYING DATES: Sudan, Sept; Ethiopia, Oct–Nov; Somalia, June (nest-building Feb); Uganda, Apr.

Key Reference Summers-Smith, J. D. (1988).

Passer cordofanicus Heuglin. Kordofan Rufous Sparrow. Moineau roux du Kordofan.

Plate 2

Passer cordofanicus Heuglin, 1871. Orn. Nordost. Afr. Nachtr., p. 141; Kordofan (ex Passer motitensis, Heuglin, ibid., p. 634).

(Opp. p. 11)

Forms a superspecies with P. motitensis, P. rufocinctus, P. shelleyi, P. insularis and P. iagoensis. Range and Status. Endemic resident. W-central Sudan between 12
and 14
N in Darfur and Kordofan Provs, east to c. 30
E. E Chad from Sudan border to Oum Hadjer in W, Biltine and Iriba in N and Am Dam in S. Common to uncommon. Description. ADULT Y: forehead to hindneck, upper mantle and rear side of neck grey; lower mantle to scapulars and short

uppertail-coverts bright cinnamon, well demarcated from top of head, lower mantle and inner scapulars with short black streaks; long uppertail-coverts grey-brown. Tail feathers blackish, both webs of T1 and outer webs of T2–T6 narrowly fringed cinnamonbuff. Broad supraloral stripe buffy white, merging above eye with bright cinnamon crescent which extends along side of crown and nape and around ear-coverts. Lores, below eye and stripe through upper ear-coverts black; cheeks and lower ear-coverts to side of throat white, forming large face patch. Chin and centre of throat

Passer cordofanicus extends along side of crown and nape and around ear-coverts to foreside of neck. Lores and narrowly below eye black, continuous with black stripe through upper ear-coverts, which turns down behind ear-coverts and borders cinnamon crescent. Lower earcoverts ashy grey, merging into buffy white cheeks and side of throat. Chin and centre of throat black, forming short bib which just extends to centre of upper breast; rest of underparts pale buffy brown, grading to buffy white on centre of lower breast, belly and undertail-coverts. Flight feathers, primary coverts and alula blackish brown, narrowly edged pale buff; tertials and greater coverts blackish brown with pale buff fringes, more whitish on greater covert tips; median coverts blackish with broad white tips, forming well-developed wing-bar; lesser coverts cinnamon. Underwing-coverts and axillaries buffy white. Bill slate; eyes brown; legs horn brown, grey-brown or slate. ADULT X: differs from ad. Y in having head pattern less bright; crown to upper mantle slightly browner, broad crescent behind eye cinnamon-buff, lores and stripe through and around upper earcoverts dusky grey-brown, cheeks buff; bib on chin and throat slightly smaller, dull grey, bordered by buffy white at sides; underparts slightly browner. SIZE (10 YY, 10 XX): wing, Y 69–76 (732), X 68–75 (718); tail, Y 42–48 (454), X 44–49 (458); bill, Y 13–145 (136), X 13–145 (137); tarsus Y 185–205 (194), X 185–195 (191). IMMATURE and NESTLING: not described.

Field Characters. Length 13–14 cm. The rufous sparrow of SE Sudan, Uganda, Ethiopia and (just) NW Kenya. Told from Kenya Rufous Sparrow P. rufocinctus by dark eye, black stripe through and behind eye curving down behind ear-coverts, below broad bright crescent which is cinnamon rather than rufous (both sexes). Grey of crown extends onto mantle, back grey-brown striped black, contrasting with bright rufous rump (mantle and back red in Kordofan Rufous Sparrow P. cordofanicus). Voice. Not known. General Habits. Inhabits mostly open, semi-arid grassy savanna and plains; especially partial to whistling thorns (Somalia); also occurs in open woodland and cultivated areas but not in and around villages; usually below 2000 m, in Ethiopia mainly at 900–1430 m (J. S. Ash, pers. comm.). Feeds regularly on ground; less sociable or gregarious than other Passer sparrows. Food. No data; presumably seeds, grain, and insects.

15

Passer shelleyi

Breeding Habits. Nest in small colonies (Somalia: Ash and Miskell 1998). NEST: untidy, ovoid, domed structure with side entrance; built of grasses, lined with feathers. Ext. diam. 250, depth 200; wedged in branches of small thorny acacia 1–2 m above ground. Built by both sexes. EGGS: 4–6, usually 4. White, streaked and blotched with rich umber-brown and violet-grey. SIZE: Somalia (n ¼ 8) av. 198  136. LAYING DATES: Sudan, Sept; Ethiopia, Oct–Nov; Somalia, June (nest-building Feb); Uganda, Apr.

Key Reference Summers-Smith, J. D. (1988).

Passer cordofanicus Heuglin. Kordofan Rufous Sparrow. Moineau roux du Kordofan.

Plate 2

Passer cordofanicus Heuglin, 1871. Orn. Nordost. Afr. Nachtr., p. 141; Kordofan (ex Passer motitensis, Heuglin, ibid., p. 634).

(Opp. p. 11)

Forms a superspecies with P. motitensis, P. rufocinctus, P. shelleyi, P. insularis and P. iagoensis. Range and Status. Endemic resident. W-central Sudan between 12
and 14
N in Darfur and Kordofan Provs, east to c. 30
E. E Chad from Sudan border to Oum Hadjer in W, Biltine and Iriba in N and Am Dam in S. Common to uncommon. Description. ADULT Y: forehead to hindneck, upper mantle and rear side of neck grey; lower mantle to scapulars and short

uppertail-coverts bright cinnamon, well demarcated from top of head, lower mantle and inner scapulars with short black streaks; long uppertail-coverts grey-brown. Tail feathers blackish, both webs of T1 and outer webs of T2–T6 narrowly fringed cinnamonbuff. Broad supraloral stripe buffy white, merging above eye with bright cinnamon crescent which extends along side of crown and nape and around ear-coverts. Lores, below eye and stripe through upper ear-coverts black; cheeks and lower ear-coverts to side of throat white, forming large face patch. Chin and centre of throat

16

PASSERIDAE

to border of upper breast centre black, forming short bib; rest of underparts buffy white, linking around bib with face patch. Flight feathers, primary coverts and alula blackish brown, narrowly edged pale buffy brown; tertials dark grey-brown, broadly edged buffy brown; greater coverts dark grey-brown, edged pale greybuff, tipped whitish; median coverts blackish, broadly tipped white; lesser coverts cinnamon. Underwing-coverts and axillaries buffy white. Bill black; eyes dark brown; legs slaty or plumbeous grey. ADULT X: similar to ad. Y, but head pattern rather less bright: crown to upper mantle light ash-grey, stripe from lores to upper ear-coverts dusky grey, white face patch tinged buff, bib light ashy grey and slightly smaller. SIZE (10 YY, 10 XX): wing, Y 73–80 (758), X 72–75 (740); tail, Y 49–57 (532), X 45–52 (497); bill, Y 13–145 (136), X 13–14 (137); tarsus, Y 19–205 (199), X 19–205 (197). IMMATURE: juvenile like ad. X, but with no distinct bib; entire chin, throat and upper breast greyish white; upperparts rather duller and browner. NESTLING: naked at hatching.

Passer cordofanicus

?

Field Characters. Length 13–14 cm. Confined to E Chad and W-central Sudan; not known to meet Shelley’s Rufous Sparrow P. shelleyi. Distinguished from Shelley’s by bright cinnamon lower mantle and back, pure white face and underparts (other rufous sparrows are variably washed grey on face, breast and flanks). Eye dark; lores and lower border of rufous neck stripe black. Voice. Not tape-recorded. Said to be indistinguishable from those of other sparrows.

Food. Not known.

feathers. Placed in thorny bush or tree, 2–3 m above ground. EGGS: 3–6 (once 9, Chad). Dull white, boldly marked with lavender, maroon, brown and grey blotches. SIZE: Sudan (n ¼ 32) av. 19  135. LAYING DATES: Chad, June–Sept; Sudan, June–July, Sept–Nov (June–July nests abandoned and breeding did not begin until Sept: Lynes 1924).

Breeding Habits. NEST: untidy, ovoid, domed, with side entrance a tunnel c. 300 mm long; lined with grass and

Key References (1988).

General Habits. Inhabits arid bush country with average annual rainfall of c. 300–400 mm; usually around villages and surrounding cultivated land; occasionally on plains and in open savanna. Behaves like House Sparrow P. domesticus in villages and towns.

Lynes, H. (1924), Summers-Smith, J. D.

Plate 2

Passer insularis Sclater and Hartlaub. Socotra Sparrow. Moineau de Socotra.

(Opp. p. 11)

Passer insularis Sclater and Hartlaub, 1881. Proc. Zool. Soc. London, p. 169; Socotra. Forms a superspecies with P. motitensis, P. rufocinctus, P. shelleyi, P. cordofanicus and P. iagoensis. Range and Status. Endemic resident Socotra and Abd-elKuri; common and widespread, population on Socotra c. 250,000 (R. F. Porter, pers. comm.). Description. P. i. insularis Sclater and Hartlaub: Socotra. ADULT Y: forehead to upper mantle and rear side of neck grey, lower mantle and scapulars ashy grey, mantle and inner scapulars streaked blackish; back, rump and uppertail-coverts grey-brown. Tail feathers dark grey-brown, both webs of T1 and outer webs of T2–T6 narrowly edged buff. Supraloral stripe buff, merging above eye with broad cinnamon crescent extending along side of crown and nape and around ear-coverts. Lores, narrowly below eye, and broad stripe through upper ear-coverts turning down behind ear-coverts dull black. Upper cheeks to lower ear-coverts

dark grey, merging with large buffy white patch on lower cheeks, side of throat and foreside of neck. Chin and throat to bib on centre of upper breast sooty black; sides of breast, lower breast and flanks ashy grey, grading to greyish white on centre of belly and undertail-coverts. Flight feathers, primary coverts and alula dark grey-brown, narrowly edged cinnamon-buff; tertials and greater coverts dark grey-brown with cinnamon-buff fringes; median coverts blackish with broad greyish white tips; lesser coverts rufous-cinnamon. Underwing-coverts and axillaries pale grey-buff. Bill black; eyes dark brown; legs dark grey-brown or slate. ADULT X: similar to ad. Y, but head pattern much less bright; forehead to hindneck slightly browner, crescent behind eye dull buff and poorly defined, lores and stripe through upper ear-coverts dusky grey-brown, lower cheek patch more buffy; lacks distinct bib, dull greyish chin and throat merging with

16

PASSERIDAE

to border of upper breast centre black, forming short bib; rest of underparts buffy white, linking around bib with face patch. Flight feathers, primary coverts and alula blackish brown, narrowly edged pale buffy brown; tertials dark grey-brown, broadly edged buffy brown; greater coverts dark grey-brown, edged pale greybuff, tipped whitish; median coverts blackish, broadly tipped white; lesser coverts cinnamon. Underwing-coverts and axillaries buffy white. Bill black; eyes dark brown; legs slaty or plumbeous grey. ADULT X: similar to ad. Y, but head pattern rather less bright: crown to upper mantle light ash-grey, stripe from lores to upper ear-coverts dusky grey, white face patch tinged buff, bib light ashy grey and slightly smaller. SIZE (10 YY, 10 XX): wing, Y 73–80 (758), X 72–75 (740); tail, Y 49–57 (532), X 45–52 (497); bill, Y 13–145 (136), X 13–14 (137); tarsus, Y 19–205 (199), X 19–205 (197). IMMATURE: juvenile like ad. X, but with no distinct bib; entire chin, throat and upper breast greyish white; upperparts rather duller and browner. NESTLING: naked at hatching.

Passer cordofanicus

?

Field Characters. Length 13–14 cm. Confined to E Chad and W-central Sudan; not known to meet Shelley’s Rufous Sparrow P. shelleyi. Distinguished from Shelley’s by bright cinnamon lower mantle and back, pure white face and underparts (other rufous sparrows are variably washed grey on face, breast and flanks). Eye dark; lores and lower border of rufous neck stripe black. Voice. Not tape-recorded. Said to be indistinguishable from those of other sparrows.

Food. Not known.

feathers. Placed in thorny bush or tree, 2–3 m above ground. EGGS: 3–6 (once 9, Chad). Dull white, boldly marked with lavender, maroon, brown and grey blotches. SIZE: Sudan (n ¼ 32) av. 19  135. LAYING DATES: Chad, June–Sept; Sudan, June–July, Sept–Nov (June–July nests abandoned and breeding did not begin until Sept: Lynes 1924).

Breeding Habits. NEST: untidy, ovoid, domed, with side entrance a tunnel c. 300 mm long; lined with grass and

Key References (1988).

General Habits. Inhabits arid bush country with average annual rainfall of c. 300–400 mm; usually around villages and surrounding cultivated land; occasionally on plains and in open savanna. Behaves like House Sparrow P. domesticus in villages and towns.

Lynes, H. (1924), Summers-Smith, J. D.

Plate 2

Passer insularis Sclater and Hartlaub. Socotra Sparrow. Moineau de Socotra.

(Opp. p. 11)

Passer insularis Sclater and Hartlaub, 1881. Proc. Zool. Soc. London, p. 169; Socotra. Forms a superspecies with P. motitensis, P. rufocinctus, P. shelleyi, P. cordofanicus and P. iagoensis. Range and Status. Endemic resident Socotra and Abd-elKuri; common and widespread, population on Socotra c. 250,000 (R. F. Porter, pers. comm.). Description. P. i. insularis Sclater and Hartlaub: Socotra. ADULT Y: forehead to upper mantle and rear side of neck grey, lower mantle and scapulars ashy grey, mantle and inner scapulars streaked blackish; back, rump and uppertail-coverts grey-brown. Tail feathers dark grey-brown, both webs of T1 and outer webs of T2–T6 narrowly edged buff. Supraloral stripe buff, merging above eye with broad cinnamon crescent extending along side of crown and nape and around ear-coverts. Lores, narrowly below eye, and broad stripe through upper ear-coverts turning down behind ear-coverts dull black. Upper cheeks to lower ear-coverts

dark grey, merging with large buffy white patch on lower cheeks, side of throat and foreside of neck. Chin and throat to bib on centre of upper breast sooty black; sides of breast, lower breast and flanks ashy grey, grading to greyish white on centre of belly and undertail-coverts. Flight feathers, primary coverts and alula dark grey-brown, narrowly edged cinnamon-buff; tertials and greater coverts dark grey-brown with cinnamon-buff fringes; median coverts blackish with broad greyish white tips; lesser coverts rufous-cinnamon. Underwing-coverts and axillaries pale grey-buff. Bill black; eyes dark brown; legs dark grey-brown or slate. ADULT X: similar to ad. Y, but head pattern much less bright; forehead to hindneck slightly browner, crescent behind eye dull buff and poorly defined, lores and stripe through upper ear-coverts dusky grey-brown, lower cheek patch more buffy; lacks distinct bib, dull greyish chin and throat merging with

Passer melanurus buffy grey breast; rest of underparts paler grey-buff, lower breast and flanks with faint narrow streaks. SIZE: wing, Y (n ¼ 8) 75–80 (766), X (n ¼ 6) 72–77 (745); tail, Y (n ¼ 8) 55–60 (575), X (n ¼ 6) 48–57 (538); bill, Y (n ¼ 6) 15–17 (161), X (n ¼ 6) 145–17 (161); tarsus, Y (n ¼ 6) 205–22 (213), X (n ¼ 5) 21–22 (213). WEIGHT: Socotra, unsexed (n ¼ 51) 20–35 (Ripley and Bond 1966); Y (n ¼ 4) 250–292 (275), X (n ¼ 4) 260–291 (271), unsexed (n ¼ 3) 239–280 (257) (Dymond 1996). IMMATURE and NESTLING: not described. P. i. hemileucus Ogilvie-Grant and Forbes: Abd-el-Kuri. Much paler than insularis, underparts nearly pure white, black patch on chin of Y smaller, X without dusky patch on throat. Smaller; 1 Y wing 74, tail 55, 1 X wing 71, tail 54.

17

Passer insularis

Field Characters. Length 13–14 cm. The only sparrow on Socotra and Abd-el-Khuri. Y has upperparts entirely grey, including rump, unlike mainland forms; rufous confined to crescent on side of head and patch from wing shoulder to scapulars. X drab, like X House Sparrow P. domesticus but with black bill and eye-line; X Cinnamon-breasted and Socotra Buntings Emberiza tahapisi and E. socotrana have striped head, cinnamon on underparts. Voice. Tape-recorded (B, DAV). Dry chirping calls, ‘chirrup’, ‘cheep’, ‘chee-che´ep’, ‘chip’, ‘jip’ ‘chlip’; nasal scolding ‘jer-jer’ and faster ‘jajajajajaja . . . ’. General Habits. In Socotra occurs widely in all habitats from sea level to 1200 m, but especially Croton scrub on coastal plains; scrub, woodland and scrub/grass mosaic on foothills and arid limestone areas; and palm groves. Sometimes common in human settlements (R. P. Martins and R. F. Porter, pers. comm.). Occurs singly but most often in groups of 180 cm long; may remain intact for several years before disintegrating. Built by Y only, using twigs 150–750 long collected up to 1000 m away; material added throughout breeding season. Y brings some green material to line chambers; once X accepts nest, she adds lining of leaves, particularly acacia leaves, and green grass. Thorny roof over nest chambers may be 30 cm thick, floor 8–10 cm thick. One nest in Namibia built mainly by X; Y occasionally stole material from neighbours (Hoesch 1952). EGGS: 2–4, (av. 33, n ¼ 16, South Africa). White, spotted and blotched with sepia and grey, or blue-green heavily marked with olive-green and ash-grey. SIZE: (Somalia, n ¼ 12) 270–300  195–210 (278  200); (South Africa, n ¼ 25) 256–325  190–209 (283  199). LAYING DATES: Ethiopia, Mar–Sept; Somalia, Apr–June; Uganda, June; Kenya, Mar–May, Dec; Tanzania, Dec, Mar (recently–fledged bird, June); Angola, Nov, Feb; Botswana, Sept–Mar, mainly Nov–Mar, also May–June;

53

54

PLOCEIDAE

Zimbabwe, Sept–Apr, chiefly Jan–Feb; Mozambique, Sept–Nov; South Africa, Oct–Apr. INCUBATION: by X only, period 11 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only (Namibia), or (Kenya) by Y (1–2 feeds for every 24 by X); both adults once continued to feed young in nest mass cut down and resting on ground. Nestling period: 20–23 days. BREEDING SUCCESS/SURVIVAL: some nests deserted after attack by Wahlberg’s Eagles Aquila wahlbergi. Nests may be heavily infested with mites and insects; in Kenya, a

distinctive grey cockroach especially common. Of 16 eggs, 2 failed to hatch, 6 chicks fledged. White-backed Vulture Gyps africanus may nest on top of nest mass or in same tree; in southern Africa Bateleur Terathopius ecaudatus often nests near buffalo-weaver nests. Key References Birkhead, T. R. et al. (1993), Collias, N. E. and Collias, E. C. (1964), Kemp, A. and Kemp, M. (1974), Winterbottom, M. et al. (1999, 2001).

Genus Dinemellia Reichenbach

Endemic, monotypic. A large, social buffalo-weaver with predominantly white plumage, unique in family, and orange-red bend of wing, rump and undertail-coverts. A seed-eater, of semi-arid E Africa, constructing bulky nest of thorny twigs with single grass-lined chamber inside. Does not weave.

Plate 3

Dinemellia dinemelli (Ru ¨ ppell). White-headed Buffalo-Weaver. Alecto a` teˆte blanche.

(Opp. p. 26)

Textor dinemelli Ru ¨ ppell, 1845. Syst. Uebers. p. 72, pl. 30: Shoa, Ethiopia. Range and Status. Endemic resident. Sudan, common in SE from Torit eastwards, occasionally west to Juba, Mongalla. Ethiopia, central and southern parts of W highlands, SE highlands, W, S, NE regions and Rift Valley. Uganda, in NE, Karamoja to E Acholi. Kenya, northern lowlands, coast north of Ras Ngomeni, S-central and SE regions. Somalia, widespread and common, absent from NE. Tanzania, Mpanda, east of L. Victoria to Kilimanjaro, south to Shinyanga, Tabora, Ruaha River.

Dinemellia dinemelli

Description. D. d. dinemelli (Ru ¨ ppell): Sudan to Somalia and Kenya (except SE). ADULT Y: forehead, crown and nape white; mantle dark brown; rump orange-red; tail dark brown; chin, throat, chest, flanks and belly white; thighs brown; undertailcoverts orange-red. Wings dark brown with small orange epaulet, white band 20 wide across base of primaries, edges of wing coverts pale. Bill black (dark brown to horn in some birds; perhaps a seasonal change); eyes brown, with bare black skin around eye and between eye and bill; legs black. Leucistic bird in Kenya with pale orange undertail-coverts, larger white flash at base of primaries; mantle, wings and tail pale creamy brown (Lewis and Ogola 1989). Sexes alike. SIZE (11 YY, 12 XX): wing, Y 102–127 (112), X 103–128 (113); tail, Y 73–88 (764), X 72–82 (758); bill, Y 226–261 (245), X 224–253 (238); tarsus, Y 262– 305 (287), X 265–317 (290). WEIGHT: (Kenya) ad. Y (n ¼ 6) 596–780, X (n ¼ 2) 568, 855, (unsexed) 61, 65. IMMATURE: like ad. but tail-coverts pale orange (not orange-red). NESTLING: fledges with conspicuous gape wattle. D. d. boehmi (Reichenow): Tanzania to SE Kenya. Mantle, wings and tail black; intergrades with nominate race.

Field Characters. Length 18 cm. A large weaver of lowland thornbush, common and conspicuous. White head

and underparts are most striking features of perched birds; in flight a unique pattern of red, white and black. Unmistakable.

54

PLOCEIDAE

Zimbabwe, Sept–Apr, chiefly Jan–Feb; Mozambique, Sept–Nov; South Africa, Oct–Apr. INCUBATION: by X only, period 11 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only (Namibia), or (Kenya) by Y (1–2 feeds for every 24 by X); both adults once continued to feed young in nest mass cut down and resting on ground. Nestling period: 20–23 days. BREEDING SUCCESS/SURVIVAL: some nests deserted after attack by Wahlberg’s Eagles Aquila wahlbergi. Nests may be heavily infested with mites and insects; in Kenya, a

distinctive grey cockroach especially common. Of 16 eggs, 2 failed to hatch, 6 chicks fledged. White-backed Vulture Gyps africanus may nest on top of nest mass or in same tree; in southern Africa Bateleur Terathopius ecaudatus often nests near buffalo-weaver nests. Key References Birkhead, T. R. et al. (1993), Collias, N. E. and Collias, E. C. (1964), Kemp, A. and Kemp, M. (1974), Winterbottom, M. et al. (1999, 2001).

Genus Dinemellia Reichenbach

Endemic, monotypic. A large, social buffalo-weaver with predominantly white plumage, unique in family, and orange-red bend of wing, rump and undertail-coverts. A seed-eater, of semi-arid E Africa, constructing bulky nest of thorny twigs with single grass-lined chamber inside. Does not weave.

Plate 3

Dinemellia dinemelli (Ru ¨ ppell). White-headed Buffalo-Weaver. Alecto a` teˆte blanche.

(Opp. p. 26)

Textor dinemelli Ru ¨ ppell, 1845. Syst. Uebers. p. 72, pl. 30: Shoa, Ethiopia. Range and Status. Endemic resident. Sudan, common in SE from Torit eastwards, occasionally west to Juba, Mongalla. Ethiopia, central and southern parts of W highlands, SE highlands, W, S, NE regions and Rift Valley. Uganda, in NE, Karamoja to E Acholi. Kenya, northern lowlands, coast north of Ras Ngomeni, S-central and SE regions. Somalia, widespread and common, absent from NE. Tanzania, Mpanda, east of L. Victoria to Kilimanjaro, south to Shinyanga, Tabora, Ruaha River.

Dinemellia dinemelli

Description. D. d. dinemelli (Ru ¨ ppell): Sudan to Somalia and Kenya (except SE). ADULT Y: forehead, crown and nape white; mantle dark brown; rump orange-red; tail dark brown; chin, throat, chest, flanks and belly white; thighs brown; undertailcoverts orange-red. Wings dark brown with small orange epaulet, white band 20 wide across base of primaries, edges of wing coverts pale. Bill black (dark brown to horn in some birds; perhaps a seasonal change); eyes brown, with bare black skin around eye and between eye and bill; legs black. Leucistic bird in Kenya with pale orange undertail-coverts, larger white flash at base of primaries; mantle, wings and tail pale creamy brown (Lewis and Ogola 1989). Sexes alike. SIZE (11 YY, 12 XX): wing, Y 102–127 (112), X 103–128 (113); tail, Y 73–88 (764), X 72–82 (758); bill, Y 226–261 (245), X 224–253 (238); tarsus, Y 262– 305 (287), X 265–317 (290). WEIGHT: (Kenya) ad. Y (n ¼ 6) 596–780, X (n ¼ 2) 568, 855, (unsexed) 61, 65. IMMATURE: like ad. but tail-coverts pale orange (not orange-red). NESTLING: fledges with conspicuous gape wattle. D. d. boehmi (Reichenow): Tanzania to SE Kenya. Mantle, wings and tail black; intergrades with nominate race.

Field Characters. Length 18 cm. A large weaver of lowland thornbush, common and conspicuous. White head

and underparts are most striking features of perched birds; in flight a unique pattern of red, white and black. Unmistakable.

54

PLOCEIDAE

Zimbabwe, Sept–Apr, chiefly Jan–Feb; Mozambique, Sept–Nov; South Africa, Oct–Apr. INCUBATION: by X only, period 11 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only (Namibia), or (Kenya) by Y (1–2 feeds for every 24 by X); both adults once continued to feed young in nest mass cut down and resting on ground. Nestling period: 20–23 days. BREEDING SUCCESS/SURVIVAL: some nests deserted after attack by Wahlberg’s Eagles Aquila wahlbergi. Nests may be heavily infested with mites and insects; in Kenya, a

distinctive grey cockroach especially common. Of 16 eggs, 2 failed to hatch, 6 chicks fledged. White-backed Vulture Gyps africanus may nest on top of nest mass or in same tree; in southern Africa Bateleur Terathopius ecaudatus often nests near buffalo-weaver nests. Key References Birkhead, T. R. et al. (1993), Collias, N. E. and Collias, E. C. (1964), Kemp, A. and Kemp, M. (1974), Winterbottom, M. et al. (1999, 2001).

Genus Dinemellia Reichenbach

Endemic, monotypic. A large, social buffalo-weaver with predominantly white plumage, unique in family, and orange-red bend of wing, rump and undertail-coverts. A seed-eater, of semi-arid E Africa, constructing bulky nest of thorny twigs with single grass-lined chamber inside. Does not weave.

Plate 3

Dinemellia dinemelli (Ru ¨ ppell). White-headed Buffalo-Weaver. Alecto a` teˆte blanche.

(Opp. p. 26)

Textor dinemelli Ru ¨ ppell, 1845. Syst. Uebers. p. 72, pl. 30: Shoa, Ethiopia. Range and Status. Endemic resident. Sudan, common in SE from Torit eastwards, occasionally west to Juba, Mongalla. Ethiopia, central and southern parts of W highlands, SE highlands, W, S, NE regions and Rift Valley. Uganda, in NE, Karamoja to E Acholi. Kenya, northern lowlands, coast north of Ras Ngomeni, S-central and SE regions. Somalia, widespread and common, absent from NE. Tanzania, Mpanda, east of L. Victoria to Kilimanjaro, south to Shinyanga, Tabora, Ruaha River.

Dinemellia dinemelli

Description. D. d. dinemelli (Ru ¨ ppell): Sudan to Somalia and Kenya (except SE). ADULT Y: forehead, crown and nape white; mantle dark brown; rump orange-red; tail dark brown; chin, throat, chest, flanks and belly white; thighs brown; undertailcoverts orange-red. Wings dark brown with small orange epaulet, white band 20 wide across base of primaries, edges of wing coverts pale. Bill black (dark brown to horn in some birds; perhaps a seasonal change); eyes brown, with bare black skin around eye and between eye and bill; legs black. Leucistic bird in Kenya with pale orange undertail-coverts, larger white flash at base of primaries; mantle, wings and tail pale creamy brown (Lewis and Ogola 1989). Sexes alike. SIZE (11 YY, 12 XX): wing, Y 102–127 (112), X 103–128 (113); tail, Y 73–88 (764), X 72–82 (758); bill, Y 226–261 (245), X 224–253 (238); tarsus, Y 262– 305 (287), X 265–317 (290). WEIGHT: (Kenya) ad. Y (n ¼ 6) 596–780, X (n ¼ 2) 568, 855, (unsexed) 61, 65. IMMATURE: like ad. but tail-coverts pale orange (not orange-red). NESTLING: fledges with conspicuous gape wattle. D. d. boehmi (Reichenow): Tanzania to SE Kenya. Mantle, wings and tail black; intergrades with nominate race.

Field Characters. Length 18 cm. A large weaver of lowland thornbush, common and conspicuous. White head

and underparts are most striking features of perched birds; in flight a unique pattern of red, white and black. Unmistakable.

Plocepasser Voice. Tape-recorded (2, B, C, F, PARK, PAY, PEA, NOR). Wide vocabulary of screechy, rather starling-like notes. Typical is down-slurred whistle with reedy quality, ‘tsweew’ or ‘tsee-tsweew’. 2 birds calling together (possible duet? North 1958) ‘peeyee-tsuyu’, ‘peesu-tseeyu’, drawn-out ‘tsluweew’, shorter ‘pee-tsew’, abrupt ‘chu-chu’ and phrases ‘tsu-weely-weely-wair’ or ‘tsu-weely-weely-tsair-tsu’. Other calls are liquid ‘twooyoo’, ‘toowee’ and ‘tooweeyu’ and phrase ‘chalaweeya-chooli’, and buzzy ‘dzwo-zheeya’, ‘dzhweeya’ and longer 3-part ‘dzzer-wazhleeyaa’. General Habits. Inhabits dry bush and savanna thornveld (Tanzania: including acacia patches in miombo); generally below 1400 m. Rare in grassland and riverine vegetation. Occurs in parties of 3–6 birds; regularly joins mixedspecies flocks in open woodland (Kenya: Pomeroy and Tengecho 1982). Forages for insects on ground, often in association with sparrow-weavers and starlings, e.g. Superb Starling Lamprotornis superbus and White-crowned Starling Spreo albicapillus in Somalia. Adept at running. In Tsavo present all year, most abundant in June–Aug. Lone Y calls from treetops. Food. Insects, also seeds and fruit. Beetles and caterpillars in stomachs. Breeding Habits. Monogamous. Captive Y flutters outstretched wings and chatters when X returns after brief separation; X responds with weaker wing-fluttering. Y occasionally feeds X. Possible cooperative breeder; 3 adults fed young in captivity (Wilkinson and McLeod 1994). NEST: large ovoid structure 05 m long, slightly flattened (A), outer shell built of thorny sticks, with short entrance tube at level of nest chamber opening downwards, inside this is dome-shaped nest of dry grass stems. Lined with grass, woolly leaves and feathers; saddled across branches, suspended on long branch or attached to underside, often surrounded with extra thorny twigs. Built by Y and X; apparently inner herbaceous shell is built first, then top and sides are covered with thorny twigs. Single nest in tree, or may be grouped, usually 2–4 m above ground. Captive birds built 3 nests; several nests may belong to single pair. Sometimes uses old nest of White-browed Sparrow-weaver Plocepasser mahali (Granvik 1934). Some

nest trees in Kenya held colonies of biting ants (Collias and Collias 1964). EGGS: 3–4. Greyish white or pale blue, thickly blotched and marbled with olive-brown and grey-brown, often forming cap at thick end. SIZE: (Kenya, n ¼ 12) 225– 265  171–190 (246  180); (Somalia, n ¼ 10) 230– 260  180–190 (250  185). LAYING DATES: Sudan, Aug–Sept, Dec; Ethiopia, Apr– Aug; Somalia, Feb–June, Sept; Uganda, May; Kenya, Feb– Mar, May, July, Nov; Tanzania, Dec (breeding condition June; nest-building Nov). INCUBATION: no information. DEVELOPMENT AND CARE OF YOUNG: in captivity, fed by both Y and X; intolerant of other birds at the nest. Nestling period >14 days. Fledgling feeds itself after 4 weeks. BREEDING SUCCESS/SURVIVAL: nests may be taken over by Pygmy Falcon Polihierax semitorquatus. Key Reference Hawkins, R. W. (1958).

Genus Plocepasser Smith

Medium-sized sparrow-like birds, with no sexual dimorphism nor seasonal change in plumage. Upperparts brownish, underparts white, prominent superciliary stripe, eyes brown. Complex, musical songs. Feed predominantly on ground, taking seeds and insects. Always in dry savanna country, in small flocks which form cooperative breeding units. Conspicuous nests constructed of straight grass stalks, not woven; in thorn trees; nests with a second entrance used for roosting, and form a permanent base for the group. Endemic, 4 species.

55

Plocepasser Voice. Tape-recorded (2, B, C, F, PARK, PAY, PEA, NOR). Wide vocabulary of screechy, rather starling-like notes. Typical is down-slurred whistle with reedy quality, ‘tsweew’ or ‘tsee-tsweew’. 2 birds calling together (possible duet? North 1958) ‘peeyee-tsuyu’, ‘peesu-tseeyu’, drawn-out ‘tsluweew’, shorter ‘pee-tsew’, abrupt ‘chu-chu’ and phrases ‘tsu-weely-weely-wair’ or ‘tsu-weely-weely-tsair-tsu’. Other calls are liquid ‘twooyoo’, ‘toowee’ and ‘tooweeyu’ and phrase ‘chalaweeya-chooli’, and buzzy ‘dzwo-zheeya’, ‘dzhweeya’ and longer 3-part ‘dzzer-wazhleeyaa’. General Habits. Inhabits dry bush and savanna thornveld (Tanzania: including acacia patches in miombo); generally below 1400 m. Rare in grassland and riverine vegetation. Occurs in parties of 3–6 birds; regularly joins mixedspecies flocks in open woodland (Kenya: Pomeroy and Tengecho 1982). Forages for insects on ground, often in association with sparrow-weavers and starlings, e.g. Superb Starling Lamprotornis superbus and White-crowned Starling Spreo albicapillus in Somalia. Adept at running. In Tsavo present all year, most abundant in June–Aug. Lone Y calls from treetops. Food. Insects, also seeds and fruit. Beetles and caterpillars in stomachs. Breeding Habits. Monogamous. Captive Y flutters outstretched wings and chatters when X returns after brief separation; X responds with weaker wing-fluttering. Y occasionally feeds X. Possible cooperative breeder; 3 adults fed young in captivity (Wilkinson and McLeod 1994). NEST: large ovoid structure 05 m long, slightly flattened (A), outer shell built of thorny sticks, with short entrance tube at level of nest chamber opening downwards, inside this is dome-shaped nest of dry grass stems. Lined with grass, woolly leaves and feathers; saddled across branches, suspended on long branch or attached to underside, often surrounded with extra thorny twigs. Built by Y and X; apparently inner herbaceous shell is built first, then top and sides are covered with thorny twigs. Single nest in tree, or may be grouped, usually 2–4 m above ground. Captive birds built 3 nests; several nests may belong to single pair. Sometimes uses old nest of White-browed Sparrow-weaver Plocepasser mahali (Granvik 1934). Some

nest trees in Kenya held colonies of biting ants (Collias and Collias 1964). EGGS: 3–4. Greyish white or pale blue, thickly blotched and marbled with olive-brown and grey-brown, often forming cap at thick end. SIZE: (Kenya, n ¼ 12) 225– 265  171–190 (246  180); (Somalia, n ¼ 10) 230– 260  180–190 (250  185). LAYING DATES: Sudan, Aug–Sept, Dec; Ethiopia, Apr– Aug; Somalia, Feb–June, Sept; Uganda, May; Kenya, Feb– Mar, May, July, Nov; Tanzania, Dec (breeding condition June; nest-building Nov). INCUBATION: no information. DEVELOPMENT AND CARE OF YOUNG: in captivity, fed by both Y and X; intolerant of other birds at the nest. Nestling period >14 days. Fledgling feeds itself after 4 weeks. BREEDING SUCCESS/SURVIVAL: nests may be taken over by Pygmy Falcon Polihierax semitorquatus. Key Reference Hawkins, R. W. (1958).

Genus Plocepasser Smith

Medium-sized sparrow-like birds, with no sexual dimorphism nor seasonal change in plumage. Upperparts brownish, underparts white, prominent superciliary stripe, eyes brown. Complex, musical songs. Feed predominantly on ground, taking seeds and insects. Always in dry savanna country, in small flocks which form cooperative breeding units. Conspicuous nests constructed of straight grass stalks, not woven; in thorn trees; nests with a second entrance used for roosting, and form a permanent base for the group. Endemic, 4 species.

55

56

PLOCEIDAE

Plate 3

Plocepasser mahali A. Smith. White-browed Sparrow-Weaver. Mahali a` sourcils blancs.

(Opp. p. 26)

Plocepasser mahali A. Smith, 1836. Rep. Exped. Centr. Afr., p. 51; Orange R., South Africa. Range and Status. Endemic resident. Sudan, southeast from Juba, Lado eastwards. Ethiopia, western highlands, Rift Valley, NE, S and SE regions. Somalia, local south of 4
N. Uganda, Bunyoro, Lango, N Teso, Karamoja, old record from Bakivungu. Kenya, north shores of Winam (Kavirondo) Gulf, L. Magadi, Amboseli, Tsavo east to Wajir and Garissa, L. Turkana basin. Tanzania, no records in N prior to 1980 (Britton 1980), local in 1980s, now well established, but real gap between Wami R. and Liwale; widespread in central highlands, including Mikumi and Selous Res., outlier on Singida highlands. Angola, from Cunene and S and central Huı´la to Namibe, north to Benguela, also in SE Cuando Cubango. Zambia, Luangwa Valley, middle Zambezi Valley, Southern Prov., SE ^ Shire Valley north to Monkey Western Province. Malawi, ^ Mozambique, Bay and Salima on SW shores of L. Malawi. Tete to Feira in Zambezi Valley. Namibia, throughout interior, largely absent from coastal desert. Botswana, throughout where sufficient trees are present; 30 colonies in one 70 km transect (P. J. Jones pers. comm.). Zimbabwe, throughout, west of 31
E and along Zambezi Valley, absent from E highlands and plateau. South Africa, throughout Northern Cape, Free State, Northwest Province, very local in KwaZulu-Natal, largely absent from E lowveld areas, and isolated population in dry interior of Eastern Cape. Common to abundant. Description. P. m. mahali A. Smith (includes ‘stentor’, ‘terricolor’): S Namibia, S Botswana, S Zimbabwe, South Africa. ADULT Y: forehead and crown chocolate-brown; broad white superciliary stripe; cheeks and ear-coverts brown; nape and mantle brown, paler than crown; rump white; tail brown, with paler tips to feathers; chin white, with dark blackish brown malar stripes; throat, breast, belly, thighs and undertail-coverts white; flanks white with some buffy wash; wing brown, with broad pale tips to greater and lesser coverts, narrow pale edges to remiges. Bill black (in some populations apparently in YY only, brown in XX; in others, black bills in both sexes); eyes rich chestnut-brown; legs brown. Sexes alike. SIZE (14 YY 11 XX): wing, Y 90–112 (101), X 91–105 (100); tail, Y 61–70 (666), X 63–67 (647); bill, Y 188– 226 (204), X 185–216 (201); tarsus, Y 230–289 (259), X 249– 280 (260). WEIGHT: Y (Namibia, n ¼ 7) 36–42 (399), (South Africa, n ¼ 13) 456–520 (495), X (Namibia, n ¼ 4) 37–42 (405), (Botswana, n ¼ 5) 380–409 (392), (South Africa, n ¼ 13) 456– 518 (489); unsexed (Botswana) 342, 439, 448 (P. J. Jones pers. comm.), (Zimbabwe, Hwange) at Main Camp (n ¼ 27) 350–467 (432), Robins Camp (n ¼ 65) 327–470 (389) but no differences at same site at different times of day (C. J. Vernon pers. comm.); (South Africa, n ¼ 811) 31–59 (440); juv. (South Africa, n ¼ 40) 31–48 (400). Within a breeding group, breeding Y heaviest, breeding X next heaviest (Earle´ 1983b). IMMATURE: bill paler than adult. NESTLING: not described. P. m. melanorhynchus Bonaparte: Sudan to Kenya. Upperparts earthen brown; black marks at side of breast continuous with black of malar region; underparts otherwise plain white. Eyes red-brown in adult, dark grey in nestling. WEIGHT: unsexed (Kenya, n ¼ 35) 310–430 (391). ^ P. m. pectoralis (Peters): Tanzania, Zambia, Malawi, Mozambique, NE Zimbabwe, N Botswana. Back reddish brown; breast feathers with triangular dark centres; bill black.

Plocepasser mahali

?

P. m. ansorgei Hartert: Angola to N Namibia. Broader white tips to tail feathers. TAXONOMIC NOTE: several additional races have been described but they intergrade, and degree of interchange between local populations is not known.

Field Characters. Length 16–18 cm. A large, noisy brown and white weaver, common and familiar around drycountry campgrounds. On ground shows broad white stripe from above eye backwards, and 2 prominent white wing-bars and wing edgings; in flight, large white patch on rump and uppertail-coverts conspicuous. Voice. Tape-recorded (22, 36, 38, 39, 58, 88, 99, B, C, F, LEM, PAY). Loud, cheerful, bustling song complex and high-pitched, with chattering and squealing notes; no two phrases exactly the same: ‘chop-cheelu-cheeyu-cheeluchop’; ‘chop-cheeyu-cheeyu-cheeyu-cheeyu’; ‘woyo-cheechu-weelu-chochiro-chiyo-chiyo-jarp-jurr’; ‘cheer, cheechee, chachacha-jeer’; in chattery versions, rattle may be all on 1 pitch, or shrill notes descending into dry chatter, ‘chee-chee-chee-chee-chachachachacha’, or ascending, accelerating ‘chochochocho-chichichichichi . . . ’. For further variations see Zimmerman et al. (1996). Territorial song used by group members in boundary display also complex, with wide frequency sweeps and low trills; much individual variation. Harsh low-pitched call when several birds perch close together; short ‘chirp’ in flight. Brief ‘chuk’ in alarm, also used as vigilance call by perched birds. In Kenya, alarm calls given to Pearl-spotted Owlet Glaucidium perlatum, Fan-tailed Raven Corvus rhipidurus, vervet monkeys and people (Collias and Collias 1978c). A

Plocepasser mahali high-pitched ‘tweet’ given by copulating Y and by begging chicks (Ferguson 1988a). General Habits. Inhabits mopane and acacia savanna mainly below 1400 m (but ranges up to 1900 m) and with 500,000 birds (Kemp et al. 2001). Density of 2.5 per ha on transect counts in N and E Botswana and 9 per ha in Kutse Game Res. (Herremans 1997); commonest of all birds near Gaborone, with up to 22 birds per km in transects (Tyler 2001a). Description. ADULT Y: forehead and crown feathers arrowshaped, black with white margins, lores black, narrow eye-ring white; cheeks and ear-coverts dull grey-brown; nape, mantle and rump dull grey-brown; tail dark brown, with white fringes all around; chin black, with black malar streak extending down onto throat, bordered above by white streak; throat white; breast white, with some light brown wash; flanks, belly, thighs and undertail-coverts whitish; wings brown, coverts and tertials very dark blackish-brown with broad white margins. Bill pink, varying from flesh-coloured to rosy (apparently unrelated to breeding condition); eyes reddish brown; legs pale horn-brown. Sexes alike. SIZE (12 YY, 14 XX): wing, Y 55–60 (577), X 54–59 (566); tail, Y 37–42 (400), X 36–42 (399); bill, Y 105–113 (109), X 105–120 (112); tarsus, Y 147–165 (157), X 146–166 (159). WEIGHT: (South Africa) 2 YY 112, 113; unsexed (n ¼ 66) 104–137 (123); juv. (n ¼ 52) 94–136 (116) (Skead 1974). IMMATURE: like ad., but crown and forehead dull grey-brown with no scaling, malar stripes lacking; bill horn coloured.

65

Sporopipes squamifrons

NESTLING:

no information.

Field Characters. Length 11 cm. Very small, rather plump and short-tailed; pale brown and white with distinctive

Plate 3 (Opp. p. 26)

66

PLOCEIDAE

head pattern – bill pink, forehead speckled black and white, black lores, brown cheeks separated by white streak from prominent black moustachial stripe; wings and tail black with white edgings conspicuous in flight; white eye-ring visible at close range. Immature duller, without speckles on forehead, wings brown and buff, bill horn-coloured. Voice. Tape-recorded (88, 99, B, F, CHA, LUT, PAY, STJ). Song a repeated ‘chisip-chisip-chap-chap’ or ‘chisipchap chisip-chap’, with variations; has been likened to Cape Sparrow Passer melanurus but is much higher-pitched. Calls include high thin ‘seep’ or ‘tsee-tsee’, irregular grating chatter, ‘jeep, jee, jer . . . ’ or higher buzzy ‘tzip, jyip, jip zyip . . . ’; wispy rolling ‘sirrip’ given frequently. Contact call in flight a pleasant ‘ching-ching-ching’ (Maclean 1993); when disturbed flies off with chattering sound (Newman 1989). X solicits copulation with high ‘zizi-zi-’. General Habits. Inhabits dry Acacia savanna, arid scrub and bush along dry watercourses, shrubs near houses. In pairs or flocks of up to 20 birds; flocks regularly with waxbills, particular Violet-eared Waxbill Granatina granatina and Blue Waxbill Uraeginthus angolensis; sometimes displaces the latter (Skead 1975). Pair allopreens, primarily around the head. Sleeps communally in nest, all year, with up to 12 birds using 1 nest. May roost on branch, 3–4 birds clustered together. Dust-bathes. Feeds largely on ground; moves mainly by hopping, seldom walks except for very short distance. Wild-caught birds survived 62 days without drinking water; increased in weight when able to drink again (Cade 1965). Post-nuptial moult in Jan (Botswana). No regular movements, but locally nomadic under unfavourable conditions. Birds from SE Botswana showed no clear moulting season, and primary moult lasts at least 200 days in individual birds. Some young birds apparently have a partial moult, which may be irregular in sequence; suspended wing-moult also occurs (Tyler 2001b). Food. Small seeds; insects fed to young, in captivity; all animal food collected on ground. Never drinks (semidesert areas of Botswana: Irwin 1956). Breeding Habits. Solitary nester; monogamous; territorial. Courtship and breeding behaviour barely known; once Y flew to X, landed on her back with quivering wings; X crouched and quivered wings also. Y leant back to make cloacal contact, but did not grasp nape feathers. Soliciting X quivered both wings and tail, gave high-pitched call (Michaelis 1970). Captive X paired with Y Speckle-fronted Weaver S. frontalis and produced young (Kunkel 1966). NEST: untidy, rugby-ball shaped mass (A) of stiff grass stems and fruiting heads of grasses Eragrostis, Urochloa, Sporobolus, Aristida, Andropogon and Digitaria; lined with plant down, spikelets of Rhynchelytrum and feathers. Sited 09–42 m above ground (av. 17 m, n ¼ 50, South Africa); in large trees up to 6 m. Often placed in Acacia mellifera (Botswana), A. karoo (South Africa) and Ziziphus (Zimbabwe). Nest-building starts with hollow cup, above

which a dome is then formed, but roof is thin and transparent in sleeping nests. May roof over old cup nest of shrike or flycatcher. Captive birds use old nests of other species, as well as building their own; Y brings material, X built nest. EGGS: 3–5 (av. 41, n ¼ 63, South Africa); up to 7 in 1 nest in Zimbabwe. Greenish, bluish or pale cream, almost covered with dense freckling or mottling in ash-grey and light brown to olive-brown. SIZE: (South Africa, n ¼ 106) 139–180  103–127 (157  113). LAYING DATES: Botswana, mostly Dec–Apr, may breed all year with records from all months except June, Aug (Skinner 1995); Zimbabwe, all year, with peak Jan–Mar; South Africa, all year, peak months Jan–June. INCUBATION: by X only. Typically for c. 19 min on eggs and 5 min off. Period 10–12 days. DEVELOPMENT AND CARE OF YOUNG: fed by both Y and X. Nestling period: 14–18 days. Young continue to sleep in nest for several weeks. BREEDING SUCCESS/SURVIVAL: 4 YY and 4 XX died during cold spell in Bulawayo, Zimbabwe, in 1968 (Steyn and Brooke 1970). Fiscal Shrike Lanius collaris attacked nest after birds had entered to roost, but failed to capture any (Collias and Collias 1964). Preyed on by Barn Owl Tyto alba in Gauteng, South Africa (Dean 1973).

Key References Michaelis, H. J. (1970), Tyler, S. J. (2001b), Vincent, A. W. (1949).

Pseudonigrita arnaudi

67

Genus Pseudonigrita Reichenow

Small, social weavers of semi-arid E Africa. Grey or black cap; no red or yellow in plumage. Cooperative breeders; single nests of straight grass stems, not woven, clustered in groups in trees. Breeding nest used by single pair, with single entrance; group roosts in nest with 2 entrances. Granivorous. Endemic, 2 species in a superspecies.

Pseudonigrita arnaudi superspecies

1

2

2

1

1 1 P. arnaudi 2 P. cabanisi

1

Pseudonigrita arnaudi (Bonaparte). Grey-headed Social Weaver. Re´publicain d’Arnaud.

Plate 5

Nigrita arnaudi Bonaparte, 1850. Consp. Gen. Av., I, p. 444; White Nile, Sudan.

(Opp. p. 74)

Forms a superspecies with P. cabinisi. Range and Status. Endemic resident. Sudan, S Darfur, and local in SE from Bor to Kobowen Swamp. Ethiopia, uncommon in extreme S. Somalia, 2 old records at Afmadow (1962, 1964). Uganda, Karamoja, Acholi. Kenya, locally common in Karasuk Hills, Laikipia to L. Magadi, Kedu, Tsavo. Tanzania, Iringa and Dodoma area, Serengeti and Siloti R. to Asambiro, Shinyanga and Tabora, west of Kilimanjaro from L. Natron and Longido to Mt Meru; colony in Dar es Salaam (Baker and Howell 1994). Description. P. a. arnaudi (Bonaparte): Sudan to NE Tanzania. ADULT Y: forehead, crown to nape pale grey, ending in sharp line on nape, and extending down to line level with mid-point of eye; lores black, cheeks and ear-coverts sandy brown with ring of white feathers around eye; mantle and rump sandy brown; tail dark brown, outer feathers with dull white tips, central 2 feathers with entire inner web also dull white; underparts uniform sandy

brown; wings dark brown, pale outer margin on secondaries, tertials with entire outer web sandy brown, coverts sandy brown except for second row of primary wing-coverts, which are darker brown with grey tips, forming small grey epaulet. Bill black; eyes red-brown to brown; legs brown. Sexes alike. SIZE (12 YY, 12 XX): wing, Y 63–70 (665), X 63–69 (660); tail, Y 32–40 (358), X 32–40 (353); bill, Y 131–150 (142), X 140–155 (146); tarsus, Y 170–190 (179), X 170–187 (176). WEIGHT: (Kenya) Y (n ¼ 4) 19–22 (203), X (n ¼ 4) 15–22 (195); unsexed (n ¼ 54) 162–259 (200). IMMATURE: lacks grey cap and ring of white feathers around eye, generally paler and buffier than adult, with dark brown patch on epaulet, buffy margins to tail feathers. Bill light brown. NESTLING: not described. P. a. dorsalis (Reichenow): W and central Tanzania. Mantle grey.

Field Characters. Length 11–12 cm. Small and plump with short pale-tipped tail, drab grey-brown except for

Pseudonigrita arnaudi

67

Genus Pseudonigrita Reichenow

Small, social weavers of semi-arid E Africa. Grey or black cap; no red or yellow in plumage. Cooperative breeders; single nests of straight grass stems, not woven, clustered in groups in trees. Breeding nest used by single pair, with single entrance; group roosts in nest with 2 entrances. Granivorous. Endemic, 2 species in a superspecies.

Pseudonigrita arnaudi superspecies

1

2

2

1

1 1 P. arnaudi 2 P. cabanisi

1

Pseudonigrita arnaudi (Bonaparte). Grey-headed Social Weaver. Re´publicain d’Arnaud.

Plate 5

Nigrita arnaudi Bonaparte, 1850. Consp. Gen. Av., I, p. 444; White Nile, Sudan.

(Opp. p. 74)

Forms a superspecies with P. cabinisi. Range and Status. Endemic resident. Sudan, S Darfur, and local in SE from Bor to Kobowen Swamp. Ethiopia, uncommon in extreme S. Somalia, 2 old records at Afmadow (1962, 1964). Uganda, Karamoja, Acholi. Kenya, locally common in Karasuk Hills, Laikipia to L. Magadi, Kedu, Tsavo. Tanzania, Iringa and Dodoma area, Serengeti and Siloti R. to Asambiro, Shinyanga and Tabora, west of Kilimanjaro from L. Natron and Longido to Mt Meru; colony in Dar es Salaam (Baker and Howell 1994). Description. P. a. arnaudi (Bonaparte): Sudan to NE Tanzania. ADULT Y: forehead, crown to nape pale grey, ending in sharp line on nape, and extending down to line level with mid-point of eye; lores black, cheeks and ear-coverts sandy brown with ring of white feathers around eye; mantle and rump sandy brown; tail dark brown, outer feathers with dull white tips, central 2 feathers with entire inner web also dull white; underparts uniform sandy

brown; wings dark brown, pale outer margin on secondaries, tertials with entire outer web sandy brown, coverts sandy brown except for second row of primary wing-coverts, which are darker brown with grey tips, forming small grey epaulet. Bill black; eyes red-brown to brown; legs brown. Sexes alike. SIZE (12 YY, 12 XX): wing, Y 63–70 (665), X 63–69 (660); tail, Y 32–40 (358), X 32–40 (353); bill, Y 131–150 (142), X 140–155 (146); tarsus, Y 170–190 (179), X 170–187 (176). WEIGHT: (Kenya) Y (n ¼ 4) 19–22 (203), X (n ¼ 4) 15–22 (195); unsexed (n ¼ 54) 162–259 (200). IMMATURE: lacks grey cap and ring of white feathers around eye, generally paler and buffier than adult, with dark brown patch on epaulet, buffy margins to tail feathers. Bill light brown. NESTLING: not described. P. a. dorsalis (Reichenow): W and central Tanzania. Mantle grey.

Field Characters. Length 11–12 cm. Small and plump with short pale-tipped tail, drab grey-brown except for

68

PLOCEIDAE

Pseudonigrita arnaudi

contrasting pale grey cap from forehead to nape, white eyering. Juv. almost featureless, with dark brown epaulet, darker cheeks; not unlike young Black-capped Social Weaver P. cabanisi but underparts brown, not white; bill smaller, tail shorter, with buffy tip. Voice. Tape-recorded (B, C, GREG, McVIC, MOY, NOR, PAY). Sonagrams in Collias (1984). Common call, especially when going to roost, a short rolling trill, ‘trrree’, ‘trrrreew’ or ‘teeerr’ with plaintive quality recalling whiteeye Zosterops. Other calls include buzzy ‘peezu-peezu’, ‘pzeeww’ or ‘zzzz’, hard ‘zeet-zeet-zeet’, rattling ‘pitty’ or ‘pittu’, ‘chreet-chewt’ and lower ‘tyoop’, also a ‘faint whistle’ (Benson 1947). Contact and flight call ‘sreep’ (contact calls used by partners at nest relief); alarm, sharp ‘cheep’; in territorial display a loud ‘chee-chee-chee’. Audible differences between voices of Kenyan and Ethiopian birds (Benson 1948). Young birds give shrill repetitive begging calls. For further renditions see Zimmerman et al. (1996). General Habits. Inhabits bush and acacia woodland, generally below 1400 m but may range up to 1900 m. Usually in wetter areas than P. cabanisi, although the 2 can occur together. Feeds predominantly on ground. Resident at nesting colonies throughout the year, roosting in nests which retain 2 entrances; also roosts in nests of other weavers. Family group of up to 7 birds sleeps together in same nest until next breeding attempt, when young from previous brood are driven out. There appears to be a consistent dominance hierarchy within the group. Food. Grass seeds and insects; grasshoppers, beetles, termites and caterpillars in stomach contents. Young fed on soft green grass seeds and insects. Birds drink regularly.

Breeding Habits. Colonial, with up to 157 nests, some in fused masses, in a single tree; monogamous, co-operative breeder. Colony consists of several groups of 2–10 birds, which may have 2–8 nests, defended against members of other groups, but most vigorously against strangers from other colonies. Communal mobbing of predators ignores territory boundaries within the colony. Each family group has cluster of nests in 1 or more trees; sites are shifted unpredictably. Sub-ad. helpers may assist in feeding young of parents or other close relatives. Timing of breeding relates to rainfall; at some colonies, eggs laid in all months of the year. Nest area is defended by head-forward threat: aggressor raises feathers of nape and mantle, gapes widely, and may also raise tail and quiver wings. Captive pair tolerates neither neighbours nor neighbours’ offspring at their nest. Y performs courtship display at nest entrance, sometimes accompanied by soft, rattling trill: he crouches across branch, wings drooped, spread and quivered, and persistently follows X with body erect, tail down. On ground, Y raises bill and tail, partly spreads and quivers wings; sometimes holds straw or grass seedhead in bill during courtship. X solicits copulation with raised tail and quivering wings. NEST: large, compact grass nest firmly attached to thin branches, usually in tall spindly saplings or ant-gall acacias; symmetrical structure, with 2 openings below when nest used for roosting (A), 1 closed when eggs laid. Lined by both sexes with feathers. Constructed of straight grass stems, laid like thatch around a central cavity (B). Birds constantly add to and modify nest structure, may pull out stems and replace them elsewhere. 6–20 nests may be more or less fused together into a mass; such masses are common in larger trees. 1 colony, in Sudan, was in Acacia

Pseudonigrita arnaudi thorny acacias such as Acacia tortilis, A. mellifera, A. senegala, but largest nest masses are generally in ant-gall acacias A. drepanolobium. Generally the larger the tree, the more nests, both within and between tree species. Most commonly used trees, Kenya: A. tortilis (n ¼ 73), A. thomasi (44), A. senegal (31), Balanites aegyptiaca (19), A. xanthophloea (13) and A. drepanolobium (12) (Collias and Collias 1977). EGGS: 3–4, greenish or bluish white, with fine olive or black specks, thickly blotched and marbled with olivebrown and ash-grey; often white, with sparse markings concentrated at thick end. Considerable variation in egg patterning, but each X lays eggs of 1 type only; only 1 X contributes eggs to a nest. SIZE: (Kenya, n ¼ 3) 173– 185  118–13, (n ¼ 6) av. 19  14. LAYING DATES: Sudan, Aug–Dec; Ethiopia, Mar–Apr (possible breeding June); E Africa, Region A, Aug (1 clutch), Dec (10 clutches), Region C, Dec–Feb, Apr–June, Region D, Dec–July (121 clutches, 21% in Mar–Apr, 67% in May, 7% in June, July and Dec); mainly in rains, sometimes before rains in Region D and after them in Region C. INCUBATION: by both Y and X. Eggs left uncovered c. 23% of time. In captivity, X did 70% of incubation (Hemmer 1976). Both Y and X roost in nest at night throughout incubation period. Other birds may sleep in nests, but do not incubate. Period: 14 days; in captivity, 13 days. DEVELOPMENT AND CARE OF YOUNG: young make opening in wall of nest chamber, and parents arrive to feed them using both opening and proper nest entrance (Granvik 1934). Young fed by Y and X and by helpers, mainly YY, which might have active nests of their own. At a Kenyan colony, 25–30% of pairs had helpers. Helped pairs raised more young than unhelped pairs; helpers contributing most at time of greatest food demand by chicks. Nestling period: 20 days. Fledged young fed by adults for 2–3 weeks after leaving nest. BREEDING SUCCESS/SURVIVAL: colony in Dar es Salaam disrupted by House Crows Corvus splendens (Baker and Howell 1994). Snakes take eggs. Some nests apparently damaged by Gabar Goshawk Melierax gabar. Adult weaver caught by African Harrier Hawk Polyboroides typus on leaving nest. Colony members mobbed a stuffed owl and a rubber snake placed in tree, but not on ground (Collias and Collias 1980). At 1 Kenyan colony, of 56 birds ringed, only 6 present after 25 years, suggesting annual survival of 38% (Collias and Collias 1981); at another site, adults had annual survival rate of 81%, but only 20% of juvs were present after 1 year. XX apparently disperse before YY (Bennun 1994a). Old nests used by Cut-throat Finches Amadina fasciata. Chestnut Sparrows Passer eminibey actively usurp newly-built nests. Captive birds defended nest against Chestnut Sparrows but tolerated other species in aviary.

drepanolobium tree with Crematogaster ants inhabiting tree galls (C, a single nest); when birds left the nest, ants did not react, whereas any other disturbance brought the ants swarming out (Moreau 1943). Colonies in Kenya often in

Key References Bartsch, C. (1991), Bennun, L. (1992, 1994a, b), Collias, N. E. (1984), Collias, N. E. and Collias E. C. (1977, 1978d, 1980), Payne, R. B. (1969).

69

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Plate 5 (Opp. p. 74)

PLOCEIDAE

Pseudonigrita cabanisi (Fischer and Reichenow). Black-capped Social Weaver. Re´publicain de Cabanis. Nigrita cabanisi Fischer and Reichenow, 1884. J. Orn., 1884, p. 54; Pare´ Mts, Tanzania. Forms a superspecies with P. arnaudi. Range and Status. Endemic resident. Ethiopia, uncommon in Arero Forest, Dawa-Wachile, Yabello Sanctuary and Hills and Rift Valley. Somalia, previously known only from 2 records in 1901 and 1 in 1922 on Kenyan border at Damassa, Wonte, Geel Waaq; recently recorded in Nugaal valley, Buulobarde, Boja swamp, Laag Dheere (Fishpool and Evans 2001). Kenya, locally common in Mandra, Wajir, Wemba and Borojoi south to Nemanga and Tsavo, nonbreeding visitor to Tsavo East Nat. Park. Tanzania, Arusha and lowlands near Kilimanjaro, south to Aruha Naherera.

Pseudonigrita cabanisi

Description. ADULT Y: forehead and crown black, extending to just below eye and ending in sharp line on nape; nape, mantle and rump light brown; tail dark brown; chin, throat and breast white; belly white, with short central line of c. 25 black feathers; flanks light brown with line of black bordering white belly, inner webs of feathers light brown, outer webs black; thighs and undertailcoverts white; wings light brown, with some black speckles on leading edge. Bill ivory white; eyes red; legs brown. Sexes alike. SIZE (11 YY, 10 XX): wing, Y 63–71 (675), X 64–68 (653); tail, Y 43–49 (469), X 43–49 (454); bill, Y 148–170 (160), X 143–161 (155); tarsus, Y 160–182 (170), X 157–178 (166). WEIGHT: (Kenya) Y (n ¼ 4) 215–240 (230), X (n ¼ 4) 180–190 (185); Tanzania, 1 Y 225. IMMATURE: head brown, not black; lacks black borders to flanks and black central streak on belly. Bill horn-brown; eyes brown. NESTLING: not described.

Field Characters. Length 13 cm. Light brown above, with black cap from forehead to nape and black tail; white below, with lines of black feathers along flanks and up centre of lower belly; bill white, eyes red. Juv. differs from juv. Grey-capped Social Weaver P. arnaudi in whitish underparts, dark brown crown, longer tail, dark without pale tip. Voice. Tape-recorded (B, KEI, McVIC, TUCK). Squeaky, chirping chatter from birds in breeding colony, notes with hard, dry, grating quality, ‘beek’, ‘gerp’ and buzzy ‘beezz’ and ‘bozzz’; rendered ‘sk’peee chwee-cher skiieer chir-chir squirrrrrr chirr-chrii-chirr’ (Zimmerman et al. 1996). General Habits. Very localized in dry thorn-bush; generally below 1300 m. At some sites occurs alongside P. arnaudi, but they are usually allopatric. Always in groups, birds often perch close together. Forages on ground. 2 birds allopreened briefly (Collias 1984). Nonbreeding flocks are nomadic. Food. Seeds, and a few insects, including grasshoppers. Breeding Habits. Colonial, with 7–61 nests in a colony; monogamous. Colony in Tanzania in Delonix tree, the largest tree in a wide radius. Bird in apparent courtship display sings and flaps wings; raises and vibrates tail

vertically (Collias 1984). Pairs often perch together in nesting tree. NEST: bulky, conical structure, built entirely of thin, dry grass stems, unlined; protruding ends of grass make it prickly (A); sometimes c. 5 nests combined into single compound one (B, recalling huge compound nest of Sociable Weaver Philetairus socius). 2 entrances when used for roosting, 1 closed for breeding (C). Frequently placed near end of slender, pendent branch, in colonies on isolated acacia trees (D); nests may merge into compound

Pseudonigrita cabanisi

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structure. At one site in Kenya, 5 nests in Acacia mellifera, 1 in A. tortilis; most nests on W side of trees; also nests in tree euphorbias. Birds active at nests, adding material, throughout year. One nest weighed 211 g, contained an estimated 9300 grass stems (Moreau and Moreau 1939). Pad built alongside twig on which bird perches, then extended and arched above twig. Arch is extended on both sides of twig so that tubular nest straddles it and sags at each end, with entrance on either side of supporting twig. 2 birds may work at nest at once.

EGGS: 2–4. White or pinkish, with brown and violet markings. SIZE: 19  14. LAYING DATES: Ethiopia, Feb–May; Kenya, Jan–May, Aug–Sept, Dec; Tanzania, Apr–June, possibly Nov. No further information.

Key References Collias, N. E. and Collias, E. C. (1964), Moreau, R. E. and Moreau, W. M. (1939).

Genus Philetairus Smith

Endemic, monotypic. Small, short-tailed, monomorphic, dull-plumaged weaver of arid SW Africa, where trees or other structures available to support the huge, domed, communal nest mass. Nests constructed of straight grass stems, with numerous (often >100) individual chambers under a common roof. Granivorous.

Plate 5

Philetairus socius (Latham). Sociable Weaver. Re ´publicain social.

(Opp. p. 74)

Loxia socia Latham, 1790. Ind. Orn. i, p. 381; Great Namaqualand, South Africa. Range and Status. Endemic resident. Namibia, Etosha and Ovambo region, separated from population on central plateau, south to Orange R. but absent from coastal desert. Botswana, abundant in SW sector, sparse and localized in SE in Pitsane area; absent from fire-prone central and N Kalahari. South Africa, abundant in Northern Cape from 18
E eastwards, south to 29
300 S and, near Vanwyksvlei, 31
S; absent from Ghaap Plateau; another population in NW Free State and in W margin of North-west Prov. east to Klerksdorp. Common to abundant but localized. Range expanded in 20th century through use of artificial structures as nest sites. Density up to 80 birds per km2. Description. ADULT Y: forehead, crown and nape pale sandybrown; mantle feathers dark brown to black with pale edges, producing scalloped effect; rump pale sandy-brown; tail dull brown; lores dark brown, cheeks dull cream, with brownish wash; chin black, extending up to lores at corners of bill; throat black, in central patch with rounded border; breast, belly, thighs and undertail-coverts dull cream with brownish wash; flanks with patch of whitish-edged black feathers; wings dull brown, coverts with broad paler margins, underwing whitish. Bill blue-grey, dark grey at base; eyes dark brown; legs dark brown. Sexes alike. SIZE (12 YY, 12 XX): wing, Y 72–79 (749), X 70–78 (746); tail, Y 42–48 (456), X 41–47 (443); bill, Y 161–183 (176), X 168–195 (178); tarsus, Y 191–213 (201), X 194–208 (202). WEIGHT: (Northern Cape, South Africa) Y (n ¼ 73) 238–319 (275), X (n ¼ 43) 240–320 (273). IMMATURE: initially lacks black throat patch, scallops on mantle and flank patches, so appears uniform brownish; some speckling on forehead and crown. Bill pale horn. By 16–18 weeks not distinguishable from adults after first moult.

Philetairus socius

NESTLING: on hatching naked and blind, with pink skin; gape swollen, creamy white, inside of mouth yellow. TAXONOMIC NOTE: various subspecies have been described but regional differences are not consistent (cf. Clancey 1989) and

72

PLOCEIDAE

structure. At one site in Kenya, 5 nests in Acacia mellifera, 1 in A. tortilis; most nests on W side of trees; also nests in tree euphorbias. Birds active at nests, adding material, throughout year. One nest weighed 211 g, contained an estimated 9300 grass stems (Moreau and Moreau 1939). Pad built alongside twig on which bird perches, then extended and arched above twig. Arch is extended on both sides of twig so that tubular nest straddles it and sags at each end, with entrance on either side of supporting twig. 2 birds may work at nest at once.

EGGS: 2–4. White or pinkish, with brown and violet markings. SIZE: 19  14. LAYING DATES: Ethiopia, Feb–May; Kenya, Jan–May, Aug–Sept, Dec; Tanzania, Apr–June, possibly Nov. No further information.

Key References Collias, N. E. and Collias, E. C. (1964), Moreau, R. E. and Moreau, W. M. (1939).

Genus Philetairus Smith

Endemic, monotypic. Small, short-tailed, monomorphic, dull-plumaged weaver of arid SW Africa, where trees or other structures available to support the huge, domed, communal nest mass. Nests constructed of straight grass stems, with numerous (often >100) individual chambers under a common roof. Granivorous.

Plate 5

Philetairus socius (Latham). Sociable Weaver. Re ´publicain social.

(Opp. p. 74)

Loxia socia Latham, 1790. Ind. Orn. i, p. 381; Great Namaqualand, South Africa. Range and Status. Endemic resident. Namibia, Etosha and Ovambo region, separated from population on central plateau, south to Orange R. but absent from coastal desert. Botswana, abundant in SW sector, sparse and localized in SE in Pitsane area; absent from fire-prone central and N Kalahari. South Africa, abundant in Northern Cape from 18
E eastwards, south to 29
300 S and, near Vanwyksvlei, 31
S; absent from Ghaap Plateau; another population in NW Free State and in W margin of North-west Prov. east to Klerksdorp. Common to abundant but localized. Range expanded in 20th century through use of artificial structures as nest sites. Density up to 80 birds per km2. Description. ADULT Y: forehead, crown and nape pale sandybrown; mantle feathers dark brown to black with pale edges, producing scalloped effect; rump pale sandy-brown; tail dull brown; lores dark brown, cheeks dull cream, with brownish wash; chin black, extending up to lores at corners of bill; throat black, in central patch with rounded border; breast, belly, thighs and undertail-coverts dull cream with brownish wash; flanks with patch of whitish-edged black feathers; wings dull brown, coverts with broad paler margins, underwing whitish. Bill blue-grey, dark grey at base; eyes dark brown; legs dark brown. Sexes alike. SIZE (12 YY, 12 XX): wing, Y 72–79 (749), X 70–78 (746); tail, Y 42–48 (456), X 41–47 (443); bill, Y 161–183 (176), X 168–195 (178); tarsus, Y 191–213 (201), X 194–208 (202). WEIGHT: (Northern Cape, South Africa) Y (n ¼ 73) 238–319 (275), X (n ¼ 43) 240–320 (273). IMMATURE: initially lacks black throat patch, scallops on mantle and flank patches, so appears uniform brownish; some speckling on forehead and crown. Bill pale horn. By 16–18 weeks not distinguishable from adults after first moult.

Philetairus socius

NESTLING: on hatching naked and blind, with pink skin; gape swollen, creamy white, inside of mouth yellow. TAXONOMIC NOTE: various subspecies have been described but regional differences are not consistent (cf. Clancey 1989) and

72

PLOCEIDAE

structure. At one site in Kenya, 5 nests in Acacia mellifera, 1 in A. tortilis; most nests on W side of trees; also nests in tree euphorbias. Birds active at nests, adding material, throughout year. One nest weighed 211 g, contained an estimated 9300 grass stems (Moreau and Moreau 1939). Pad built alongside twig on which bird perches, then extended and arched above twig. Arch is extended on both sides of twig so that tubular nest straddles it and sags at each end, with entrance on either side of supporting twig. 2 birds may work at nest at once.

EGGS: 2–4. White or pinkish, with brown and violet markings. SIZE: 19  14. LAYING DATES: Ethiopia, Feb–May; Kenya, Jan–May, Aug–Sept, Dec; Tanzania, Apr–June, possibly Nov. No further information.

Key References Collias, N. E. and Collias, E. C. (1964), Moreau, R. E. and Moreau, W. M. (1939).

Genus Philetairus Smith

Endemic, monotypic. Small, short-tailed, monomorphic, dull-plumaged weaver of arid SW Africa, where trees or other structures available to support the huge, domed, communal nest mass. Nests constructed of straight grass stems, with numerous (often >100) individual chambers under a common roof. Granivorous.

Plate 5

Philetairus socius (Latham). Sociable Weaver. Re ´publicain social.

(Opp. p. 74)

Loxia socia Latham, 1790. Ind. Orn. i, p. 381; Great Namaqualand, South Africa. Range and Status. Endemic resident. Namibia, Etosha and Ovambo region, separated from population on central plateau, south to Orange R. but absent from coastal desert. Botswana, abundant in SW sector, sparse and localized in SE in Pitsane area; absent from fire-prone central and N Kalahari. South Africa, abundant in Northern Cape from 18
E eastwards, south to 29
300 S and, near Vanwyksvlei, 31
S; absent from Ghaap Plateau; another population in NW Free State and in W margin of North-west Prov. east to Klerksdorp. Common to abundant but localized. Range expanded in 20th century through use of artificial structures as nest sites. Density up to 80 birds per km2. Description. ADULT Y: forehead, crown and nape pale sandybrown; mantle feathers dark brown to black with pale edges, producing scalloped effect; rump pale sandy-brown; tail dull brown; lores dark brown, cheeks dull cream, with brownish wash; chin black, extending up to lores at corners of bill; throat black, in central patch with rounded border; breast, belly, thighs and undertail-coverts dull cream with brownish wash; flanks with patch of whitish-edged black feathers; wings dull brown, coverts with broad paler margins, underwing whitish. Bill blue-grey, dark grey at base; eyes dark brown; legs dark brown. Sexes alike. SIZE (12 YY, 12 XX): wing, Y 72–79 (749), X 70–78 (746); tail, Y 42–48 (456), X 41–47 (443); bill, Y 161–183 (176), X 168–195 (178); tarsus, Y 191–213 (201), X 194–208 (202). WEIGHT: (Northern Cape, South Africa) Y (n ¼ 73) 238–319 (275), X (n ¼ 43) 240–320 (273). IMMATURE: initially lacks black throat patch, scallops on mantle and flank patches, so appears uniform brownish; some speckling on forehead and crown. Bill pale horn. By 16–18 weeks not distinguishable from adults after first moult.

Philetairus socius

NESTLING: on hatching naked and blind, with pink skin; gape swollen, creamy white, inside of mouth yellow. TAXONOMIC NOTE: various subspecies have been described but regional differences are not consistent (cf. Clancey 1989) and

Philetairus socius there is variation within a single breeding colony. We therefore treat the species as monotypic.

Field Characters. Length 14 cm. Chunky, with heavy pale bill; brown cap, pale cheeks and small black bib suggest sparrow Passer, but scaled upperparts and flanks distinctive. Juv. lacks black on face, flank markings indistinct. Voice. Tape-recorded (72, 88, 99, B, F, LUT, WAT). Birds at nest maintain fast, high-pitched chippering chatter; individual notes have dry, grating tone but overall effect can be slightly melodious. Song a chatter of 11 single notes followed by 3 paired ones, also described as short series of pleasing ‘chi-chi-chi-chi’ notes falling in pitch (Maclean 1993). All except lowest-ranking YY sing; no song from X. Threat a series of 16 rapid notes, with 2 or 3 at longer intervals. Nest entry call a long phrase of 16–17 notes. On take-off, birds call for several s, until well away from nest, at rate of 7–8 calls per s; flight call a rapid high-pitched ‘chipchip-chip-chip’. Contact call a single ‘chip’. Alarm a sharp ‘tip-tip’, and at nest a burst of noise covering wide frequency range. General Habits. Inhabits open, arid country with scattered trees and bare ground. Usually in flocks of 10–40 birds, occasionally in hundreds. Huge nests a feature of arid southern African landscapes (A), with up to 500 birds in a single nest; birds resident all year. Forages in radius of

15 km from nest site. Usual gait on ground a hop, but runs after insect prey. Collects most food on ground; occasionally hawks flying insects. Seldom drinks, but will do so when water available. Roosts in nest at night and during heat of day, with structure providing some insulation against extreme temperature fluctuations; in winter up to 5 birds may roost together in single chamber, 1–2 in summer (White et al. 1975, Bartholomew et al. 1976). Pygmy Falcon Polihierax semitorquatus in southern Africa occurs only within range of Sociable Weaver, and is apparently entirely dependent on nest masses for roosting and breeding. Falcons occupy 2–4 chambers in active or deserted nests; usually only 1 pair per nest mass. Sociable Weavers tolerate them but often react to their presence with alarm calls, and they may abandon a nest mass if >5 chambers taken over by falcons (Maclean 1970). Ignored Ashy Tits Parus cinerascens roosting in nest but drove off Rosy-faced Lovebirds Agapornis roseicollis trying to enter nests (Macdonald 1983). Other species which may roost or breed in nest chambers include Pied Barbet Tricholaema leucomelaena, Familiar Chat Cercomela familiaris and Redheaded Finch Amadina erythrocephala. Y normally dominant over X; in aggressive interactions, dominant bird is very erect with bill pointing down, wings folded, tail down, head and nape feathers erect, and blackmarked flank feathers erect. Subordinate bird may crouch and gape; rarely, dominant bird briefly mounts subordinate one. At roosting time, dominant bird may eject

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subordinates from chamber. In captivity, initially all 10 birds roosted in single nest chamber. Allopreens rarely (usually subordinate Y preening dominant bird). Resident at nest site all year. Wing-moult protracted; each primary feather takes a month to grow, so moulting overlaps with breeding. Body moult occurs rapidly, during period of greatest food availability. Food. Seeds and insects, especially harvester termites, also grasshoppers, caterpillars and beetle larvae, sometimes termite alates and moths; occasionally soft fruit, ovaries of flowers of Rhigozum trichotomum. By weight, proportions vary according to season, from 16% insects/84% seeds to 80% insects/20% seeds. Young fed insects only. Breeding Habits. Monogamous; colonial, with numerous pairs in single massive communal structure with a common roof and separate nesting chambers; co-operative breeder, adults and sub-adults serving as helpers. Does not pair for life, and successive nesting attempts are commonly with different partners, even if former mate still present in the colony. During courtship, Y, approaching X on ground, holds body more and more erect until tail touches ground, his plumage sleeked; he often holds bit of material in bill. Following X about, he flips tail deliberately over back and down again (B), at rate of 4–24 flips per min., continuing for up to 10 min. Becoming more excited, Y approaches X slowly and deliberately, walking (rather than hopping) with body upright, plumage sleeked, bill pointing up, wings slightly opened and quivering, tail just touching ground (C); he utters 1–2 soft notes per s, circles soliciting X (D), the display ending in copulation (Collias and Collias 1978). Once birds are paired, Y prevents birds other than mate from entering nest chamber. Y sings to advertise nest site and to greet X entering nest. X solicits copulation by crouching low and quivering wings. Up to 4 broods per season; breeding may continue for 9 months. NEST: huge mass of dry grass stems, up to 4 m deep and 72 m long, supported by branches or other structure (E). Largest single structure built by any bird; can weigh up to 1 tonne. Contains numerous individual nests with entrances on underside of nest mass; each is started by bird pushing in straws to form a depression, which becomes a rounded chamber 100–150 in diam., then sides are built out to form a tunnel 60–70 in diam. and up to 250 long. Walls lined with

spiky inward-pointing grass stems. Chamber lined with soft plant material, animal fur, sometimes feathers. Nest material added throughout year, and new chambers added especially after rain; all birds may participate in building. In Kalahari, main building material is the stiff grass Aristida. Sited in tree 35–7 m tall, often on N side (Burger and Gochfeld 1981). Favours Acacia erioloba trees, also uses A. giraffae, A. haematoxylon, Boscia albitrunca, Colophospermum mopane, Aloe dichotoma (Namaqualand, Namibia) and, rarely, Eucalyptus trees (Northern Cape: Anderson 1995). Usually only a single nest mass per tree (86 records), but rarely up to 4 nest masses (8 records); typically 4 m above ground, seldom below 3 m. Sometimes nests in A. giraffae trees which have ant colonies in their thorn bases (Collias and Collias 1977). Since 1949 has nested commonly on telephone poles in treeless areas (Clancey 1950); 1 colony was on 10 poles in a row, each nest mass small, 5 m diam., with max. of 95 chambers, in use for at least 20 years.

Malimbus In captive colony of 10 birds, all participated in nestbuilding as soon as they were released into aviary; dominant YY did most of the work. Bird nips off ends of straws to shorten them; nest lined with pampas grass rather than feathers. EGGS: 2–6 (av. 36, n ¼ 32, South Africa). Dull white, densely spotted with grey in different shades. Over 9-month period, captive X laid 47 eggs in 12 clutches. SIZE: (n ¼ 76, South Africa). 181–228  132–159 (209  151). Up to 4 successive broods; replacement clutches generally smaller than first clutches. LAYING DATES: Namibia, Dec–Aug in N, not seasonal in S but dependent on rainfall; Botswana, all months; South Africa, eastern population Nov–Jan (Tarboton et al. 1987), in Northern Cape not seasonal, timing dependent on irregular rains. INCUBATION: by both Y and X; starts with 1st or 2nd egg; Y contribution variable, generally much less than X, averaging shorter periods on nest. Mean incubation shift 14 min (2–40, n ¼ 22). Period: 13–14 days. DEVELOPMENT AND CARE OF YOUNG: young fed by both parents and at least one ad. helper; in some years up to 9 helpers (i.e. young from previous broods) may assist with feeding. Chicks’ eyes open at 7–10 days, by 14 days all feathers have emerged from sheaths. Nestling period 21–24

days; fledglings dependent on parents for at least 16 days after leaving nest. BREEDING SUCCESS/SURVIVAL: at Kalahari colony, 42% of eggs hatched, 13% of eggs produced flying chicks. Medium-weight fledglings survive better than light or heavy ones (Anon. [M. Anderson, R. Covas] 2003, Afr. Birds & Birding 8 (1), 11). Eggs and young taken from nests by cobra Naja nivea and boomslang Dispholidus typus, also by honey badger Mellivora capensis which climbs tree and rips open nest mass. Occasionally nestlings or adults taken by Pygmy Falcon. Adult birds may be taken by other raptors such as Gabar Goshawk Accipiter gabar. Birds very agitated when Pearl-spotted Owl Glaucidium perlatum entered nest chambers (Etosha, Namibia: Baron and Baron 1976). Large birds of prey such as Giant Eagle Owl Bubo lacteus, White-backed Vulture Gyps africanus and Bateleur Terathopius ecaudatus sometimes use nest mass as a platform for their own nests. Nest masses are susceptible to bush fires. Tree occasionally collapses under weight of nest mass. Key References Bartholomew, G. A. et al. (1976), Channing, A. and Maclean, G. L. (1976), Collias, E. C. and Collias, N .E. (1978, 1981), Maclean, G. L. (1973a, b, c, d, e, 1987), Rudebeck, G. (1956), White, F. N. et al. (1975).

Genus Malimbus Vieillot

Distinctive W and equatorial African lowland rainforest weavers, morphologically very similar to Ploceus, but with black and scarlet or crimson plumage, except for racheliae (black and orange) and ballmanni (black and yellow). Sexes dissimilar, XX having lesser areas of red or yellow (or slightly greater in nitens and erythrogaster); XX of cassini and coronatus lack any red. One sp. crested (malimbicus). Juvs black and red, usually differing in pattern from either adult; juv. rubricollis like adult. Almost exclusively insectivorous. Live mainly high in trees and palms. Nests are closely and neatly woven, some spp. with medium or long downward-hanging entrance tubes, finely but strongly woven and basket-like in scutatus and cassini. Relationships within genus difficult to discern (Moreau 1958, Hall and Moreau 1970). 8 species occur in Cameroon and in Gabon; up to 6 can occur at a single locality. M. rubricollis, nitens and malimbicus all range throughout W African and Congo basin forests and are independent species by all criteria. M. coronatus, a blackish Lower Guinea malimbe with unique nest architecture, is also independent. We concur with Dowsett and Forbes-Watson (1993) in regarding allopatric ballmanni and racheliae as a superspecies. Upper Guinea scutatus is parapatric with the similar Lower Guinea cassini and we treat them as a superspecies, notwithstanding that there may be a tiny enclave of cassini in range of scutatus in Ghana. M. ballmanni/racheliae and M. scutatus/cassini form a group, with relatively short bills and similar plumage patterns, mating systems and nests. M. ibadanensis might form a superspecies either with erythrogaster (Hall and Moreau 1970) or with cassini (Dowsett 1993); pending better knowledge, we treat it, and erythrogaster, as independent. Closely related with Ploceus (Moreau 1958); plumage of M. ballmanni is like Ploceus YY, with yellow merging into chestnut; nest of M. nitens is Ploceus-like (and nest of e.g. P. nigricollis is Malimbus-like). Voices of some Malimbus and Ploceus spp. very alike. [Ploceus] flavipes has been placed in Malimbus (so has Anaplectes rubriceps). A case can be made for uniting Malimbus and Ploceus (Field 1979) when, since Malimbus has date priority over Ploceus, ‘there might be a tendency to supersede the well-known names Ploceinae and Ploceidae’ (Moreau 1960)!

77

Malimbus In captive colony of 10 birds, all participated in nestbuilding as soon as they were released into aviary; dominant YY did most of the work. Bird nips off ends of straws to shorten them; nest lined with pampas grass rather than feathers. EGGS: 2–6 (av. 36, n ¼ 32, South Africa). Dull white, densely spotted with grey in different shades. Over 9-month period, captive X laid 47 eggs in 12 clutches. SIZE: (n ¼ 76, South Africa). 181–228  132–159 (209  151). Up to 4 successive broods; replacement clutches generally smaller than first clutches. LAYING DATES: Namibia, Dec–Aug in N, not seasonal in S but dependent on rainfall; Botswana, all months; South Africa, eastern population Nov–Jan (Tarboton et al. 1987), in Northern Cape not seasonal, timing dependent on irregular rains. INCUBATION: by both Y and X; starts with 1st or 2nd egg; Y contribution variable, generally much less than X, averaging shorter periods on nest. Mean incubation shift 14 min (2–40, n ¼ 22). Period: 13–14 days. DEVELOPMENT AND CARE OF YOUNG: young fed by both parents and at least one ad. helper; in some years up to 9 helpers (i.e. young from previous broods) may assist with feeding. Chicks’ eyes open at 7–10 days, by 14 days all feathers have emerged from sheaths. Nestling period 21–24

days; fledglings dependent on parents for at least 16 days after leaving nest. BREEDING SUCCESS/SURVIVAL: at Kalahari colony, 42% of eggs hatched, 13% of eggs produced flying chicks. Medium-weight fledglings survive better than light or heavy ones (Anon. [M. Anderson, R. Covas] 2003, Afr. Birds & Birding 8 (1), 11). Eggs and young taken from nests by cobra Naja nivea and boomslang Dispholidus typus, also by honey badger Mellivora capensis which climbs tree and rips open nest mass. Occasionally nestlings or adults taken by Pygmy Falcon. Adult birds may be taken by other raptors such as Gabar Goshawk Accipiter gabar. Birds very agitated when Pearl-spotted Owl Glaucidium perlatum entered nest chambers (Etosha, Namibia: Baron and Baron 1976). Large birds of prey such as Giant Eagle Owl Bubo lacteus, White-backed Vulture Gyps africanus and Bateleur Terathopius ecaudatus sometimes use nest mass as a platform for their own nests. Nest masses are susceptible to bush fires. Tree occasionally collapses under weight of nest mass. Key References Bartholomew, G. A. et al. (1976), Channing, A. and Maclean, G. L. (1976), Collias, E. C. and Collias, N .E. (1978, 1981), Maclean, G. L. (1973a, b, c, d, e, 1987), Rudebeck, G. (1956), White, F. N. et al. (1975).

Genus Malimbus Vieillot

Distinctive W and equatorial African lowland rainforest weavers, morphologically very similar to Ploceus, but with black and scarlet or crimson plumage, except for racheliae (black and orange) and ballmanni (black and yellow). Sexes dissimilar, XX having lesser areas of red or yellow (or slightly greater in nitens and erythrogaster); XX of cassini and coronatus lack any red. One sp. crested (malimbicus). Juvs black and red, usually differing in pattern from either adult; juv. rubricollis like adult. Almost exclusively insectivorous. Live mainly high in trees and palms. Nests are closely and neatly woven, some spp. with medium or long downward-hanging entrance tubes, finely but strongly woven and basket-like in scutatus and cassini. Relationships within genus difficult to discern (Moreau 1958, Hall and Moreau 1970). 8 species occur in Cameroon and in Gabon; up to 6 can occur at a single locality. M. rubricollis, nitens and malimbicus all range throughout W African and Congo basin forests and are independent species by all criteria. M. coronatus, a blackish Lower Guinea malimbe with unique nest architecture, is also independent. We concur with Dowsett and Forbes-Watson (1993) in regarding allopatric ballmanni and racheliae as a superspecies. Upper Guinea scutatus is parapatric with the similar Lower Guinea cassini and we treat them as a superspecies, notwithstanding that there may be a tiny enclave of cassini in range of scutatus in Ghana. M. ballmanni/racheliae and M. scutatus/cassini form a group, with relatively short bills and similar plumage patterns, mating systems and nests. M. ibadanensis might form a superspecies either with erythrogaster (Hall and Moreau 1970) or with cassini (Dowsett 1993); pending better knowledge, we treat it, and erythrogaster, as independent. Closely related with Ploceus (Moreau 1958); plumage of M. ballmanni is like Ploceus YY, with yellow merging into chestnut; nest of M. nitens is Ploceus-like (and nest of e.g. P. nigricollis is Malimbus-like). Voices of some Malimbus and Ploceus spp. very alike. [Ploceus] flavipes has been placed in Malimbus (so has Anaplectes rubriceps). A case can be made for uniting Malimbus and Ploceus (Field 1979) when, since Malimbus has date priority over Ploceus, ‘there might be a tendency to supersede the well-known names Ploceinae and Ploceidae’ (Moreau 1960)!

77

78

PLOCEIDAE

Malimbus racheliae superspecies

1

Malimbus scutatus superspecies

1

1

2 2

2? 1 M. ballmanni 2 M. racheliae

1 M. scutatus 2 M. cassini

Plate 4

Malimbus ballmanni Wolters. Gola Malimbe. Malimbe de Gola.

(Opp. p. 27)

Malimbus ballmanni Wolters, 1974. Bonn. Zool. Beitr. 25, p. 283; between Cavally R. and Keibli R., near Taı¨, Ivory Coast. Forms a superspecies with M. racheliae. Range and Status. Endemic resident. Sierra Leone, Gola Forest on Liberian border: seen 15 times in Dec 1971 to June 1972, recorded until 1976 (Field 1979), but not found in Gola survey in 1988/1889 (Allport et al. 1989, Allport 1991). W Liberia, Gola Forest, nesting pair at 7
250 N, 11
500 W in 1989 and several groups at 7
220 N, 11
380 W (population covers area of 200–300 km2); 300 km away in E Liberia, locally common in Grand Gedeh and Sinoe Counties; in Grand Gedeh, from Cess R. to Liberian border near Taı¨; not in Grebo Nat. Forest; in Sinoe, occurs along Sinoe R., in Sapo Nat. Park, south to Juazohn (24 sightings in 2 months in 1987: Gatter and Gardner 1993); first found in Sapo, nesting, in 1988 (Gore 1994). Ivory Coast, type specimen collected near Taı¨, not found in S Taı¨ Nat. Park (Gartshore et al. 1995) but widespread and locally common northwest of Taı¨, along Cavalla R. from near Bakoubli east to 6
220 N 8
000 W in Foreˆt de la Goin and 6
100 N 7
400 W in Foreˆt du Cavally (Gatter and Gardner 1993). Bird ‘resembling this species’ glimpsed in forest near Se´re´dou, Guinea (Halleux 1994). E Liberia and W Ivory Coast population estimated at 20,000–50,000 birds, in area of c. 18,000 km2. Density of at least 3 breeding pairs per 12 ha (Zwedru, Liberia); flocks seen several times per day at separate points along a 1 km forest path (so density much greater than that of allospecies M. racheliae, q.v.). The species, restricted to 1 small and 1 tiny area of Upper Guinea rain forest, is

Malimbus ballmanni

Endangered (Collar et al. 1994); forest between the 2 ranges had already been massively destroyed by early 1990s, and

78

PLOCEIDAE

Malimbus racheliae superspecies

1

Malimbus scutatus superspecies

1

1

2 2

2? 1 M. ballmanni 2 M. racheliae

1 M. scutatus 2 M. cassini

Plate 4

Malimbus ballmanni Wolters. Gola Malimbe. Malimbe de Gola.

(Opp. p. 27)

Malimbus ballmanni Wolters, 1974. Bonn. Zool. Beitr. 25, p. 283; between Cavally R. and Keibli R., near Taı¨, Ivory Coast. Forms a superspecies with M. racheliae. Range and Status. Endemic resident. Sierra Leone, Gola Forest on Liberian border: seen 15 times in Dec 1971 to June 1972, recorded until 1976 (Field 1979), but not found in Gola survey in 1988/1889 (Allport et al. 1989, Allport 1991). W Liberia, Gola Forest, nesting pair at 7
250 N, 11
500 W in 1989 and several groups at 7
220 N, 11
380 W (population covers area of 200–300 km2); 300 km away in E Liberia, locally common in Grand Gedeh and Sinoe Counties; in Grand Gedeh, from Cess R. to Liberian border near Taı¨; not in Grebo Nat. Forest; in Sinoe, occurs along Sinoe R., in Sapo Nat. Park, south to Juazohn (24 sightings in 2 months in 1987: Gatter and Gardner 1993); first found in Sapo, nesting, in 1988 (Gore 1994). Ivory Coast, type specimen collected near Taı¨, not found in S Taı¨ Nat. Park (Gartshore et al. 1995) but widespread and locally common northwest of Taı¨, along Cavalla R. from near Bakoubli east to 6
220 N 8
000 W in Foreˆt de la Goin and 6
100 N 7
400 W in Foreˆt du Cavally (Gatter and Gardner 1993). Bird ‘resembling this species’ glimpsed in forest near Se´re´dou, Guinea (Halleux 1994). E Liberia and W Ivory Coast population estimated at 20,000–50,000 birds, in area of c. 18,000 km2. Density of at least 3 breeding pairs per 12 ha (Zwedru, Liberia); flocks seen several times per day at separate points along a 1 km forest path (so density much greater than that of allospecies M. racheliae, q.v.). The species, restricted to 1 small and 1 tiny area of Upper Guinea rain forest, is

Malimbus ballmanni

Endangered (Collar et al. 1994); forest between the 2 ranges had already been massively destroyed by early 1990s, and

Malimbus ballmanni fragmentation and clearing of forest for slash-and-burn farming together with heavy logging even in Sapo Nat. Park, was accelerating during civil strife when population estimates were made (Gatter and Gardner 1993). Description. ADULT Y: glossy deep black, but nape golden yellow or orange-yellow, upper quarter and sides with chestnut or cinnamon tinge; large patch on breast bright cadmium yellow with faint chestnut tinge visible in some lights, undertail-coverts bright daffodil or cadmium yellow (rarely washed with orange). Yellow nape and yellow breast usually separated by a few black feathers. A clear indentation formed by a few small black feathers at lower edge of breast. Underside of wing and tail matt grey, underwing-coverts greyish black. P6–9 longest. Bill shiny black; eyes bright wine-red or dark brown; legs black or dark grey, claws and underside of toes sepia. ADULT X: like Y but nape black and breast purer yellow, with lower part divided centrally to about half way up by thin dark line (Field 1979). SIZE: wing, Y (n ¼ 10) 85–94, X (n ¼ 7) 84–92; tail, Y (n ¼ 10) 57–64, X (n ¼ 7) 60–64; bill to skull, Y (n ¼ 8) 165–175, X (n ¼ 5) 160–175; tarsus, Y (n ¼ 8) 200–225, X (n ¼ 5) 200–215. WEIGHT: not recorded. IMMATURE: Y like ad. Y but greyer and less shiny; forehead, crown and throat pale yellow, orange-yellow or matt orangebrown; yellow of throat confluent with yellow breast; nape bar paler yellow and less extensive, and breast indentation less clear than in ad. Y. Imm. bill horn-yellow or pale flesh-pink; later, bill black with only underside of lower mandible yellow; eyes paler red than in ad.; legs dark sepia, claws horn-brown. X like imm. Y but lacks yellow nape bar.

Field Characters. Length 14–155 cm. The only malimbe (or ploceid) in Upper Guinea high forest with combination of yellow breast, black belly and yellow undertail-coverts; Y with yellow and X with black nape. Immatures greyer, with some yellow on forehead, crown and throat. Voice. (Not tape-recorded). Song of Y a rough, discordant series of chatters followed by wheeze lasting 3 s, ‘cheg chig cheg cheg chega zzzzzzzzzzzzzzzzzzzzz’, strikingly like song of Village Weaver Ploceus cucullatus. Song of X similar but without the wheeze: ‘cheg cheg chig chag chaaag cheg chiiig’. Songs carry for 100 m (Gatter and Gardner 1993). Call a low, conversational ‘chch chchchch’, given whilst foraging (Field 1979). General Habits. Inhabits middle storey of primary lowland rain forest, up to 400 m, also moderately and heavily logged high forest and very old, mature secondary forest; keeps mainly to outermost, open parts of understorey tree-tops. Solitary, in pairs or trios; of 159 observations, 43 of single birds, 72 of pairs and 44 of trios; but habitually occurs in mixed-species foraging flocks, at all times of year, when up to 8 Gola Malimbes together. Forages actively, rather like a tit Parus, by gleaning open green leaves and horizontal twigs 3–10 mm in diam. for insects; also investigates vertical twigs and thick tangles of lianacovered branches; rarely, probes into dry, rolled leaves. Forages generally between 8 and 22 m above forest floor (79% of 239 observations), rarely down to 2 m and above 25 m. Sedentary. Food. Insects in range 1–30 mm long, including grasshoppers and mantids.

Breeding Habits. Solitary nester, evidently co-operative. Displaying Y sings, facing X, with yellow breast feathers lifted and yellow undertail-coverts expanded (A). NEST: a free-hanging ‘inverted sock’ with globular egg chamber and tubular entrance pointing down, suspended from tip of thin, vertically hanging liana by means of anchoring structure; very like nest of M. racheliae (Dowsett 1993). Whole structure est. 60–110 cm long: anchoring structure 25–70 cm long, nest with entrance tunnel 35–45 (tunnel being longer than diam. of egg chamber). Sited (n ¼ 13) between 8 and 21 m above ground (6 at 10–15 m, 5 at 16–20 m), always suspended from vines (never on climbing palms) 15–6 m horizontally from nearest branch or trunk. 2–5 birds at a time, from a group of up to 8 in a roving mixed-species foraging flock, build nest ‘on the move’, visiting it several times a day at irregular intervals, whenever mixed-species flock passes by. Group members can make >10 nest visits in 5 min; some work at weaving whilst others fly to and fro with building material and wait nearby for a place to become vacant so that they can build. Construction can cease for several hours when mixedspecies flock moves away, but resumes immediately flock returns. Initial anchoring of nest at tip of thin hanging liana is ‘chief task of the YY’; entrance tunnel and interior built primarily by XX (Gatter and Gardner 1993). EGGS: not known. Clutch size probably 2; 23 family parties were of 1–2 juvs accompanying 1–2 adults (and a 24th: 1 juv. with 3 adults). LAYING DATES: Liberia, (nest-building mainly in 2nd half of rainy season: Sept–Nov; a few nests built in July– Aug; young being fed out of nest from late Oct to mid Mar; independent young from late Jan to mid May). DEVELOPMENT AND CARE OF YOUNG: nestling fed by at least 3 adults, but only when mixed-species foraging flock passes nearby. Key Reference Gatter, W. and Gardner, R. (1993).

79

80

PLOCEIDAE

Plate 4

Malimbus racheliae (Cassin). Rachel’s Malimbe. Malimbe de Rachel.

(Opp. p. 27)

Sycobius racheliae Cassin, 1857. Proc. Acad. Nat. Sci. Philadelphia, p. 36; River Muni (Spanish Guinea). Forms a superspecies with M. ballmanni. Range and Status. Endemic resident, extreme SE Nigeria, W Cameroon, Mbini and Gabon. Nigeria, frequent to common within 20 km of Calabar, in Oban Hills and Cross River Nat. Park (Ikpan block and Oban West: up to 10 birds seen daily). Cameroon, forest mainly within 100 km of coast (Ndian, Kumba, Massaka in W, Campo, Efulen, Nlo Ayong in SW, inland east to Minkalli at 3
030 N, 11
350 E); 2 populations, in W and SW (12 and 5 localities respectively: M. Languy, pers. comm., 2002). Mbini, at 600–900 m in Mt Alen Nat. Park. Gabon, widespread in Ivondo catchment region (Makokou, Sin R., Ilindi) but not common; Lope´. Density estimated at pair per 15 km2 of primary forest (Gabon).

Malimbus racheliae

Description. ADULT Y: forehead to hindneck orange-red; rest of upperparts black. Tail black. Lores and above eye to ear-coverts, cheeks, chin and upper throat black. Upper sides of neck orangeyellow, continuous with orange nape and hindneck and linking narrowly with large patch on lower throat and upper breast, orange-scarlet, grading to yellow at sides and at rear. Lower breast, belly, flanks and thighs black; undertail-coverts yellow. Entire upperwing black; underwing-coverts sooty black. P7–8 longest, P6 3 shorter, P5 c. 5 shorter, P9 c. 4 shorter. P10 rather small and pointed, 8 mm longer than primary coverts. Bill black; eyes dark red or red-brown; legs grey. ADULT X: like ad. Y, but head and neck entirely black. SIZE (8 YY, 7 XX): wing, Y 76–82 (796), X 74–83 (792); tail, Y 44–48 (462), X 44–47 (454); bill, Y 18–20 (187), X 175–195 (183); tarsus, Y 20–205 (202), X 20–205 (203). IMMATURE: juvenile like ad. X, but black areas tinged brown, and chin and upper throat yellowish brown.

Field Characters. Length 14–155 cm. Black face and orange-red crown like other malimbes in its range, e.g. Red-vented Malimbe M. scutatus, but the only one with lower breast and undertail-coverts yellow. X like Y but head and neck black; juv. a browner version of X with touch of yellow on chin and upper throat. Voice. Tape-recorded (B, MAC, ROD). Chattering ‘chop’ and ‘chip’ notes; also said to give long-drawn buzzing ‘cheeee-ee’ like Village Weaver Ploceus cucullatus (Bannerman 1949).

General Habits. Inhabits primary rain forest, keeping in canopy; occasionally in understorey down to 10 m above ground; comes to edge of forest in groves of parasol trees. Solitary in pairs or more often trios, or in small family parties in mixed foraging flocks of insectivores such as barbets and drongos (Nigeria, Gabon); in 43% of mixedspecies flocks (Korup, Cameroon). Usual social unit is trio of 2 YY and 1 X which keep together all year, accompanied by their young after breeding season. Forages actively amongst foliage, like tit Parus sp. Bathes in small stream on forest floor, and then perches in under-shrub to preen. Sedentary.

Food. Insects, including caterpillars, moths, orthopterans and mantids (Brosset and Erard 1986).

Breeding Habits. Solitary nester, evidently monogamous and territorial. NEST: 3 were all suspended from tip of slender liana, 8–12 m above forest floor, well below forest canopy; each built by 2 YY and 1 X, working together at same time. EGGS: not known. LAYING DATES: Nigeria, (nests Calabar Apr, June, Nov; nest-building, and dependent juvs, Cross River Nat. Park, Apr); Gabon (nests Mar, June, Nov; feeding young just out of nest, Dec). DEVELOPMENT AND CARE OF YOUNG: fledgling fed by trio of adults (2 YY, 1 X).

Key References (1986).

Brosset, A. (1978), Brosset, A. and Erard, C.

Malimbus scutatus

81

Malimbus scutatus (Cassin). Red-vented Malimbe. Malimbe a` queue rouge.

Plate 4

S[y]cobius scutatus Cassin, 1849. Proc. Acad. Nat. Sci. Philadelphia, 4, p. 157; Western Africa (inferred to be Sierra Leone).

(Opp. p. 27)

Forms a superspecies with M. cassini. Range and Status. Endemic resident. Sierra Leone, widespread in S and SE, closely associated with oil-palms: west to Freetown Peninsula (only around Russell), north to Rokupr (Makeni) and Sefadu. Guinea, common in SE, close to Sierra Leone and Liberia borders: Gue´kedou, Macenta, Nze´re´kore´. Liberia, common resident in coastal lowlands and N highlands, absent or at least uncommon in intervening country. Ivory Coast, common throughout forest zone, north to Sipilou, Lamto and Bossematie´. Ghana, widespread in forest zone and coastal gallery forests. Togo, recorded only at Misahohe¨, 1990. Benin, forest zone in extreme S and SE. Nigeria, common in forest zone and patches of derived savanna, from coast north to Ilaro, Abeokuta, Ibadan, Ife, Ubiaja, Enugu and Nsukka; often the commonest malimbe. Cameroon, in W, north to Kumba and Massaka (40
500 N), south to Douala and Sanaga R., east to Bakebe. Description. M. s. scutatus (Cassin): Sierra Leone to Ghana. ADULT Y: forehead to hindneck and sides of neck orange-red; rest of upperparts black, slightly glossy. Tail black. Lores and above eye to ear-coverts, cheeks, chin and upper throat black; lower throat to upper breast and sides of breast orange-red, linked to red neck collar. Lower breast, belly, flanks and thighs black; undertail-coverts orange-red. Upperwing black, coverts, tertials and outer edges of flight feathers slightly glossy; underwing-coverts sooty black. P7–8 longest, P6 2 shorter, P5 4 shorter, P9 1–3 shorter, P10 small and pointed, c. 8 mm longer than primary coverts. Bill black; eyes brown or chestnut-brown; legs blackish brown. ADULT X: differs from ad. Y in having head and neck entirely black; scarlet confined to patch on lower throat and upper breast and undertail-coverts. SIZE (10 YY, 10 XX): wing, Y 86–94 (908), X 86–91 (887); tail, Y 51–57 (534), X 47–53 (507); bill, Y 19–20 (193), X 18–21 (193); tarsus, Y 22–24 (230), X 21–23 (222). WEIGHT: Liberia, Y (n ¼ 3) 28, 29, 33, X (n ¼ 3) 29, 29, 32. IMMATURE: juvenile similar to ad. X but more sooty black; orange on breast and undertail-coverts duller; pale orange extends to chin and upper throat and sometimes to forecheeks and forehead. M. s. scutopartitus Reichenow: SW Nigeria to SW Cameroon. Red of breast sides extends further posteriorly. X has narrow black mid-ventral band separating orange-red breast sides. Slightly larger: wing, Y (n ¼ 10) 88–97 (927).

Field Characters. Length 14–145 cm. Black face surrounded by red crown, collar and breast; differs from similar Cassin’s Malimbe M. cassini by red vent. X has head and neck black, with red patch on breast; immature has chin to breast and undertail-coverts dull pale orange, sometimes orange on cheeks or forehead. Voice. Tape-recorded (104, B, CHA). Song melodious and pleasant, with repetition of phrase, e.g. ‘peeyo-peeyo-tsitsipyurr’, often lengthened, ‘peeyo-peeyo-tsi-tsi-pyurrr-tsuwee-tsurr-tsurr-tsu-wee-tsitsi-wer’; another variation a low-key mixture of chattering and clear whistles, with bustling almost canary-like quality. Calls include a long, nasal ‘zu-weeya’ and frequent loud, harsh ‘chirp’; contact

Malimbus scutatus

calls from party harsh ‘zee-zee-zee’ and ‘chit-it-zeer-zeer’; both sexes said to utter low soft notes varying in tone (F. C. Holman in Bannerman 1949). General Habits. Original habitat thought to be upper canopy and emergents of mature lowland forest; now inhabits mature and secondary forest, small and large forest clearings, forest edges, gallery forest, swamp forest, and well-wooded derived savanna. Perhaps more closely associated with swamp forest in E of range than in W. At Owerri, Nigeria, occurs in tall bush along stream courses filled with Raphia palms and among open stands of palms, also in gallery forests around villages and in derived savanna (Marchant 1953). Ranges from sea level up to 1070 m (Mt Cameroon). Closely associated also with Raphia, rattan and oil palms Elaeis guineensis, living near and nesting in them in both forest and open, cultivated land and around villages. Keeps mainly to upper strata in forest, foraging at 15–>50 m, mainly at 30–50 m (the highest of any malimbe: Gatter 1997) but nests lower down, in palms. In pairs or groups of up to 5 adults; once ‘a dozen’ (Sierra Leone), but ‘usually occurs in large parties’ (SE Nigeria, Marchant 1953). Sometimes joins mixed-species foraging flocks of insectivores, but less frequently than some congeners do. Forages for 70% of time on horizontal outer twigs, 50 m, mainly 15–45 (Gatter 1997). Clings to bark, often on dead tree, creeping and climbing like a nuthatch Sitta, examining crevices and fissures and using head as a hammer to dig in with bill like a small woodpecker. Takes grubs out of dead wood. Gleans mainly on horizontal structures, hopping along them; on vertical ones jumps, hops or shuffles upward, sideways and downward, sometimes balancing by opening wings, and makes short flight down to next vine or trunk. On large, horizontal branch moves forward constantly turning 90
to right and left, leaning right over to examine underside of branch (Brosset and Erard 1986,

Anaplectes rubriceps Gatter 1997); runs after a fleeing insect. Searches amongst lichens, thick masses of moss and young shoots of epiphytes; clings and hops up full length of long hanging liana. Dislodges fruit of oil-palm Elaeis guineensis from inflorescence, holds it under feet, wedging it into axil of palm frond, and strips and eats small pieces (Pettet 1969). Occasionally keeps watch from isolated dead branch at edge of clearing and flies out to catch passing insect; exploits hatches of flying termites. Sedentary. Food. Invertebrates, including orthopterans, winged termites, caterpillars, beetles, beetle larvae, spiders and tiny snails; also fruits of oil-palm. Breeding Habits. Poorly known. Often 2–5 nests clustered together, probably all built by single pair. X defoliates immediate surrounds of nest. NEST: large, globular, untidy mass with short but wide entrance tunnel (A); built largely of tendrils of climber Paullinia pennata (Ibadan, Nigeria) or of dry roots of epiphytic orchid (Owerri, Nigeria). Entrance tunnel c. 22–25 cm long (Button 1967). Suspended by its top just below branch; one was also supported by a small branch

97

passing beneath brood chamber. 2 or 3 nests sometimes fused together. Sited at 30–45 m above ground in mature forest, 6–30 m high in secondary habitats; one cluster of 5 nests thought to be ‘200 ft’ (65 m) above forest floor. Sometimes nests in colonies of weaver Ploceus tricolor and often 1–2 m from occupied nests of Palm-nut Vulture Gypohierax angolensis or very close to wasps’ and bees’ nests; also sometimes nests in same tree as other Malimbus spp., drongos, barbets or rollers (Marchant 1953). EGGS: 2. Pure white. LAYING DATES: Liberia, (nest-building mainly Oct; occupied nests Jan–Apr, mainly Mar–Apr); Ivory Coast, Taı¨, (nest with young Dec, carrying food Feb, collecting nest material Mar); Ghana, (at nests Nov, Dec, Mar); Togo, Mar; Benin, (at nests Apr, May); Nigeria, (at nests in nearly all months, particularly Nov–Apr: Marchant 1953); Angola, (fledglings Feb–Mar); Zaı¨re, Uele, (enlarged gonads Apr, May, Oct; oviduct egg Oct), Itombwe, June (and juvs Mar, Aug); Uganda, Apr, May, Aug. Nothing further known. Key References (1997).

Brosset, A. and Erard, C. (1986), Gatter, W.

Genus Anaplectes Reichenbach Endemic, monotypic. A medium-sized weaver with moderately long, slender, pointed bill, culmen rounded at base, straight for basal two-thirds, descending towards tip, red or orange. Wings long and pointed; P9 to P6 nearly equal, P10 reduced. Tail short and square-tipped. Feet and tarsi fairly strong; claws short and sharply decurved. Nest suspended from savanna tree, woven from dry twigs with a long entrance tunnel. Previously placed in Malimbus, based on red plumage of breeding Y and nest form, or in Ploceus, based on yellow in plumage, particularly of X, and resemblance of nest to that of some forest weavers (e.g. Ploceus bicolor, perhaps as a consequence of using similar dry materials). Y and X lack black plumage (as occurs in all Malimbus spp.) and juvenile resembles X (whereas Malimbus spp. have distinctive juvenile plumages). Elgood (1982) argued for retention of Anaplectes, and we follow Sibley and Monroe (1990) and Dowsett and Forbes-Watson (1993) in recognizing this genus, perhaps the savanna derivative of a forest genus.

Anaplectes rubriceps (Sundevall). Red-headed Weaver. Tisserin e ´carlate.

Plate 12

Ploceus (Hyphantornis) rubriceps Sundevall, 1850. Oefv. K. Sv. Vet.-Akad. Fo ¨rhandl., vol. vii, p. 97; ‘Caffraria superiore’; type from Mohapoani, Rustenberg district, W Transvaal (Gyldenstolpe, 1927, Ark. Zool. vol. 19A(1), p.12).

(Opp. p. 155)

Range and Status. Endemic resident, widespread in Africa south of 14
N; frequent to common or uncommon. 2 records in Gambia: Barra and Mile 205; also Seleti in Senegal on Gambian border. Mapped by Borrow and Demey (2001) in broad band through SE Senegal, NE Guinea, extreme SW Mali and N Ivory Coast, but we do not know of any published records from those areas. N guinea savanna belt from SE Mali to Nigeria, Cameroon and Central African Republic; south to Mambilla Plateau in Nigeria. Savanna in S Sudan; Ethiopia and NW Somalia at 1200–1800 m in woodlands, absent from dry E Somalia, race jubaensis south

of 2
300 N in lowlands along mid and lower Jubba R., south to Kiwayu in Kenya in moist coastal bush. In Uganda and Kenya, generally below 2000 m; absent from arid areas of NE Kenya, and from coastal lowlands to 10
S in Tanzania, where otherwise widespread. Rwanda, Burundi. Frequent in E and S Zaı¨re; vagrant Ngula, Bandundu (Louette 1988). Angola, locally common in woodlands of central plateau, north to Cuanza Norte, Bie´ and Moxico. Sparse in N Namibia; throughout Zambia, where commonest in Braˆ i widespread. Local in drier interior of chystegia. Malaw Mozambique, absent from coastal littoral. Uncommon to

Anaplectes rubriceps Gatter 1997); runs after a fleeing insect. Searches amongst lichens, thick masses of moss and young shoots of epiphytes; clings and hops up full length of long hanging liana. Dislodges fruit of oil-palm Elaeis guineensis from inflorescence, holds it under feet, wedging it into axil of palm frond, and strips and eats small pieces (Pettet 1969). Occasionally keeps watch from isolated dead branch at edge of clearing and flies out to catch passing insect; exploits hatches of flying termites. Sedentary. Food. Invertebrates, including orthopterans, winged termites, caterpillars, beetles, beetle larvae, spiders and tiny snails; also fruits of oil-palm. Breeding Habits. Poorly known. Often 2–5 nests clustered together, probably all built by single pair. X defoliates immediate surrounds of nest. NEST: large, globular, untidy mass with short but wide entrance tunnel (A); built largely of tendrils of climber Paullinia pennata (Ibadan, Nigeria) or of dry roots of epiphytic orchid (Owerri, Nigeria). Entrance tunnel c. 22–25 cm long (Button 1967). Suspended by its top just below branch; one was also supported by a small branch

97

passing beneath brood chamber. 2 or 3 nests sometimes fused together. Sited at 30–45 m above ground in mature forest, 6–30 m high in secondary habitats; one cluster of 5 nests thought to be ‘200 ft’ (65 m) above forest floor. Sometimes nests in colonies of weaver Ploceus tricolor and often 1–2 m from occupied nests of Palm-nut Vulture Gypohierax angolensis or very close to wasps’ and bees’ nests; also sometimes nests in same tree as other Malimbus spp., drongos, barbets or rollers (Marchant 1953). EGGS: 2. Pure white. LAYING DATES: Liberia, (nest-building mainly Oct; occupied nests Jan–Apr, mainly Mar–Apr); Ivory Coast, Taı¨, (nest with young Dec, carrying food Feb, collecting nest material Mar); Ghana, (at nests Nov, Dec, Mar); Togo, Mar; Benin, (at nests Apr, May); Nigeria, (at nests in nearly all months, particularly Nov–Apr: Marchant 1953); Angola, (fledglings Feb–Mar); Zaı¨re, Uele, (enlarged gonads Apr, May, Oct; oviduct egg Oct), Itombwe, June (and juvs Mar, Aug); Uganda, Apr, May, Aug. Nothing further known. Key References (1997).

Brosset, A. and Erard, C. (1986), Gatter, W.

Genus Anaplectes Reichenbach Endemic, monotypic. A medium-sized weaver with moderately long, slender, pointed bill, culmen rounded at base, straight for basal two-thirds, descending towards tip, red or orange. Wings long and pointed; P9 to P6 nearly equal, P10 reduced. Tail short and square-tipped. Feet and tarsi fairly strong; claws short and sharply decurved. Nest suspended from savanna tree, woven from dry twigs with a long entrance tunnel. Previously placed in Malimbus, based on red plumage of breeding Y and nest form, or in Ploceus, based on yellow in plumage, particularly of X, and resemblance of nest to that of some forest weavers (e.g. Ploceus bicolor, perhaps as a consequence of using similar dry materials). Y and X lack black plumage (as occurs in all Malimbus spp.) and juvenile resembles X (whereas Malimbus spp. have distinctive juvenile plumages). Elgood (1982) argued for retention of Anaplectes, and we follow Sibley and Monroe (1990) and Dowsett and Forbes-Watson (1993) in recognizing this genus, perhaps the savanna derivative of a forest genus.

Anaplectes rubriceps (Sundevall). Red-headed Weaver. Tisserin e ´carlate.

Plate 12

Ploceus (Hyphantornis) rubriceps Sundevall, 1850. Oefv. K. Sv. Vet.-Akad. Fo ¨rhandl., vol. vii, p. 97; ‘Caffraria superiore’; type from Mohapoani, Rustenberg district, W Transvaal (Gyldenstolpe, 1927, Ark. Zool. vol. 19A(1), p.12).

(Opp. p. 155)

Range and Status. Endemic resident, widespread in Africa south of 14
N; frequent to common or uncommon. 2 records in Gambia: Barra and Mile 205; also Seleti in Senegal on Gambian border. Mapped by Borrow and Demey (2001) in broad band through SE Senegal, NE Guinea, extreme SW Mali and N Ivory Coast, but we do not know of any published records from those areas. N guinea savanna belt from SE Mali to Nigeria, Cameroon and Central African Republic; south to Mambilla Plateau in Nigeria. Savanna in S Sudan; Ethiopia and NW Somalia at 1200–1800 m in woodlands, absent from dry E Somalia, race jubaensis south

of 2
300 N in lowlands along mid and lower Jubba R., south to Kiwayu in Kenya in moist coastal bush. In Uganda and Kenya, generally below 2000 m; absent from arid areas of NE Kenya, and from coastal lowlands to 10
S in Tanzania, where otherwise widespread. Rwanda, Burundi. Frequent in E and S Zaı¨re; vagrant Ngula, Bandundu (Louette 1988). Angola, locally common in woodlands of central plateau, north to Cuanza Norte, Bie´ and Moxico. Sparse in N Namibia; throughout Zambia, where commonest in Braˆ i widespread. Local in drier interior of chystegia. Malaw Mozambique, absent from coastal littoral. Uncommon to

Anaplectes rubriceps Gatter 1997); runs after a fleeing insect. Searches amongst lichens, thick masses of moss and young shoots of epiphytes; clings and hops up full length of long hanging liana. Dislodges fruit of oil-palm Elaeis guineensis from inflorescence, holds it under feet, wedging it into axil of palm frond, and strips and eats small pieces (Pettet 1969). Occasionally keeps watch from isolated dead branch at edge of clearing and flies out to catch passing insect; exploits hatches of flying termites. Sedentary. Food. Invertebrates, including orthopterans, winged termites, caterpillars, beetles, beetle larvae, spiders and tiny snails; also fruits of oil-palm. Breeding Habits. Poorly known. Often 2–5 nests clustered together, probably all built by single pair. X defoliates immediate surrounds of nest. NEST: large, globular, untidy mass with short but wide entrance tunnel (A); built largely of tendrils of climber Paullinia pennata (Ibadan, Nigeria) or of dry roots of epiphytic orchid (Owerri, Nigeria). Entrance tunnel c. 22–25 cm long (Button 1967). Suspended by its top just below branch; one was also supported by a small branch

97

passing beneath brood chamber. 2 or 3 nests sometimes fused together. Sited at 30–45 m above ground in mature forest, 6–30 m high in secondary habitats; one cluster of 5 nests thought to be ‘200 ft’ (65 m) above forest floor. Sometimes nests in colonies of weaver Ploceus tricolor and often 1–2 m from occupied nests of Palm-nut Vulture Gypohierax angolensis or very close to wasps’ and bees’ nests; also sometimes nests in same tree as other Malimbus spp., drongos, barbets or rollers (Marchant 1953). EGGS: 2. Pure white. LAYING DATES: Liberia, (nest-building mainly Oct; occupied nests Jan–Apr, mainly Mar–Apr); Ivory Coast, Taı¨, (nest with young Dec, carrying food Feb, collecting nest material Mar); Ghana, (at nests Nov, Dec, Mar); Togo, Mar; Benin, (at nests Apr, May); Nigeria, (at nests in nearly all months, particularly Nov–Apr: Marchant 1953); Angola, (fledglings Feb–Mar); Zaı¨re, Uele, (enlarged gonads Apr, May, Oct; oviduct egg Oct), Itombwe, June (and juvs Mar, Aug); Uganda, Apr, May, Aug. Nothing further known. Key References (1997).

Brosset, A. and Erard, C. (1986), Gatter, W.

Genus Anaplectes Reichenbach Endemic, monotypic. A medium-sized weaver with moderately long, slender, pointed bill, culmen rounded at base, straight for basal two-thirds, descending towards tip, red or orange. Wings long and pointed; P9 to P6 nearly equal, P10 reduced. Tail short and square-tipped. Feet and tarsi fairly strong; claws short and sharply decurved. Nest suspended from savanna tree, woven from dry twigs with a long entrance tunnel. Previously placed in Malimbus, based on red plumage of breeding Y and nest form, or in Ploceus, based on yellow in plumage, particularly of X, and resemblance of nest to that of some forest weavers (e.g. Ploceus bicolor, perhaps as a consequence of using similar dry materials). Y and X lack black plumage (as occurs in all Malimbus spp.) and juvenile resembles X (whereas Malimbus spp. have distinctive juvenile plumages). Elgood (1982) argued for retention of Anaplectes, and we follow Sibley and Monroe (1990) and Dowsett and Forbes-Watson (1993) in recognizing this genus, perhaps the savanna derivative of a forest genus.

Anaplectes rubriceps (Sundevall). Red-headed Weaver. Tisserin e ´carlate.

Plate 12

Ploceus (Hyphantornis) rubriceps Sundevall, 1850. Oefv. K. Sv. Vet.-Akad. Fo ¨rhandl., vol. vii, p. 97; ‘Caffraria superiore’; type from Mohapoani, Rustenberg district, W Transvaal (Gyldenstolpe, 1927, Ark. Zool. vol. 19A(1), p.12).

(Opp. p. 155)

Range and Status. Endemic resident, widespread in Africa south of 14
N; frequent to common or uncommon. 2 records in Gambia: Barra and Mile 205; also Seleti in Senegal on Gambian border. Mapped by Borrow and Demey (2001) in broad band through SE Senegal, NE Guinea, extreme SW Mali and N Ivory Coast, but we do not know of any published records from those areas. N guinea savanna belt from SE Mali to Nigeria, Cameroon and Central African Republic; south to Mambilla Plateau in Nigeria. Savanna in S Sudan; Ethiopia and NW Somalia at 1200–1800 m in woodlands, absent from dry E Somalia, race jubaensis south

of 2
300 N in lowlands along mid and lower Jubba R., south to Kiwayu in Kenya in moist coastal bush. In Uganda and Kenya, generally below 2000 m; absent from arid areas of NE Kenya, and from coastal lowlands to 10
S in Tanzania, where otherwise widespread. Rwanda, Burundi. Frequent in E and S Zaı¨re; vagrant Ngula, Bandundu (Louette 1988). Angola, locally common in woodlands of central plateau, north to Cuanza Norte, Bie´ and Moxico. Sparse in N Namibia; throughout Zambia, where commonest in Braˆ i widespread. Local in drier interior of chystegia. Malaw Mozambique, absent from coastal littoral. Uncommon to

98

PLOCEIDAE

Anaplectes rubriceps

? ? ?

?

frequent in N and E Botswana, absent from the central Kalahari and SW. Zimbabwe, throughout. N Transvaal south to the Magaliesberg, commonest in E, extending into E Swaziland and extreme N Natal to Ndumu Nature Reserve. Population in S Mozambique probably >10,000 birds (Parker 1999). Density in Acacia savanna, Swaziland, 5 birds per 100 ha; Swaziland population c. 300 (Parker 1994). Density in Moremi Wildlife Res., Botswana, 021 birds per ha (Milewski and Campbell 1976). Description. A. r. leuconotus Mu ¨ ller: Senegal to Ethiopia, N Somalia, Uganda, Kenya, N and W Tanzania, E Zaı¨re (south to N Katanga) and N Angola. ADULT Y (breeding): forehead to hindneck and sides of neck scarlet; mantle to uppertail-coverts and scapulars brownish grey, upper mantle sometimes darker sooty grey, and often with a few red feathers. Tail dark greybrown, edges of outer webs red, becoming narrower and greyish distally. Lores and narrowly above eye to ear-coverts, cheeks and upper chin sooty black, forming a face mask; lower chin, throat and upper breast scarlet, sometimes with whitish feathers showing through; scarlet often intense on lower throat and upper breast and may extend to centre of lower breast; rest of underparts greyish white. Remiges dark grey brown, primaries edged orange-red proximal to emarginations, secondaries and outermost tertial with red outer edges, becoming pale greyish distally and around tip, inner two tertials fringed pale grey buff; upperwing-coverts dark grey-brown, greaters fringed pale greybuff, bases of outer edges tinged red, medians tipped pale greyish; lesser coverts sometimes dark sooty grey. Underwing-coverts and axillaries pale grey. Bill orange or orange-red; eyes light redbrown to dark red or deep orange; legs pale flesh to dusky brown. ADULT Y (non-breeding): like ad. X, but often with a few red feathers on top of head and from chin to upper breast. ADULT X: top and sides of head and neck brownish grey; underparts white, washed grey-buff or olive-buff, most strongly on lower throat and breast; rest of upperparts, tail and wings as ad. Y. Bill orange or pinkish orange; eyes and legs as ad. Y. SIZE (10 YY, 10 XX): wing, Y 82–87 (843), X 77–82 (787); tail, Y 46–56 (493), X

41–51 (479); bill, Y 19–21 (198), X 17–185 (180); tarsus, Y 20– 215 (209), X 19–20 (196). WEIGHT: Ghana, Y (n ¼ 1) 22. IMMATURE: similar to ad. X, but head and breast tinged more olive; edges of wing and tail feathers orange or orange-yellow. NESTLING: undescribed. A. r. rubriceps Sundevall: N Botswana, Transvaal and Mozambique north to central Angola, Zambia, SE Zaı¨re (S Katanga), S and central Tanzania, intergrading with leuconotus. Remiges, primary coverts and greater coverts broadly edged bright olive-yellow (not red); rectrices edged dull olive-yellow or olive-green. Breeding Y differs from leuconotus in having cheeks and ear-coverts scarlet, black on head restricted to lores; scarlet of head and breast slightly more orange, typically extending further onto lower breast and often invading mantle; mantle to uppertailcoverts tinged olive. X has top of head to hindneck olive-yellow, rest of upperparts olive-grey; sides of head to chin, throat and breast pale olive yellow. Non-breeding Y like X but top of head typically deeper golden-brown, and crown and breast with a few orange feathers. A. r. jubaensis van Someren: S Somalia and adjacent coastal Kenya (Kiunga to Kiwayu). Y has head and body entirely bright red except for jet black outer edges of scapulars which form contrasting V-shaped line; wing and tail feathers blackish brown, broadly edged red. Apparently no non-breeding plumage. X like that of leuconotus.

Field Characters. Length 15–165 cm. A highly distinctive weaver of wooded savanna, with red or yellow wing and tail edgings in all plumages, orange-pink bill (brown in immature), much red in plumage of breeding YY, and clear white belly in all except breeding Y jubaensis. Subspecies differ considerably: YY of nominate rubriceps (southern Africa) have red head, breast and upper back and yellow wing and tail edgings; in northern race leuconotus they have black face patch, mainly black back, red wing and tail edgings; race jubaensis (S Somalia, NE Kenya) entirely red except for some black in wings and tail. XX and non-breeding YY are grey-brown with white belly, pale red wing and tail edgings (leuconotus, jubaensis) or yellowish on head and breast, with yellow feather edgings (rubriceps). Immatures similar to XX but head and breast washed yellowish in leuconotus. Insectivorous foraging habits draw attention to XX and non-breeding YY. Cardinal and Red-headed Queleas Quelea cardinalis and Q. erythrops are smaller and shorter-tailed with short, thick black bills, no red on back or in wings and tail. Voice. Tape-recorded (86, 88, 91, 104, B, F, GREG, McVIC). Song is a series of rapid, high-pitched squeaky swizzling notes ‘chu-tsee-tsi’, ‘chu-tsi’, ‘tsee-tsi’, ‘tswi-tsitswee’, ‘tzirrrr’, often running into a long nasal trill, ‘tchuthi-tseee-iiiiii-i’, ‘swizzzzzzz’ or ‘sizzi-sizzi-sizzi-sizzi’. Display calls of Y while hanging from nest, ‘ka-chi-kachi’ and ‘tsi-tsi-tswiiii-i’, in aggression ‘tswi-wee’. Contact calls while perched, ‘tswi-i-tsee’, ‘tsweei’, and twittering ‘tsi-tsi-tsi-tsi’; flight call ‘tsi’ or ‘tsi-whoo’; alarm, highpitched ‘chi-chi’. General Habits. Inhabits broad-leaved woodland, bushveld, miombo and acacia savanna in subtropical to tropical moist regions; gardens, especially on farms. Dry forests (Senegal); small forest patches (Ivory Coast); woodland savanna (Ghana); Isoberlinia doka woodland of N guinea savanna (Nigeria). Well-wooded savanna and open bush-

Anaplectes rubriceps land, up to 2100 m (Sudan). Below 2130 m in woodland, riverine woods, bushland, wooded grassland, and gardens, generally in somewhat dry areas (E Africa). Scattered treegrassland, and mixed woodland (Tanzania). In Somalia, woodland and forest edge, also open bush surrounding plains and along streams; lowlands in S but in N occurs at 1200–1800 m. Miombo and other woodlands (Angola). In Zambia, open woodland of all types (but rare in acacia); usually in thin miombo, but also mopane, Burkea, and other woodlands of similar structure; 330–1700 m. Below 1500 m, normally in canopy of any open woodland; sparse ˆ i). Broad-leaved and mixed woodlands below 330 m (Malaw in high rainfall regions and on alluvial soils (Botswana). Broad-leaved and acacia woodland; avoids forest and dense woodland (S Mozambique). Broad-leaved woodland in hot, relatively moist areas (South Africa). Usually solitary or in pairs, but also joins mixed bird parties. Forages mainly in trees, bushes and creepers; among foliage, on dry fruit capsules and along branches, often hanging upside down. Foraging height in trees 79 m (standard error 13, n ¼ 9, Kenya). Pecks at spider nests to extract spiders; hawks insects in air. Bathes in bird-baths. Quiet and easily overlooked. Frequents vicinity of old nests, which remain hanging for many months, and sometimes roosts in them. Prenuptial moult not complete (rump and belly feathers not moulted). Sometimes YY are in breeding plumage in winter in South Africa (Apr and July). In Ethiopia YY are in breeding plumage in every month except Jan and Aug, indicating irregularity in seasonality of moult (and thus breeding). Local movements between habitats occur seasonally in Botswana when some of its preferred deciduous habitat becomes inhospitable in dry season. In Zimbabwe range contracts in non-breeding season. Food. Insects, including beetles, grasshoppers, mantids, Hemiptera, termites; spiders (e.g. social spiders Stegodyphus); fruit (e.g. mistletoe Tapinanthus leendertziae, figs); seeds. Snails. Young fed on insects and spiders. Breeding Habits. Polygynous; sometimes monogamous. Usually only 1 Y in each colony; sometimes up to 9 nests, once 40 (Zimbabwe). Nest-building and courtship are spread out over several days. Between spells of building pair flies off to surrounding bush and remains for some time. When X inspects final nest, Y calls with wings quivering, while hanging from nest or nearby twigs. Occasionally Y flies into entrance tube and immediately flies out again. Copulation occurs near nest; Y sometimes pulls X out of nest to copulate (Benson 1946). NEST: rough, strong, retort-shaped, made of twigs (sometimes thorny), leaf midribs from Combretum and other trees, grass stems, tendrils and broad leaves. A ring is built first (A), then nest chamber; ceiling of leaves or acacia pods is added; often leaves on twig are intertwined at top of nest to provide additional waterproof covering. Nest not closely woven and appears untidy; length 160–180 from front to rear, height 140–180; vertical entrance spout 180–200 long is attached, whose many projecting ends give it untidy appearance (B); inside diam. of spout 60–70. Built

by Y, may be helped by X, also by imm. YY; sometimes a juv. accompanies the pair during initial building but does not help. When gathering nest material, strips midribs and twigs of their leaflets, tugs twigs until they break off, and nips off pliable twigs to knot them into the nest structure (Crook 1963). Alternate reverse winding is used in building (i.e. regularly looping strands back and forth), as in some Ploceus species (Collias and Collias 1964). Accepted nest is lined by X with dry grass, bark fibres, feathers or leaves. Placed 2–13 m above ground, suspended from ends of twigs in tree, sometimes hanging under the shady canopy, or below an old nest. Same site used each

99

100

PLOCEIDAE

(single records); S Mozambique, Sept–Oct; Namibia, Sept– Dec; Transvaal, Oct–Jan (peak Nov); Natal, Oct–Nov. INCUBATION: by both sexes, mostly by X; 61% of day spent incubating. Period: 11–13 days. DEVELOPMENT AND CARE OF YOUNG: young fed insects, by both adults, mostly by X (50 feeds per h) and helped by Y (27 feeds per h). Period c. 17 days (captivity). 2 young in one nest were of unequal size. Age at which crimson bill develops unknown. BREEDING SUCCESS/SURVIVAL: occasional host to Diederik Cuckoo Chysococcyx caprius. A snake in one nest ate ad. and eggs. Adult found struggling in a spider web (observer

year, and may include many nests from previous seasons. Site often conspicuous, e.g. overhanging a road, cattle kraal, small river, or on a steep hillside. Nest trees include Acacia albida and other acacias, mopane Colophospermum mopane, baobab Adansonia digitata, kiaat Pterocarpus angolensis, P. erinaceus, msasa Brachystegia spiciformis, mountain acacia B. glaucescens and other brachystegias, silver raisin Grewia monticola, marula Sclerocarya birrea and exotic eucalyptus trees. Foliage surrounding nest usually stripped off, as in some Ploceus spp. 1 nest was built in a bamboo. Man-made structures often used, e.g. windmill vanes, telephone wires (C) edges of thatched roofs of buildings, even inside buildings. Often nests in same tree as large raptor, especially Red-necked Buzzard Buteo auguralis, also African Hawk Eagle Hieraaetus spilogaster. Sometimes breeds near other weavers, e.g. Village Ploceus cucullatus, Southern Masked P. velatus, Lesser Masked P. intermedius and Red-billed Buffalo-Weaver Bubalornis niger. Old nests are often left hanging; the more dilapidated ones leave no trace of occupation and may be used for breeding by other bird species. Some nests may be multiple (D). EGGS: 1–4, usually 2–3; laid at 2-day intervals (Medland ˆ i (n ¼ 5) 1–4 (26); Zimbabwe (n ¼ 15) 2–4 1990); Malaw ˆ i (n ¼ 105) 1–4 (26); (26); Zimbabwe, Zambia, Malaw southern Africa (n ¼ 23) 2–3 (25). Oval; pale blue, often darker at thick end or clouded with darker blue. SIZE: (W and E Africa) av. c. 21  13; (Nigeria) 210  135; (n ¼ 3, Ethiopia) 205–22  145–147; (Ethiopia and Sudan, n ¼ 5) 204–222  143–150 (216  147); Zaı¨re: rubriceps 188–219  138–150; leuconotos 205–220  135– 147; (Zambia, n ¼ 2) 198–210  143–153; (Zimbabwe, n ¼ 39) 190–219  129–148 (205  140); (southern Africa, n ¼ 60) 187–219  129–149 (204  140). LAYING DATES: Ivory Coast, Jan–Feb; Ghana, Dec–Jan, Apr; Togo, Mar, Dec; Nigeria, Nov–May; Sudan, Aug– Oct; Ethiopia, Feb–Apr, Oct, Dec; Somalia, May, building nest Aug; E Africa: Kenya, Jan–Feb, Mar–Apr; Tanzania, Jan–Mar; Region B, Feb, May–July; Region C, Feb–Mar, May, Sept, Nov–Dec; Region D, Mar, Apr–June, Nov; Zaı¨re, Sept–Dec; Angola, Sept–Nov; Zambia, Aug–Nov ˆ i, Sept–Jan (peak Oct); Botswana, Sept– (peak Sept); Malaw Feb; Zimbabwe, July–Feb (mainly Aug–Dec), Mar–Apr

helped bird escape). Adults infected with Haemoproteus, Leucocytozoon, Trypanosoma and Microfilaria blood parasites and Hyalomma marginatum ticks. 1 bird recaptured after 5 years 8 months.

Key References

Joubert, E. (1985), Medland, R. D. (1990).

Ploceus

Genus Ploceus Cuvier

Small to medium-sized weavers. Sexes similar or dimorphic; some species with and others without seasonal plumage change. YY of most species predominantly yellow, often with black areas (mask, head, back, wings; YY of some forest species all black); YY usually with olive in plumage and sometimes brown or chestnut (never red); XX of most species olive-green and brown, generally stripe-backed. Eyes often yellow or red, colour varying with age and sex. Bill hard, conical, robust in most species, slender in a few. Strong birds; bite powerful; feet with strong grip. Nest woven, made of grass, sturdy, with or without a short entrance tunnel; nest always entered from below. Y display is focused at nest; commonly includes sizzling song and wing-beating whilst bird hanging upside down under nest. Y always builds nest, with or without active participation of X; Y may incubate and/or assist in feeding young, but in many species makes no contribution to parental care. Social organizations range from permanent monogamous pairs to colonial and highly polygynous species. Inhabit forest and savanna woodland; some species in marshland and papyrus; wide range of feeding habits – granivory, insectivory, omnivory. No shared morphological characters unite this large genus, and the diversity of skeletal characters, which tend to be conservative at a generic level (Craig 1999), suggests that Ploceus is not monophyletic and that further study will lead to the recognition of additional genera. A full revision is in progress, but for the present, based on morphological and biological characters, we keep the genus intact, except for removing superciliosus into Pachyphantes, q.v. Within the genus, several groups of species were recognized by Hall and Moreau (1970), some composing superspecies, others clustered as ‘speciesgroups’, yet others grouped quite speculatively. We recognize most of Hall and Moreau’s superspecies and a few of their species-groups, but at the present limited state of knowledge we cannot discern relationships between these groups or between most of the independent species. It is also hard to identify primitive and derived characters, as the basis for a rational species sequence (from ‘primitive’ to ‘advanced’). We therefore fall back, with minor changes, on the sequence of Dowsett and Forbes-Watson (1993). 63 species; 57 in Africa (all endemic except P. galbula of NE Africa and SW Arabia and P. manyar, introduced to Egypt), 5 Indo–Malayan (including P. manyar) and 2 Malagasy. Of the 57 African species, 33 are independent and 24 fall into 9 superspecies: baglafecht/bertrandi/nigrimentus; luteolus/pelzelni/subpersonatus; xanthops/subaureus; aurantius/castaneiceps/bojeri; castanops/xanthopterus; velatus/vitellinus/katangae/ruweti/reichardi; weynsi/golandi; rubiginosus/badius; and olivaceiceps/nicolli.

Ploceus baglafecht superspecies

Ploceus luteolus superspecies

1

1

1

3 1 P. baglafecht 2 P. bertrandi 3 P. nigrimentum

3

1

1 2 3,2

2 2

1

2

2

1 P. luteolus 2 P. pelzelni 3 P. subpersonatus

1,2 2

1 12

1,2

101

102

PLOCEIDAE

Ploceus xanthops superspecies

Ploceus aurantius superspecies

2

1 1 1 P. xanthops 2 P. subaureus

2 1

1 3

1 P. aurantius 2 P. bojeri 3 P. castaneiceps

2

2

Ploceus velatus superspecies

Ploceus castanops superspecies

1 1 3 1 P. castanops 2 P. xanthopterus

2

2

2

1 P. vitellinus 2 P. reichardi 3 P. katangae upembae 4 P. k. katangae 5 P. ruweti 6 P. velatus

4

5

6

2

1

Ploceus

Ploceus weynsi superspecies

1

Ploceus rubiginosus superspecies

1

1

2

2 1 P. weynsi 2 P. golandi

1 P. badius 2 P. rubiginosus

Ploceus olivaceiceps superspecies

1 1 P. olivaceiceps 2 P. nicolli

2 1

1

103

104

PLOCEIDAE

Plate 10

Ploceus flavipes (Chapin). Yellow-legged Weaver. Tisserin a` pieds jaunes.

(Opp. p. 139)

Malimbus flavipes Chapin, 1916. Bull. Am. Mus. Nat. Hist., 35, p. 27; Avakubi, Zaı¨re. Range and Status. Endemic resident, known only from Ituri forest, Zaı¨re, from Avakubi (01
240 N, 27
400 E) east to Simbo and Campi ya Wanbuti, south to Ukaika (0
360 N, 28
510 E) whence east to Lima, Tungudu and Makayobe (0
450 N, 29
440 E), a range of c. 100  200 km. Rare and vulnerable, considered endangered by forest clearance which is known to have begun in its range 20 years ago (Collar and Stuart 1985). None seen by Chapin (1954), known only by 9 specimens collected from 1910 to 1959 and 2 sightings in Okapi Faunal Res., Ituri, in 1980s (Collar et al. 1994). Description. ADULT Y: forehead to tail black, some gloss on crown, nape and mantle; underparts black except for belly and undertail-coverts which are blackish-brown. Bill black; eyes not described; legs and feet dull yellow. May assume X-like plumage in part of year (Louette 1988). ADULT X: like ad. Y, but a little less glossy, undertail-coverts duller. Bill black, eyes yellow, legs and feet dull yellow. Some specimens are blackish green dorsally, greenish grey ventrally; this may represent a non-breeding dress rather than imm. plumage (Louette 1988). SIZE: wing, Y 83, X 78–79; tail, Y 47, X 44–45; bill, Y 20, X 191–192; tarsus, Y 165, X 183–190. IMMATURE: crown to tail chocolate brown, throat and belly brown, wings chocolate brown; some birds blackish green dorsally, greenish grey ventrally (Louette 1988); faint oliveyellow collar on nape (Gyldenstolpe 1924); bill black, eyes yellowwhite, legs brown. SIZE: (2 YY, 3 XX) wing, Y 75–80, X 75–82; tail, Y 45–52, X 40–45; bill, Y 165–175, X 160–170; tarsus, Y 170–180, X 160–170 (Prigogine 1960). NESTLING: unknown. TAXONOMIC NOTE: placed in Malimbus by Chapin (1916) and Sibley and Monroe (1990); the latter remark only that ‘it seems closer to Malimbus than to Ploceus and perhaps represents a monotypic genus’; placed in monotypic genus Rhinoploceus by Gyldenstolpe (1924) and Wolters (1975–1982). Nostrils are large, exposing a peculiar turbinal tubercle.

Field Characters. Length 15 cm. Rare, restricted to lowland forest in E Zaı¨re. Black, with green gloss on tips of feathers of crown, back and breast giving slightly scaly effect (these areas wholly black in Vieillot’s and Maxwell’s Black Weavers P. nigerrimus and P. albinucha); belly and undertailcoverts dark brown. Sexes similar. Dull yellow legs and feet distinctive; further told from Maxwell’s by yellow eye. Y Vieillot’s has yellow eye but pale brown legs, wholly black underparts; lone Y could be difficult but presence of streaky olive and yellow X Vieillot’s will help identification.

Ploceus flavipes

Voice. Not known. General Habits. Very little-known. Occurs in canopy of dense lowland forest; not seen by J. P. Chapin, and only 9 known specimens. Since 1959 (specimen), only 2 sightings (Collar et al. 1994). Birds with wing-moult in Dec, Aug. Food. Caterpillars in 1 stomach. Breeding Habits. Nothing known except that X had enlarged ovary and egg in oviduct in Sept; another specimen from Sept described as in breeding condition; evidence from moult and ad. plumages that breeding may occur May–Sept (Louette 1988). Key References Collar, N. J. et al. (1994), Louette, M. (1988), Prigogine, A. (1960).

Plate 6

Ploceus bertrandi (Shelley). Bertram’s Weaver. Tisserin de Bertrand.

(Opp. p. 75)

^ Hyphantornis bertrandi Shelley, 1893. Ibis, 1893, p. 23, pl. 2; Zomba, Malawi. Forms a superspecies with P. baglafecht and P. nigrimentus. ˆ i, from Mulanje Range and Status. Endemic resident. Malaw northwards, widespread; NE Zambia, Nyika Plateau and nearby highlands, also SW corner of L. Tanganyika;

Mozambique, near Furancungo. Tanzania, local in southern highlands, rare in Udzungwa Mts, commoner in Uluguru and Ukaguru Mts; also in Ngurus (Britton 1980).

104

PLOCEIDAE

Plate 10

Ploceus flavipes (Chapin). Yellow-legged Weaver. Tisserin a` pieds jaunes.

(Opp. p. 139)

Malimbus flavipes Chapin, 1916. Bull. Am. Mus. Nat. Hist., 35, p. 27; Avakubi, Zaı¨re. Range and Status. Endemic resident, known only from Ituri forest, Zaı¨re, from Avakubi (01
240 N, 27
400 E) east to Simbo and Campi ya Wanbuti, south to Ukaika (0
360 N, 28
510 E) whence east to Lima, Tungudu and Makayobe (0
450 N, 29
440 E), a range of c. 100  200 km. Rare and vulnerable, considered endangered by forest clearance which is known to have begun in its range 20 years ago (Collar and Stuart 1985). None seen by Chapin (1954), known only by 9 specimens collected from 1910 to 1959 and 2 sightings in Okapi Faunal Res., Ituri, in 1980s (Collar et al. 1994). Description. ADULT Y: forehead to tail black, some gloss on crown, nape and mantle; underparts black except for belly and undertail-coverts which are blackish-brown. Bill black; eyes not described; legs and feet dull yellow. May assume X-like plumage in part of year (Louette 1988). ADULT X: like ad. Y, but a little less glossy, undertail-coverts duller. Bill black, eyes yellow, legs and feet dull yellow. Some specimens are blackish green dorsally, greenish grey ventrally; this may represent a non-breeding dress rather than imm. plumage (Louette 1988). SIZE: wing, Y 83, X 78–79; tail, Y 47, X 44–45; bill, Y 20, X 191–192; tarsus, Y 165, X 183–190. IMMATURE: crown to tail chocolate brown, throat and belly brown, wings chocolate brown; some birds blackish green dorsally, greenish grey ventrally (Louette 1988); faint oliveyellow collar on nape (Gyldenstolpe 1924); bill black, eyes yellowwhite, legs brown. SIZE: (2 YY, 3 XX) wing, Y 75–80, X 75–82; tail, Y 45–52, X 40–45; bill, Y 165–175, X 160–170; tarsus, Y 170–180, X 160–170 (Prigogine 1960). NESTLING: unknown. TAXONOMIC NOTE: placed in Malimbus by Chapin (1916) and Sibley and Monroe (1990); the latter remark only that ‘it seems closer to Malimbus than to Ploceus and perhaps represents a monotypic genus’; placed in monotypic genus Rhinoploceus by Gyldenstolpe (1924) and Wolters (1975–1982). Nostrils are large, exposing a peculiar turbinal tubercle.

Field Characters. Length 15 cm. Rare, restricted to lowland forest in E Zaı¨re. Black, with green gloss on tips of feathers of crown, back and breast giving slightly scaly effect (these areas wholly black in Vieillot’s and Maxwell’s Black Weavers P. nigerrimus and P. albinucha); belly and undertailcoverts dark brown. Sexes similar. Dull yellow legs and feet distinctive; further told from Maxwell’s by yellow eye. Y Vieillot’s has yellow eye but pale brown legs, wholly black underparts; lone Y could be difficult but presence of streaky olive and yellow X Vieillot’s will help identification.

Ploceus flavipes

Voice. Not known. General Habits. Very little-known. Occurs in canopy of dense lowland forest; not seen by J. P. Chapin, and only 9 known specimens. Since 1959 (specimen), only 2 sightings (Collar et al. 1994). Birds with wing-moult in Dec, Aug. Food. Caterpillars in 1 stomach. Breeding Habits. Nothing known except that X had enlarged ovary and egg in oviduct in Sept; another specimen from Sept described as in breeding condition; evidence from moult and ad. plumages that breeding may occur May–Sept (Louette 1988). Key References Collar, N. J. et al. (1994), Louette, M. (1988), Prigogine, A. (1960).

Plate 6

Ploceus bertrandi (Shelley). Bertram’s Weaver. Tisserin de Bertrand.

(Opp. p. 75)

^ Hyphantornis bertrandi Shelley, 1893. Ibis, 1893, p. 23, pl. 2; Zomba, Malawi. Forms a superspecies with P. baglafecht and P. nigrimentus. ˆ i, from Mulanje Range and Status. Endemic resident. Malaw northwards, widespread; NE Zambia, Nyika Plateau and nearby highlands, also SW corner of L. Tanganyika;

Mozambique, near Furancungo. Tanzania, local in southern highlands, rare in Udzungwa Mts, commoner in Uluguru and Ukaguru Mts; also in Ngurus (Britton 1980).

Ploceus bertrandi

Ploceus bertrandi

swizzzzzzz, swerrzzzzz, swizzzzzzz, titititititititi, dzerrrrrrti-dzerrrrrr, swizzzzzzzz . . . ’. Call a sparrow-like chirp. General Habits. Inhabits open woodland and bush along rivers, forest edges and cultivation in hilly country; at 900– 1800 m in Tanzania; trees and bracken-briar along stream lines above 1850 m in Zambia; only above 1000 m, up to ˆ i. Apparently present all year around 2000 m, in Malaw ˆ i in Mar–July, birds Blantyre. Post-juvenile moult in Malaw acquiring ad. plumage in Oct–Nov; ad. in moult during Sept (Lane 1995). In pairs or small groups of 5–6, gleaning insects from leaves. Food. Insects (Lane 1995).

Description. ADULT Y: forehead and crown chestnut-brown; narrow yellow band across hindcrown; nape golden-yellow with sickle-shaped black patch; mantle, rump and tail olive-green; lores, supercilium, cheeks and ear-coverts black; chin and throat black; breast golden yellow with some buffy wash; belly, thighs and undertail-coverts yellow, flanks with greenish wash; wings olive-green with narrow yellow edges to remiges, broader yellow fringes on coverts; underwing yellow. Bill black; eyes yellow; legs dark brown. No eclipse plumage. ADULT X: like Y, but forehead and crown black; edges of wing coverts greenish. SIZE (12 YY, 12 XX): wing, Y 80–86 (832), X 78–81 (796); tail, Y 56–63 (595), X 55–60 (578); bill, Y 200–230 (215), X 203–225 (214); ˆ i) tarsus, Y 213–233 (224), X 207–227 (220). WEIGHT: (Malaw Y (n ¼ 14) 359–420; X (n ¼ 15) 358–410 (Lane 1995). IMMATURE: sexes alike. Forehead, crown, lores, supercilium and ear-coverts olive-green; nape dull yellow; mantle, rump and tail olive-green; chin, throat, breast, belly, and undertail-coverts yellow, paling to white in centre of belly. Upper mandible dark horn, lower pinkish horn; eyes grey; legs brown. At first moult, head is mottled with black plumage, upper mandible becomes dark brown, lower mandible light brown, and eyes pale strawcoloured. NESTLING: no information.

Field Characters. Length 15 cm. Y distinctive, with chestnut crown, black nape patch separated from black face and throat by narrow yellow line; golden yellow below with saffron wash on neck and breast; X has head solid black to nape and upper throat; both sexes have yellow eyes, unstreaked upperparts. Juv. like X but black on head mixed with green and yellow. Sympatric forms of Baglafecht Weaver P. baglafecht also have yellow eyes in black mask, but throat yellow, upperparts streaked. Voice. Tape-recorded (102, B, CHA). Song a series of rather screechy swizzles lasting 8–12 s, starting with short squeaky phrase, then 3 swizzles, clicking notes, 3-syllable swizzle, and a final swizzle which tapers off: ‘p’tsi-chu,

Breeding Habits. Solitary nester, monogamous. NEST: ball made of broad grass strips, without any entrance tunnel (A), lined with grass seed heads. Resembles nest of P. xanthops, but rounder (Belcher 1924). Sited typically near tip of lower branch of camelthorn acacias, or in leafy end of evergreen tree branch, in tree usually near a stream. EGGS: 2, deep green, heavily blotched and spotted with ˆ i) av. 225  160 (Belcher 1924). red-brown. SIZE: (Malaw LAYING DATES: Tanzania, (breeding condition Apr); ˆ i, Aug–Nov (X in breeding condition Zambia, Dec; Malaw Oct, very young juvs Nov–Dec (Lane 1995); nest-building Feb (Johnston-Stewart 1982), nests active and adults feeding fledged young May (Benson and Benson 1977)). INCUBATION: no information. DEVELOPMENT AND CARE OF YOUNG: both Y and X feed young. BREEDING SUCCESS/SURVIVAL: birds ringed as adults ˆ i: recaptured after 25, 36 and 52 months (Blantyre, Malaw Lane 1995). Key References (1995).

Johnston-Stewart, N. G. B. (1982), Lane, S.

105

106

PLOCEIDAE

Plate 5

Ploceus baglafecht (Daudin). Baglafecht Weaver. Tisserin baglafecht.

(Opp. p. 74)

Loxia baglafecht Daudin, 1802. In Buffon, Hist. Nat. ´ed. Lace´pe`de, Quad. 14, p. 245; Abyssinia. Forms a superspecies with P. bertrandi and P. nigrimentus. Range and Status. Endemic resident. Nigeria, local on Mambilla Plateau, Gashaka-Gumti Nat. Park; Cameroon, c. 35 localities (M. Languy, pers. comm., 2002): from Bakossi in SW north to Hossere Vokre, Bamenda Highlands, Adamawa Plateau, isolated population at Mieri; Central African Republic, Ngoundji, Marali, Bogoin, north to Sangba; Zaı¨re, Bambili, L. Albert and Semliki valley, south to Ruwenzori, Beni, Kivu and Karagwe to Marungu plateau; Sudan, Boma hills, Imatong Mts, Issore, Li Rangu; Ethiopia, Dessa’a forest, Simen Mts Nat. Park, Hugumburda forest, L. Tana, Denkoro forest, Jemma R., Ankober, Finchara swamp, Mt Entoto, Jibat forest, Mugo highlands, Benga forest, Bale Mts Nat. Park, Aletta, Gidabo; Eritrea, Keren, Asmara and Senafe; Rwanda, Rugezi marsh, Volcans Nat. Park, Akagera Nat. Park, Nyabarongo, Akanyaru, Nyungwe forest; Burundi, L. Rwihinda, Kibira Nat. Park, Ruvubu Nat. Park, Ruizi Nat. Park; Uganda, NW east to Kitgum, Mt Lonyili, Mt Morongole, Busoga, W Elgon, Teso, S Karamoja; Kenya, Mt Elgon to Turkwell and Kerio R., Kakamega Forest, Eldoret, Maralal, Mt Marsabit, throughout highlands, Nguruman, Chyulu, Taita Hills; Tanzania, Crater and Mbulu Highlands, Arusha, Mt Meru, Kilimanjaro, Pare Mts, Usambaras, Irinja south to Mbeya and Ufipa, Tatanda, Mahari Mt, West Lake and Kigoma region; Zambia, Nyika Plateau and ˆ i, Nyika Uningi (Leonard and Van Daele 2001); Malaw Plateau. Common to abundant in most areas. Description. P. b. neumanni (Bannerman): E Nigeria, Cameroon, Central African Republic. ADULT Y (breeding): forehead and forecrown bright yellow, merging to olive on hindcrown and nape; lores, around eyes and ear-coverts black, the last bordered behind with semicircle of bright yellow, forming black mask surrounded by yellow; mantle and back bright green with illdefined grey shaft streaks, rump greyer, uppertail-coverts bright green; tail olive-green. Chin, throat and breast bright yellow, belly off-white, flanks whitish tinged with grey, thighs and undertail-coverts yellow. In wing, lesser coverts blackish, margined with bright green, median and greater coverts olive, margined bright green, primaries and secondaries brownish black with greenish yellow outer edges and pale yellow inner edges, tertials blackish with wide greenish yellow fringes; underwingcoverts pale yellow. Bill black, eyes cream-white, legs and feet flesh-brown. ADULT Y (non-breeding): forehead to nape olivegreen; lores, around eyes and ear-coverts olive-brown; mantle and back olive, heavily streaked with brown, rump greyer, uppertailcoverts olive; tail olive-brown; underparts pale brownish, flanks greyer, thighs and undertail-coverts pale yellow; wing-coverts olive and flight feathers dark brown, all edged pale yellowish. ADULT X (breeding): like breeding Y but mask olive-brown and forehead, forecrown and semicircle behind mask olive. ADULT X (non-breeding): like non-breeding Y. SIZE (4 XX, 3 XX): wing, Y 80–84, X 76–80; tail, Y 55–59, X 50–58; bill, Y 17–19, X 17–18; tarsus Y 21–23, X 21–23. WEIGHT: no data. IMMATURE: olive-brown upperparts, heavily streaked; face and crown dark olive (later head dark yellowish green, still with no mask); underparts buffy yellow. Bill horn-brown; eyes brown.

Ploceus baglafecht

NESTLING: newly-fledged young bird had greenish brown upperparts, tawny underparts; bill and legs pale grey. P. b. reichenowi (Fischer) (includes ‘nigrotemporalis’): Kenya, N Tanzania. ADULT Y: head like that of neumanni but hindcrown and nape black, sharply divided from bright yellow forecrown and sides of neck; mantle to uppertail-coverts black; tail blackish olive; wings black, feathers with yellow edges; underparts uniform golden yellow. Eyes bright yellow. No seasonal change: non-breeding same as breeding plumage. ADULT X: like Y, except that whole of head down to and including lores and ear-coverts black, in sharp contrast with yellow throat. SIZE: wing, Y (n ¼ 76) 79–86 (822), X (n ¼ 43) 77–83 (795); tail, Y (n ¼ 6) 56–59 (579), X (n ¼ 6) 55–60 (575); bill, Y (n ¼ 4) 168–207 (190), X (n ¼ 6) 160–206 (192); tarsus, Y (n ¼ 72) 258–293 (275), X (n ¼ 44) 260–280 (268) (Nairobi, Kenya; C. Jackson, pers. comm.). WEIGHT: (Kenya) Y (n ¼ 52) 240–375 (324), X (n ¼ 56) 265– 340 (299). P. b. baglafecht (includes ‘fricki’): Ethiopian highlands, W Eritrea. Breeding Y has yellow crown, black mask around eyes, greenish upperparts, yellow underparts paling to whitish on belly. X in breeding plumage like Y but forehead and crown greenish, mask around eyes dull black with green wash. Non-breeding birds of both sexes have ashy upperparts, and sides of head with buffy wash; dusky streaks in front of and behind eyes, and on mantle. Buff throat and breast, rest of underparts whitish. WEIGHT: (Ethiopia) Y (n ¼ 22) 260–358 (326), X (n ¼ 16) 285– 359 (310); unsexed (n ¼ 6) 26–31 (290). P. b. emini (Hartlaub) (includes: ‘budongoensis’) SE Sudan, N Uganda. Like baglafecht, breeding Y has yellow crown and black mask, but nape and mantle black, with green or grey edges to some mantle feathers; rump grey. Chin to breast golden yellow, belly to undertail-coverts white. Breeding X like Y, forehead and

Ploceus baglafecht crown black. Non-breeding plumage has nape and mantle grey with black streaks. WEIGHT: 1 Y (Sudan) 302. P. b. eremobius (Hartlaub): NE Zaı¨re, SW Sudan. Like baglafecht but smaller, underparts white from lower breast to undertailcoverts. Both sexes have breeding and non-breeding plumage. P. b. stuhlmanni (Reichenow): E Zaı¨re, S Uganda to W Tanzania. Black crown of breeding Y extends into face mask; upperparts from nape yellowish green, underparts ochre yellow. X has duller dark cap. No non-breeding plumage (Chapin 1954). Eye ochre in Y, light yellow in X (Hartert 1900). WEIGHT: 1 Y (Uganda) 32. P. b. sharpii (Shelley): highlands of SW Tanzania. Like stuhlmanni but breeding Y has green upperparts rather than yellowish green, underparts paler yellow. No non-breeding plumage. ˆ i. Dark P. b. nyikae Benson: Nyika plateau, Zambia and Malaw capped like stuhlmanni and sharpii, but apparently has a nonbreeding plumage (Chapin 1954). TAXONOMIC NOTE: visual differences among some of the subspecies are so striking that in the past at least 4 have been raised to the rank of full species. YY of neumanni, eremobius, emini, baglafecht and reichenowi have forehead and forecrown bright yellow, and of stuhlmanni, sharpii and nyikae, black. 2 races are black-backed (emini, reichenowi), all others green-backed. Some races have an eclipse plumage, others do not. However, most or all races intergrade (‘fricki’, ‘nigrotemporalis’ and ‘budongoensis’ are intergrades: Chapin 1954). No regional variations in voice are known, at least in E Africa (Stevenson and Fanshawe 2002). In Sudan, eremobius occurs in SW (Yei, Maridi, Li Rangu, Tambura), baglafecht in SE border areas (Boma, Imatong Mts: Nikolaus 1987), and emini commonly in Imatong, Dongotona and Didinga Mts, up to 2400 m, and in Yambio and Boma (Nikolaus 1989). It appears that baglafecht and emini overlap in Imatongs and on Boma Plateau, and van den Elzen and Ko ¨nig (1983) and Nikolaus (1989) treat them as separate species. But the 2 may be segregated altitudinally, and pending field studies we retain emini as a subspecies of P. baglafecht.

Field Characters. Length 14–15 cm. Extremely variable; yellow eye set in black face and yellow or green wing edgings common to all races and sexes. Race reichenowi, ‘Reichenow’s Weaver’, black above, bright yellow below, Y with black face mask surrounded by yellow, X with top of head black to below eye; in stuhlmanni, ‘Stuhlmann’s Weaver’, Y like X reichenowi, X like Y but top of head dark green; in emini, ‘Emin’s Weaver’, Y has yellow crown and black mask like reichenowi, but grey rump contrasts with black back and green tail, belly white; X has top of head black like reichenowi but back grey. Birds in W Africa different again, with white belly but green upperparts; face mask black in Y, narrowly surrounded by yellow, dark green in X. Races reichenowi, stuhlmanni and sharpii have no non-breeding plumage; in other races, both sexes ashy or brown above with dark streaking, pale brown or whitish below. Juvs generally resemble non-breeding birds; in races with no non-breeding plumage they are duller versions of ad. X, black on head replaced by dark green or dark brown. Voice. Tape-recorded (86, 102, 104, B, C, CHA, GREG, HOR, LEM, McVIC, PAR, ZIM). Song of X neumanni a high, thin lilting whistle ending with a few metallic cheeps; response to playback an aggressive snoring ‘snaarrrrl’, loud ‘jeep’, ‘chip’ and chattering ‘chichichichichi’. Call described as a distinctive repeated ‘zwenk’ (Borrow and Demey 2001). Song of nyikae short (c. 2 s) and varied, high sweet whistles and buzzy notes. Calls of reichenowi varied,

including dry ‘rink’ or ‘errink’, chirping ‘chweeeup’, chattering ‘chwi, chi-chi-chichit’, longer ‘sweeeeee . . . tchit’, and near nest a sharp ‘spi! spi! spi!’; song said to be nondescript and infrequent (Zimmerman et al. 1996); typical vocalization a chatter, ‘swii, chee chee cheechit’, and warning call ‘swii . . . chit’ (van Someren 1956). Songs and calls of different races not well studied, but no regional variation in E Africa. General Habits. Inhabits forest clearings and open vegetation outside forest: marshes, montane heathland, scrub and road verges, cultivation and gardens in cities such as Kampala and Nairobi; in Ethiopia, in montane and forest vegetation on forest fringes, apparently always above 1200 m; and in Zambia only above 1800 m. In W Cameroon in trees around rural villages, clearings in mountain forest and cleared patches of small farms, open grassy mountainsides dotted with shrubs; altitudinal range 1400–2285 m. Often in small parties, otherwise singly or in pairs, but never in large flocks. Aggressive towards other birds at bird-baths and food tables. Feeds on nectar by dipping bill into open flowers and swallowing droplets; probes clusters of dry leaves, searching for insects, gleans leaves and branches. In Usambara Mts, Tanzania, a bird followed a driver ant raid, picking insect prey off stems c. 1 m above ground (Willis 1986). 2 birds (Nairobi Nat. Park, Kenya) fed by clinging to stems of herb Leonotis nepetifolia, poking bills into dying flower heads, extracting small whitish objects (unripe seeds?), transferring one to foot holding stem and manipulating it there by constantly moving toes to adjust grip and improve object’s exposure, whilst pecking it into fragments which were swallowed (C.H. Fry, pers. comm.). Food. Mainly insects, also seeds and plant matter. Nectar of Prunus, Agave, Ruttya fruticosa, Grevillea robusta, Acrocarpus fraxinifolius, Leonotis nepetifolia. Takes fruits, including mango, banana and pawpaw at garden feeders; may damage seedlings and growing peas. Young fed spiders, moth larvae, grasshopper nymphs, small crickets, mantids, beetles, termite alates, winged Crematogaster ants. Breeding Habits. Solitary nester; monogamous. Sometimes 2 pairs nest in same tree. Y uses both long and short songs in courting X away from nest (Crook 1969). When not building nest, Y often perches near it and sings. One colour-marked pair in Kenya nested in Apr, again in Sept. NEST: (A) oval, lacking a spout (but short spout on 1 in Ethiopia), slightly flattened below, with entrance fairly well up one side, and an internal ledge to prevent eggs falling out; thick-walled, rather coarsely woven of strong strips from green grass blades or grass stems; lined, by both sexes, with grass seedheads, especially Pennisetum and Chloris, also a few feathers and plant down; lining may project as a porch over entrance. Suspended using tough grass stems, looped around branch above; attached to tips of branches or palm fronds (Sudan), built in outer edge of ˆ i), or attached to banana pine tree (Nyika Plateau, Malaw leaves (Kenya), with loops supporting nest threaded through leaf blades (van Someren 1956). One nest in Cameroon 25 m up in 2–3 stout grass stalks. Also uses

107

108

PLOCEIDAE

Eucalyptus trees, Pinus and Grevillea. SIZE: 175–205 long, 165–180 high, entrance 45–50  50–70; 18–10 m above ground. Successive nests may be built at same site, so that several present but only one used for breeding, although Y may roost in unlined nest. Does not strip leaves from around nest, which may be well concealed by dense foliage.

In a Nairobi garden, nest built alternately by 2 YY, with 1 X present (Collias 1978). X may select nest site but only Y builds, X accompanying him on trips to and from nest (Serle 1950); in captivity, X assisted Y to build nest. EGGS: 1–3, ovate to long ovate, slightly glossy and boldly marked, of 2 types: either blue-green, blotched and spotted with blackish brown especially towards larger end; or white to pinkish, evenly covered with reddish brown spots and blotches. In captivity, X laid on successive days between 11h00 and 17h00. SIZE: (n ¼ 5, Cameroon) 209– 225  144–154 (216  147); (n ¼ 12, Ethiopia) 200– 230  146–159 (212  151); also, 23  15–16 (Sudan, oval ‘white’ eggs), 225  160 (Uganda), 197–20  147 (Tanzania). WEIGHT: 051 g. LAYING DATES: Cameroon, Aug–Nov; Zaı ¨re, Kivu, Mar– June, Ituri, May, Uele, Aug–Oct (juvs July); Sudan, July, Nov; Ethiopia, Mar–Oct, possibly also Jan–Feb; Rwanda, Dec; Uganda, Dec–Jan, June–July on Mt Elgon, elsewhere Mar–Apr, June, Oct, Dec; Kenya, Nairobi, all months except Aug with peak in May, elsewhere peak Apr–June; Tanzania, Arusha, all months except Aug–Sept, Nov; Zambia, Dec (nest-building Nov, Feb) INCUBATION: by X only, Y often perching nearby. Period: 11–12 days; in captivity, av. period 15 days. DEVELOPMENT AND CARE OF YOUNG: initially only X feeds young, on small insects; from 4th day Y also feeds them. Nestling period 15–17 days, in captivity 17–19 days. Young fed for 2–3 weeks after fledging; once, new clutch laid 5 days after previous brood fledged, and Y took over care of fledglings. Adult plumage acquired at 13 months (Kenya: Jackson 2001). BREEDING SUCCESS/SURVIVAL: X ringed as adult, recaptured 2 years 10 months later (Urban 1975). Parasitized by Emerald Cuckoo Chrysococcyx cupreus (Kenya, van Someren 1956). Key References Rost, R. and Siebenrock, K.-H. (1992), Serle, W. (1950), van Someren, V. G. L. (1956).

Plate 5

Ploceus nigrimentus Reichenow. Black-chinned Weaver. Tisserin a ` menton noir.

(Opp. p. 74)

Ploceus nigrimentum Reichenow, 1904. Vo ¨gel Afrikas 3, p. 39; Galanga, Benguella, Angola. [For gender of nigrimentus see David and Gosselin 2002.] Forms a superspecies with P. baglafecht and P. bertrandi. Range and Status. Endemic resident. Gabon, Le´koni, Bate´ke´ Plateau. Congo, Bate´ke´ Plateau, Le´fini, BomboLumene and 40 km north of Brazzaville. Zaı¨re, Bankana, Bembo-Lunene. Angola, Bailundu highlands, Huambo and Cuanza Sul, last recorded 1960. Recent surveys indicate that habitat in Congo is secure and extensive, population estimated at >10,000 pairs (Stattersfield and Capper 2000). Description. ADULT Y: forehead and crown saffron-yellow; nape golden yellow; mantle black; rump and uppertail-coverts black, feathers edged with golden yellow; tail olive-green; lores, cheeks,

ear-coverts, chin and throat black; upper breast saffron-yellow; lower breast, belly, flanks, thighs and undertail-coverts golden yellow; wings black, with golden yellow fringes to coverts, reminges and tertials. Bill black; eyes cream or greenish yellow; legs brown. ADULT X: like Y, but forehead, crown and nape black and rump duller. SIZE: (3 YY, 2 XX: wing, Y 88–93, X 85, 88; tail, Y 59–64, X 61, 62; bill Y 214–226, X 220, 225; tarsus, Y 244–270, X 243, 247. IMMATURE: like X but black areas greenish grey, bill brown, eyes dark. NESTLING: no information. TAXONOMIC NOTE: placed in a superspecies with Baglafecht Weaver P. baglafecht and Bertram’s Weaver P. bertrandi by Hall and Moreau (1970), but affinities of all 3 species are uncertain.

108

PLOCEIDAE

Eucalyptus trees, Pinus and Grevillea. SIZE: 175–205 long, 165–180 high, entrance 45–50  50–70; 18–10 m above ground. Successive nests may be built at same site, so that several present but only one used for breeding, although Y may roost in unlined nest. Does not strip leaves from around nest, which may be well concealed by dense foliage.

In a Nairobi garden, nest built alternately by 2 YY, with 1 X present (Collias 1978). X may select nest site but only Y builds, X accompanying him on trips to and from nest (Serle 1950); in captivity, X assisted Y to build nest. EGGS: 1–3, ovate to long ovate, slightly glossy and boldly marked, of 2 types: either blue-green, blotched and spotted with blackish brown especially towards larger end; or white to pinkish, evenly covered with reddish brown spots and blotches. In captivity, X laid on successive days between 11h00 and 17h00. SIZE: (n ¼ 5, Cameroon) 209– 225  144–154 (216  147); (n ¼ 12, Ethiopia) 200– 230  146–159 (212  151); also, 23  15–16 (Sudan, oval ‘white’ eggs), 225  160 (Uganda), 197–20  147 (Tanzania). WEIGHT: 051 g. LAYING DATES: Cameroon, Aug–Nov; Zaı ¨re, Kivu, Mar– June, Ituri, May, Uele, Aug–Oct (juvs July); Sudan, July, Nov; Ethiopia, Mar–Oct, possibly also Jan–Feb; Rwanda, Dec; Uganda, Dec–Jan, June–July on Mt Elgon, elsewhere Mar–Apr, June, Oct, Dec; Kenya, Nairobi, all months except Aug with peak in May, elsewhere peak Apr–June; Tanzania, Arusha, all months except Aug–Sept, Nov; Zambia, Dec (nest-building Nov, Feb) INCUBATION: by X only, Y often perching nearby. Period: 11–12 days; in captivity, av. period 15 days. DEVELOPMENT AND CARE OF YOUNG: initially only X feeds young, on small insects; from 4th day Y also feeds them. Nestling period 15–17 days, in captivity 17–19 days. Young fed for 2–3 weeks after fledging; once, new clutch laid 5 days after previous brood fledged, and Y took over care of fledglings. Adult plumage acquired at 13 months (Kenya: Jackson 2001). BREEDING SUCCESS/SURVIVAL: X ringed as adult, recaptured 2 years 10 months later (Urban 1975). Parasitized by Emerald Cuckoo Chrysococcyx cupreus (Kenya, van Someren 1956). Key References Rost, R. and Siebenrock, K.-H. (1992), Serle, W. (1950), van Someren, V. G. L. (1956).

Plate 5

Ploceus nigrimentus Reichenow. Black-chinned Weaver. Tisserin a ` menton noir.

(Opp. p. 74)

Ploceus nigrimentum Reichenow, 1904. Vo ¨gel Afrikas 3, p. 39; Galanga, Benguella, Angola. [For gender of nigrimentus see David and Gosselin 2002.] Forms a superspecies with P. baglafecht and P. bertrandi. Range and Status. Endemic resident. Gabon, Le´koni, Bate´ke´ Plateau. Congo, Bate´ke´ Plateau, Le´fini, BomboLumene and 40 km north of Brazzaville. Zaı¨re, Bankana, Bembo-Lunene. Angola, Bailundu highlands, Huambo and Cuanza Sul, last recorded 1960. Recent surveys indicate that habitat in Congo is secure and extensive, population estimated at >10,000 pairs (Stattersfield and Capper 2000). Description. ADULT Y: forehead and crown saffron-yellow; nape golden yellow; mantle black; rump and uppertail-coverts black, feathers edged with golden yellow; tail olive-green; lores, cheeks,

ear-coverts, chin and throat black; upper breast saffron-yellow; lower breast, belly, flanks, thighs and undertail-coverts golden yellow; wings black, with golden yellow fringes to coverts, reminges and tertials. Bill black; eyes cream or greenish yellow; legs brown. ADULT X: like Y, but forehead, crown and nape black and rump duller. SIZE: (3 YY, 2 XX: wing, Y 88–93, X 85, 88; tail, Y 59–64, X 61, 62; bill Y 214–226, X 220, 225; tarsus, Y 244–270, X 243, 247. IMMATURE: like X but black areas greenish grey, bill brown, eyes dark. NESTLING: no information. TAXONOMIC NOTE: placed in a superspecies with Baglafecht Weaver P. baglafecht and Bertram’s Weaver P. bertrandi by Hall and Moreau (1970), but affinities of all 3 species are uncertain.

Ploceus bannermani

Ploceus nigrimentum

109

with yellow underparts. Y resembles ‘Reichenow’s Weaver’ P. baglafecht reichenowi of E Africa, with black back, bright yellow wing edgings and yellow eye in black face mask, but entire top of head to nape yellow, and black mask extends onto throat; X like Y but top of head black. Compact Weaver P. superciliosus has black nape, no yellow in wings. Voice. Tape-recorded (LEM). Songs rather short, distinctive ‘whit-pu-pui-trrrr-pui’; harsh ‘chi-chi-cheep’ and dry ‘jijiji’. General Habits. Inhabits open savanna with trees and bushes dominated by Hymenocardia acida up to 6–8 m tall, often near gallery forest, at 500–700 m on Bate´ke´ Plateau but above 1500 m in Bailundu highlands. In pairs or small groups. Shy; keeps to tops of low trees. Moults in May– June. Food. Grass seeds and insects. Breeding Habits. Semi-colonial; no further information.

Field Characters. Length 17 cm. A large, dark-backed weaver with a yellow rump; black chin and throat contrast

Key References Collar, N. J. et al. (1994), Demey, R. et al. (2000), Stattersfield, A. and Capper, D. (2000).

Ploceus bannermani Chapin. Bannerman’s Weaver. Tisserin de Bannerman.

Plate 5

Ploceus bannermani Chapin, 1932. Amer. Mus. Novit. no. 570, p. 17; Djang district, Cameroon.

(Opp. p. 74)

Range and Status. Endemic resident, montane areas of W Cameroon and extreme E Nigeria. Cameroon, Bakossi and Manenguba Mts at 1950–2250 m (from Kodmin and Ninong to near Nkongsamba); locally common in Bamboutos Mts and Bamenda Highlands at 1700–2600 m (Dschang, BafutNgemba Forest Res., L. Bambili, L. Bambulue, Sabga Pass, Bali-Ngemba Forest Res., Mbengwi, Tinachong); Jauro Masali at 1400 m on Nigerian border near Mambilla Plateau; Wum Crater Lake at 1100 m, and Mt Oku at 2100–2850 m; and Mt Tchabal Mbaba (Smith and McNiven 1993). Nigeria, frequent on Obudu Plateau with up to 12 seen daily, and common on Mambilla Plateau with up to 40 seen daily: Gangirwal at 2000–2300 m, Chappal Waddi, Leinde Fadali at 1400 m, Gotel Mts and Gashaka-Gumti Nat. Park. Vulnerable; a major conservation programme at Kilum on Mt Oku (Collar et al. 1994). Description. ADULT Y: forehead and crown golden yellow, passing into greenish on nape; mantle, rump and tail olive-green; chin, lores, sides of head to just above eye and ear-coverts black, forming a sharply defined mask extending onto throat; breast golden yellow; flanks, belly, thighs yellow with greenish wash; undertail-coverts golden yellow; wings dull brown with olivegreen fringes on feathers, broad on wing-coverts. Bill black; eyes cream; legs dark brown. Sexes alike. SIZE (5 YY, 8 XX): wing, Y 775–80 (786), X 74–80 (762); tail, Y 565–590 (58), X 53–59 (554); bill, Y 180–196 (187), X 185–202 (192); tarsus, (1 Y)

Ploceus bannermani

Ploceus bannermani

Ploceus nigrimentum

109

with yellow underparts. Y resembles ‘Reichenow’s Weaver’ P. baglafecht reichenowi of E Africa, with black back, bright yellow wing edgings and yellow eye in black face mask, but entire top of head to nape yellow, and black mask extends onto throat; X like Y but top of head black. Compact Weaver P. superciliosus has black nape, no yellow in wings. Voice. Tape-recorded (LEM). Songs rather short, distinctive ‘whit-pu-pui-trrrr-pui’; harsh ‘chi-chi-cheep’ and dry ‘jijiji’. General Habits. Inhabits open savanna with trees and bushes dominated by Hymenocardia acida up to 6–8 m tall, often near gallery forest, at 500–700 m on Bate´ke´ Plateau but above 1500 m in Bailundu highlands. In pairs or small groups. Shy; keeps to tops of low trees. Moults in May– June. Food. Grass seeds and insects. Breeding Habits. Semi-colonial; no further information.

Field Characters. Length 17 cm. A large, dark-backed weaver with a yellow rump; black chin and throat contrast

Key References Collar, N. J. et al. (1994), Demey, R. et al. (2000), Stattersfield, A. and Capper, D. (2000).

Ploceus bannermani Chapin. Bannerman’s Weaver. Tisserin de Bannerman.

Plate 5

Ploceus bannermani Chapin, 1932. Amer. Mus. Novit. no. 570, p. 17; Djang district, Cameroon.

(Opp. p. 74)

Range and Status. Endemic resident, montane areas of W Cameroon and extreme E Nigeria. Cameroon, Bakossi and Manenguba Mts at 1950–2250 m (from Kodmin and Ninong to near Nkongsamba); locally common in Bamboutos Mts and Bamenda Highlands at 1700–2600 m (Dschang, BafutNgemba Forest Res., L. Bambili, L. Bambulue, Sabga Pass, Bali-Ngemba Forest Res., Mbengwi, Tinachong); Jauro Masali at 1400 m on Nigerian border near Mambilla Plateau; Wum Crater Lake at 1100 m, and Mt Oku at 2100–2850 m; and Mt Tchabal Mbaba (Smith and McNiven 1993). Nigeria, frequent on Obudu Plateau with up to 12 seen daily, and common on Mambilla Plateau with up to 40 seen daily: Gangirwal at 2000–2300 m, Chappal Waddi, Leinde Fadali at 1400 m, Gotel Mts and Gashaka-Gumti Nat. Park. Vulnerable; a major conservation programme at Kilum on Mt Oku (Collar et al. 1994). Description. ADULT Y: forehead and crown golden yellow, passing into greenish on nape; mantle, rump and tail olive-green; chin, lores, sides of head to just above eye and ear-coverts black, forming a sharply defined mask extending onto throat; breast golden yellow; flanks, belly, thighs yellow with greenish wash; undertail-coverts golden yellow; wings dull brown with olivegreen fringes on feathers, broad on wing-coverts. Bill black; eyes cream; legs dark brown. Sexes alike. SIZE (5 YY, 8 XX): wing, Y 775–80 (786), X 74–80 (762); tail, Y 565–590 (58), X 53–59 (554); bill, Y 180–196 (187), X 185–202 (192); tarsus, (1 Y)

Ploceus bannermani

110

PLOCEIDAE

234, (5 XX) 222–230 (227). WEIGHT: Cameroon (unsexed, n ¼ 8) 290–332 (310). IMMATURE and NESTLING: undescribed.

Field Characters. Length 14 cm. Both sexes golden yellow with green upperparts and pale eye in small black face mask; nothing like it in montane forest habitat within its restricted range. Voice. Tape-recorded (104, B, LEM). Calls short and sharp with dry reedy quality, ‘kik’, ‘kook’, ‘chooko’; jerky, 3–4 s territorial song starts with these and accelerates slightly into faster 3-syllable calls, ‘chop-kik-kik-ko, choo-ko, chokwydo-kweedip’, ‘chop-kik-kik-ko-ko-chokwydokwideekwo-chip’. Also said to give a soft ‘chi-chi-chi-chichirrr-chirrr’ (Eisentraut 1973). General Habits. Occupies forest edge and dense shrubby habitat in more open sectors, including clearings, of montane forest at 1100–2850 m; along strips of forest in deep ravines of Obudu Plateau, and in secondary scrub.

Absent from wetter mountains in Cameroon south of 5
N. Usually in pairs, sometimes in small parties. Wing-moult in 1 bird in Mar. Food. No information. Breeding Habits. Apparently monogamous, solitary nests. NEST: retort-shaped with entrance pointing downwards; no tunnel. Outer wall compactly woven with grass stems and thin grass leaf blades, sparsely lined with grass husks and soft plant down; attached to thin branches of thorn bushes, 2–3 m above ground. EGGS: clutch reported as 2; apparently not described. LAYING DATES: Cameroon, Feb; breeding activity Nov–Feb; young accompanying adults in Feb. No other information. Key References Collar, N. J. and Stuart, S. N. (1985), Stattersfield, A. and Capper, D. (2000), Stuart, S. N. and Jensen, F. P. (1986).

Plate 6

Ploceus batesi (Sharpe). Bates’s Weaver. Tisserin de Bates.

(Opp. p. 75)

Othyphantes batesi Sharpe, 1908. Ibis, p. 348; Dja R, Cameroon. Range and Status. Endemic resident, S Cameroon. Rare; known from 9 localities: Mt Kupe´ (2 seen in 1990 but none since, despite intensive searching), Limbe (Victoria) (1 seen 1979), Tissongo (3
350 N, 9
540 E, 1 and 2 seen in 1978), Lolodorf (3
140 N, 10
440 E, 1 in early 1940s, 1 in 1949), Sangme´lima (2
560 N, 11
590 E, 1 in early 1940s), Bitye (3
010 N, 12
220 E, 4 specimens in 1909), Kumangola (3
140 N, 12
230 E, 1 in 1908), near Dja R. (type specimen, 1906), and Moloudou (2
020 N, 15
130 E, on Dja R. on Congo border). Description. ADULT Y: forehead and crown, cheeks and earcoverts bright chestnut-brown; lores black; sharp line on nape with feathers with yellow fringes, then dark olive-green; mantle, rump and tail olive-green; wings and wing-coverts uniform dark olive-green, paler outer edges to primary feathers; underwing greenish; chin chestnut between rami, otherwise black; throat black, ending in a sharp line; breast bright yellow, flanks yellow with some green wash, belly, thighs and undertail-coverts yellow. Bill black; eyes dark brown; legs blue-grey. Probably no seasonal change in plumage. ADULT X: like Y except that forehead, crown, cheeks and ear-coverts to level of lower mandible black; chin and throat yellow like breast. Bill black; eyes brown; legs blue-grey. SIZE (2 YY, 5 XX): wing, Y 73, 75, X 73–76 (74); tail, Y 45, 52, X 48–52 (49); bill, Y 188, 190, X 171–189 (178); tarsus, Y 184, 187, X 167–177 (172). IMMATURE: like X but head pale olive-green, not black; bill pale horn brown. NESTLING: not described.

Field Characters. Length 14 cm. Y green above, yellow below, with unique head pattern – rich chestnut crown and face, black throat; X has black crown and face, yellow throat.

Ploceus batesi

Voice. Unknown. General Habits. In lowland rainforest, up to 900 m on Mt Kupe´. Single birds, pairs, and once in a mixed-species flock of insectivorous birds. One near Limbe was in patch of dense forest on small, rocky river overhung by large trees and creepers; it moved in zig-zags up a creeper-covered

110

PLOCEIDAE

234, (5 XX) 222–230 (227). WEIGHT: Cameroon (unsexed, n ¼ 8) 290–332 (310). IMMATURE and NESTLING: undescribed.

Field Characters. Length 14 cm. Both sexes golden yellow with green upperparts and pale eye in small black face mask; nothing like it in montane forest habitat within its restricted range. Voice. Tape-recorded (104, B, LEM). Calls short and sharp with dry reedy quality, ‘kik’, ‘kook’, ‘chooko’; jerky, 3–4 s territorial song starts with these and accelerates slightly into faster 3-syllable calls, ‘chop-kik-kik-ko, choo-ko, chokwydo-kweedip’, ‘chop-kik-kik-ko-ko-chokwydokwideekwo-chip’. Also said to give a soft ‘chi-chi-chi-chichirrr-chirrr’ (Eisentraut 1973). General Habits. Occupies forest edge and dense shrubby habitat in more open sectors, including clearings, of montane forest at 1100–2850 m; along strips of forest in deep ravines of Obudu Plateau, and in secondary scrub.

Absent from wetter mountains in Cameroon south of 5
N. Usually in pairs, sometimes in small parties. Wing-moult in 1 bird in Mar. Food. No information. Breeding Habits. Apparently monogamous, solitary nests. NEST: retort-shaped with entrance pointing downwards; no tunnel. Outer wall compactly woven with grass stems and thin grass leaf blades, sparsely lined with grass husks and soft plant down; attached to thin branches of thorn bushes, 2–3 m above ground. EGGS: clutch reported as 2; apparently not described. LAYING DATES: Cameroon, Feb; breeding activity Nov–Feb; young accompanying adults in Feb. No other information. Key References Collar, N. J. and Stuart, S. N. (1985), Stattersfield, A. and Capper, D. (2000), Stuart, S. N. and Jensen, F. P. (1986).

Plate 6

Ploceus batesi (Sharpe). Bates’s Weaver. Tisserin de Bates.

(Opp. p. 75)

Othyphantes batesi Sharpe, 1908. Ibis, p. 348; Dja R, Cameroon. Range and Status. Endemic resident, S Cameroon. Rare; known from 9 localities: Mt Kupe´ (2 seen in 1990 but none since, despite intensive searching), Limbe (Victoria) (1 seen 1979), Tissongo (3
350 N, 9
540 E, 1 and 2 seen in 1978), Lolodorf (3
140 N, 10
440 E, 1 in early 1940s, 1 in 1949), Sangme´lima (2
560 N, 11
590 E, 1 in early 1940s), Bitye (3
010 N, 12
220 E, 4 specimens in 1909), Kumangola (3
140 N, 12
230 E, 1 in 1908), near Dja R. (type specimen, 1906), and Moloudou (2
020 N, 15
130 E, on Dja R. on Congo border). Description. ADULT Y: forehead and crown, cheeks and earcoverts bright chestnut-brown; lores black; sharp line on nape with feathers with yellow fringes, then dark olive-green; mantle, rump and tail olive-green; wings and wing-coverts uniform dark olive-green, paler outer edges to primary feathers; underwing greenish; chin chestnut between rami, otherwise black; throat black, ending in a sharp line; breast bright yellow, flanks yellow with some green wash, belly, thighs and undertail-coverts yellow. Bill black; eyes dark brown; legs blue-grey. Probably no seasonal change in plumage. ADULT X: like Y except that forehead, crown, cheeks and ear-coverts to level of lower mandible black; chin and throat yellow like breast. Bill black; eyes brown; legs blue-grey. SIZE (2 YY, 5 XX): wing, Y 73, 75, X 73–76 (74); tail, Y 45, 52, X 48–52 (49); bill, Y 188, 190, X 171–189 (178); tarsus, Y 184, 187, X 167–177 (172). IMMATURE: like X but head pale olive-green, not black; bill pale horn brown. NESTLING: not described.

Field Characters. Length 14 cm. Y green above, yellow below, with unique head pattern – rich chestnut crown and face, black throat; X has black crown and face, yellow throat.

Ploceus batesi

Voice. Unknown. General Habits. In lowland rainforest, up to 900 m on Mt Kupe´. Single birds, pairs, and once in a mixed-species flock of insectivorous birds. One near Limbe was in patch of dense forest on small, rocky river overhung by large trees and creepers; it moved in zig-zags up a creeper-covered

Ploceus luteolus tree-trunk, in manner strongly reminiscent of Bar-winged Weaver Ploceus angolensis (Taylor 1981). Food. Insects in stomachs.

111

Breeding Habits. No information; juv. in Mar. Key References Collar, N. J. and Stuart, S. N. (1985), Stattersfield, A. and Capper, D. (2000), Taylor, P. B. (1981).

Ploceus luteolus (Lichtenstein). Little Weaver. Tisserin minule.

Plate 6

Fringilla luteola Lichtenstein, 1823. Verz. Doubl. p. 23; Senegal.

(Opp. p. 75)

NOMENCLATURAL NOTE:

also called Slender-billed Weaver (e.g. Serle 1943, Morel 1977), now the name of P. pelzelni.

Forms a superspecies with P. pelzelni and P. subpersonatus. Range and Status. Endemic resident; common. Mauritania, uncommon south of 17
N, may extend north to Nouakchott during rains. Senegal, Gambia, resident throughout. Mali, Boucle de Baoule´, along Niger R. to 17
N at Timbuktu, some movement to north during rains. Burkina Faso, in E at Arli, Yilou, Ouagadougou, north to Ouahigouva. Ivory Coast, local as at Korgho, Ferke´sse´dougou. Ghana, Mole, Tamale, Bolgatanga, Lawra, seasonal at Tumu. Togo, along rivers in northern savannas. Benin, Be´te´rou and savanna areas. Nigeria, northern savanna, south of rivers in Borgu Game Reserve. Niger, Niamey, Gotheye, Tahoua, Tessaoua, Zinder, Agades, Aı¨r. Cameroon, Benue Plain and N districts, south to Majo Lifi. Chad, south of L. Chad on Cameroon border. Central African Republic, Manovo-Gounda-Saint Floris Nat. Park, Bamingui-Bangoran Nat. Park. Sudan, from Atbara south along Nile R. and tributaries. Ethiopia, W, NE, SE and S regions. Eritrea, in W. Zaı¨re, Yakoma to Rutshuru in NE. Uganda, south to Masaka and Ruwenzori. Kenya, L. Victoria basin to Muhoroni, Rapogi, Turkana to Kongelai, Kerio, L. Baringo and L. Bogoria. Tanzania, old record from Biharamulo, recently from Serengeti near Kenya border. Description. ADULT Y (breeding): forehead, crown to just behind eyes, lores, cheeks and ear-coverts black; hindcrown and nape bright yellow; mantle dull yellow, feathers with faint central streaks; rump bright yellow; tail olive-brown; chin and throat black, extending onto breast as a bib; breast surrounding bib bright yellow; belly, flanks, thighs and undertail-coverts bright yellow; wings olive-brown with pale yellow edges to remiges, broader margins on wing-coverts; underwing grey. Bill black; eyes hazel-brown; legs blue-grey. ADULT Y (non-breeding): forehead, crown, nape and mantle yellowish green, mantle feathers with dark central streaks; pale buff superciliary stripe; rump yellower than rest of upperparts; tail and wings as in breeding Y; underparts from chin to belly washed with pale yellow; flanks and thighs buffy; undertail-coverts yellow. Bill brown; eyes hazel-brown; legs blue-grey. ADULT X: like nonbreeding Y, but eyes dark brown; yellow underparts when breeding, whitish in non-breeding plumage. Bill dark brown when breeding, light brown afterwards. SIZE (12 YY, 10 XX): wing, Y 59–65 (617), X 58–63 (606); tail, Y 37–41 (391), X 35– 40 (382); bill, Y 130–146 (141), X 127–143 (135); tarsus, Y 150–173 (161), X 150–165 (157). WEIGHT: Y (Ghana) 130, 140, 150; (Nigeria, Apr–June, n ¼ 10) 130–149 (138); (Kenya, n ¼ 9) 125–146 (137); (Cameroon) 125, 150; X (Nigeria, n ¼ 4) 119–135 (127); (Kenya, n ¼ 7) 112–148 (131); (Cameroon) 130, 130; unsexed (Ghana) 125, 129, 13, 135; (Nigeria)

Ploceus luteolus

?

? ?

(Oct–Mar, n ¼ 28) 115–147 (130) (Apr–June, n ¼ 14) 119–149 (135); (Kenya) 135 IMMATURE: like non-breeding X, but upperparts grey-brown rather than greenish yellow; bill brown, with pale lower mandible. NESTLING: sparse, whitish natal down (Chapin 1954). TAXONOMIC NOTE: birds from Uganda, W Kenya and N Tanzania have been separated as P. 1. kavirondensis on basis of greener upperparts with more pronounced streaking, less yellow on nape.

Field Characters. Length 11 cm. Smallest weaver. Y has black forecrown, face and throat surrounded by bright yellow, without any of the saffron wash typical of larger weavers; separable from Slender-billed Weaver P. pelzelni where they meet, e.g. at L. Kanyaboli, W Kenya, by dry woodland habitat, much shorter bill (14 cm vs 17 cm), from Lesser Masked Weaver P. intermedius by brown, not white eye.

Ploceus luteolus tree-trunk, in manner strongly reminiscent of Bar-winged Weaver Ploceus angolensis (Taylor 1981). Food. Insects in stomachs.

111

Breeding Habits. No information; juv. in Mar. Key References Collar, N. J. and Stuart, S. N. (1985), Stattersfield, A. and Capper, D. (2000), Taylor, P. B. (1981).

Ploceus luteolus (Lichtenstein). Little Weaver. Tisserin minule.

Plate 6

Fringilla luteola Lichtenstein, 1823. Verz. Doubl. p. 23; Senegal.

(Opp. p. 75)

NOMENCLATURAL NOTE:

also called Slender-billed Weaver (e.g. Serle 1943, Morel 1977), now the name of P. pelzelni.

Forms a superspecies with P. pelzelni and P. subpersonatus. Range and Status. Endemic resident; common. Mauritania, uncommon south of 17
N, may extend north to Nouakchott during rains. Senegal, Gambia, resident throughout. Mali, Boucle de Baoule´, along Niger R. to 17
N at Timbuktu, some movement to north during rains. Burkina Faso, in E at Arli, Yilou, Ouagadougou, north to Ouahigouva. Ivory Coast, local as at Korgho, Ferke´sse´dougou. Ghana, Mole, Tamale, Bolgatanga, Lawra, seasonal at Tumu. Togo, along rivers in northern savannas. Benin, Be´te´rou and savanna areas. Nigeria, northern savanna, south of rivers in Borgu Game Reserve. Niger, Niamey, Gotheye, Tahoua, Tessaoua, Zinder, Agades, Aı¨r. Cameroon, Benue Plain and N districts, south to Majo Lifi. Chad, south of L. Chad on Cameroon border. Central African Republic, Manovo-Gounda-Saint Floris Nat. Park, Bamingui-Bangoran Nat. Park. Sudan, from Atbara south along Nile R. and tributaries. Ethiopia, W, NE, SE and S regions. Eritrea, in W. Zaı¨re, Yakoma to Rutshuru in NE. Uganda, south to Masaka and Ruwenzori. Kenya, L. Victoria basin to Muhoroni, Rapogi, Turkana to Kongelai, Kerio, L. Baringo and L. Bogoria. Tanzania, old record from Biharamulo, recently from Serengeti near Kenya border. Description. ADULT Y (breeding): forehead, crown to just behind eyes, lores, cheeks and ear-coverts black; hindcrown and nape bright yellow; mantle dull yellow, feathers with faint central streaks; rump bright yellow; tail olive-brown; chin and throat black, extending onto breast as a bib; breast surrounding bib bright yellow; belly, flanks, thighs and undertail-coverts bright yellow; wings olive-brown with pale yellow edges to remiges, broader margins on wing-coverts; underwing grey. Bill black; eyes hazel-brown; legs blue-grey. ADULT Y (non-breeding): forehead, crown, nape and mantle yellowish green, mantle feathers with dark central streaks; pale buff superciliary stripe; rump yellower than rest of upperparts; tail and wings as in breeding Y; underparts from chin to belly washed with pale yellow; flanks and thighs buffy; undertail-coverts yellow. Bill brown; eyes hazel-brown; legs blue-grey. ADULT X: like nonbreeding Y, but eyes dark brown; yellow underparts when breeding, whitish in non-breeding plumage. Bill dark brown when breeding, light brown afterwards. SIZE (12 YY, 10 XX): wing, Y 59–65 (617), X 58–63 (606); tail, Y 37–41 (391), X 35– 40 (382); bill, Y 130–146 (141), X 127–143 (135); tarsus, Y 150–173 (161), X 150–165 (157). WEIGHT: Y (Ghana) 130, 140, 150; (Nigeria, Apr–June, n ¼ 10) 130–149 (138); (Kenya, n ¼ 9) 125–146 (137); (Cameroon) 125, 150; X (Nigeria, n ¼ 4) 119–135 (127); (Kenya, n ¼ 7) 112–148 (131); (Cameroon) 130, 130; unsexed (Ghana) 125, 129, 13, 135; (Nigeria)

Ploceus luteolus

?

? ?

(Oct–Mar, n ¼ 28) 115–147 (130) (Apr–June, n ¼ 14) 119–149 (135); (Kenya) 135 IMMATURE: like non-breeding X, but upperparts grey-brown rather than greenish yellow; bill brown, with pale lower mandible. NESTLING: sparse, whitish natal down (Chapin 1954). TAXONOMIC NOTE: birds from Uganda, W Kenya and N Tanzania have been separated as P. 1. kavirondensis on basis of greener upperparts with more pronounced streaking, less yellow on nape.

Field Characters. Length 11 cm. Smallest weaver. Y has black forecrown, face and throat surrounded by bright yellow, without any of the saffron wash typical of larger weavers; separable from Slender-billed Weaver P. pelzelni where they meet, e.g. at L. Kanyaboli, W Kenya, by dry woodland habitat, much shorter bill (14 cm vs 17 cm), from Lesser Masked Weaver P. intermedius by brown, not white eye.

112

PLOCEIDAE

Voice. Tape-recorded (104, 109, B, GREG, McVIC, MOR, PAY). Song a pleasant continuum of chuckling bulbul-like notes, ‘chip-wop-weep-wop-wop-weep-wip-wop-cheepwop-wip . . . ’, quieter than other weavers. Calls mainly scratchy and grating, with weaver swizzles and chatters, and a soft ‘tsip’ or ‘tsssp’; alarm calls shrill and squeaky. General Habits. Inhabits savanna woodland with large Acacia trees, open scrubland with small trees, edges of farmland and large gardens, arid thornscrub and red sandy country (Kordofan, Sudan). In E Africa at 400–1500 m. Not gregarious; usually in pairs, occasionally singly or in small flocks (family parties?). Forages mainly in acacias and other small trees, working methodically through foliage, searching carefully, slightly warbler-like. Sometimes feeds on ground, in dry season may forage with waxbills (Gambia). Resident, but may expand north in Mauritania in rainy season; in N Nigeria, common at all seasons. Post-nuptial moult in Mali in Sept-Oct (Paludan 1936) and in Nigeria started by Oct (Fry 1971). Food. Seeds and insects; 4 stomachs contained only seeds, 6 had insects, including small beetles and caterpillars (Chapin 1954). Breeding Habits. Solitary nester (rarely, in colonies); monogamous. Pair often returns to same site in successive years. Y sings to X from a perch or hanging from nest, but does not tremble wings when singing. NEST: globular structure with short, vertical entrance tube at side (A), densely woven using fine, narrow herb and grass stems, grass blades, and small vine tendrils (Collias and Collias 1964). Strips torn from fronds of fan palm Borassus flabellifer often used (Mali: Lamarche 1981). Lined with grass seedheads and other fine material. Size, 100 long by 80 high (Collias and Collias 1964). Length of tunnel varies from 50 to 300 with int. diam. 25; 3–55 m above ground, suspended from thorny branches of small tree. Nests often built in Ziziphus and ‘heglig’ trees within 50 cm of occupied wasp nests (Sudan: Lynes 1924–1925); associated with hornets in Gambia (Barlow et al. 1997), but not in Zaı¨re. Pair seems to select thorny trees (Lynes 1924–1925, Moreau 1943). Nest construction takes 3–4

days, after which Y may continue work on tunnel, and tunnel may be added to nest long after breeding has ended (Lippens and Wille 1976). EGGS: 2–3. Plain white, ovate, slightly glossy. SIZE: (n ¼ 9, Nigeria) 175–206  125–139 (184  131,); (Cameroon) 165–190  125–130; (n ¼ 7, Sudan) 19  13. LAYING DATES: Mauritania, June–Oct; Gambia, June– Oct; Senegal, May–Nov; Mali, Aug–Oct, in S from July; Ghana, July–Oct (nest-building, Mar); Togo, Aug; Burkina Faso, (nesting June–Oct); Niger, May–Sept (fledged young, Nov); Nigeria, June–Oct; Zaı¨re, Aug–Nov; Sudan, May, Aug–Oct; Ethiopia, Mar–Aug; Uganda, May–June, Aug– Sept, Nov; Kenya, Feb, Apr, Sept. INCUBATION: by both Y and X. Period: 12 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: chicks hatch on successive days (Nigeria). In captivity, both Y and X feed young. Nestling period: 18–20 days. Captive Y first sang at 4 months, moulted into breeding plumage at 1 year. BREEDING SUCCESS/SURVIVAL: no information. Key References

Nakonzer, B. (1986), Serle, W. (1943b).

Plate 6

Ploceus pelzelni (Hartlaub). Slender-billed Weaver. Tisserin de Pelzeln.

(Opp. p. 75)

Sitagra pelzelni Hartlaub, 1887. Zool. Jahrb., 2, p. 243; Magungo, Uganda. NOMENCLATURAL NOTE:

also called Little Weaver (e.g. Bannerman 1949, Serle and Morel 1977), now the name of P. luteolus.

Forms a superspecies with P. luteolus and P. subpersonatus. Range and Status. Endemic resident. Liberia, unconfirmed sightings at Marshall and Sinoe (Gatter 1997). Ivory Coast, coastal area, Grand Bassam, Azagny. Ghana, coast at Panbros, Cape Coast, Ada, Densu. Togo, wetlands near coast (Sio R., Tokpli). Nigeria, coastal belt from Lagos to Calabar. Cameroon, Wouri R., Debundscha, Victoria,

Yabassi. Gabon, Omboue´, Port Gentil, Massouka. Congo, along river. Zaı¨re, in NE from L. Edward to Rutshuru, at base of Kivu volcanoes, along major rivers and swamps in Kasai and Katanga. Rwanda. Burundi. Uganda, Nile drainage area, L. Victoria basin, Lango and SW region. Kenya and Tanzania, L. Victoria margins. Angola, Cabinda

112

PLOCEIDAE

Voice. Tape-recorded (104, 109, B, GREG, McVIC, MOR, PAY). Song a pleasant continuum of chuckling bulbul-like notes, ‘chip-wop-weep-wop-wop-weep-wip-wop-cheepwop-wip . . . ’, quieter than other weavers. Calls mainly scratchy and grating, with weaver swizzles and chatters, and a soft ‘tsip’ or ‘tsssp’; alarm calls shrill and squeaky. General Habits. Inhabits savanna woodland with large Acacia trees, open scrubland with small trees, edges of farmland and large gardens, arid thornscrub and red sandy country (Kordofan, Sudan). In E Africa at 400–1500 m. Not gregarious; usually in pairs, occasionally singly or in small flocks (family parties?). Forages mainly in acacias and other small trees, working methodically through foliage, searching carefully, slightly warbler-like. Sometimes feeds on ground, in dry season may forage with waxbills (Gambia). Resident, but may expand north in Mauritania in rainy season; in N Nigeria, common at all seasons. Post-nuptial moult in Mali in Sept-Oct (Paludan 1936) and in Nigeria started by Oct (Fry 1971). Food. Seeds and insects; 4 stomachs contained only seeds, 6 had insects, including small beetles and caterpillars (Chapin 1954). Breeding Habits. Solitary nester (rarely, in colonies); monogamous. Pair often returns to same site in successive years. Y sings to X from a perch or hanging from nest, but does not tremble wings when singing. NEST: globular structure with short, vertical entrance tube at side (A), densely woven using fine, narrow herb and grass stems, grass blades, and small vine tendrils (Collias and Collias 1964). Strips torn from fronds of fan palm Borassus flabellifer often used (Mali: Lamarche 1981). Lined with grass seedheads and other fine material. Size, 100 long by 80 high (Collias and Collias 1964). Length of tunnel varies from 50 to 300 with int. diam. 25; 3–55 m above ground, suspended from thorny branches of small tree. Nests often built in Ziziphus and ‘heglig’ trees within 50 cm of occupied wasp nests (Sudan: Lynes 1924–1925); associated with hornets in Gambia (Barlow et al. 1997), but not in Zaı¨re. Pair seems to select thorny trees (Lynes 1924–1925, Moreau 1943). Nest construction takes 3–4

days, after which Y may continue work on tunnel, and tunnel may be added to nest long after breeding has ended (Lippens and Wille 1976). EGGS: 2–3. Plain white, ovate, slightly glossy. SIZE: (n ¼ 9, Nigeria) 175–206  125–139 (184  131,); (Cameroon) 165–190  125–130; (n ¼ 7, Sudan) 19  13. LAYING DATES: Mauritania, June–Oct; Gambia, June– Oct; Senegal, May–Nov; Mali, Aug–Oct, in S from July; Ghana, July–Oct (nest-building, Mar); Togo, Aug; Burkina Faso, (nesting June–Oct); Niger, May–Sept (fledged young, Nov); Nigeria, June–Oct; Zaı¨re, Aug–Nov; Sudan, May, Aug–Oct; Ethiopia, Mar–Aug; Uganda, May–June, Aug– Sept, Nov; Kenya, Feb, Apr, Sept. INCUBATION: by both Y and X. Period: 12 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: chicks hatch on successive days (Nigeria). In captivity, both Y and X feed young. Nestling period: 18–20 days. Captive Y first sang at 4 months, moulted into breeding plumage at 1 year. BREEDING SUCCESS/SURVIVAL: no information. Key References

Nakonzer, B. (1986), Serle, W. (1943b).

Plate 6

Ploceus pelzelni (Hartlaub). Slender-billed Weaver. Tisserin de Pelzeln.

(Opp. p. 75)

Sitagra pelzelni Hartlaub, 1887. Zool. Jahrb., 2, p. 243; Magungo, Uganda. NOMENCLATURAL NOTE:

also called Little Weaver (e.g. Bannerman 1949, Serle and Morel 1977), now the name of P. luteolus.

Forms a superspecies with P. luteolus and P. subpersonatus. Range and Status. Endemic resident. Liberia, unconfirmed sightings at Marshall and Sinoe (Gatter 1997). Ivory Coast, coastal area, Grand Bassam, Azagny. Ghana, coast at Panbros, Cape Coast, Ada, Densu. Togo, wetlands near coast (Sio R., Tokpli). Nigeria, coastal belt from Lagos to Calabar. Cameroon, Wouri R., Debundscha, Victoria,

Yabassi. Gabon, Omboue´, Port Gentil, Massouka. Congo, along river. Zaı¨re, in NE from L. Edward to Rutshuru, at base of Kivu volcanoes, along major rivers and swamps in Kasai and Katanga. Rwanda. Burundi. Uganda, Nile drainage area, L. Victoria basin, Lango and SW region. Kenya and Tanzania, L. Victoria margins. Angola, Cabinda

Ploceus pelzelni

Ploceus pelzelni

Field Characters. Length 11 cm. A small weaver with extensive black patch from forecrown through face to throat, ending in point on breast; X lacks black and has top of head green, black bill; juv. like X but olive above with some streaks on back, dull yellow below, bill pale. At all stages told from similar Little Weaver P. luteolus by slender bill, swamp habitat. Voice. Tape-recorded (104, B, CHA, GREG, McVIC). Song-like phrase of hard, piercing, reedy notes, repeated ‘tyutyook, tyook-tyook’; other calls include harsh ‘check’ or ‘chyack’, high ‘ti-tyik-tyik-tyik’, single ‘tyook’ and buzzy ‘bzzzzzt’ recalling Sedge Warbler Acrocephalus schoenobaenus. General Habits. Restricted to wetland areas for breeding, foraging in trees and bushes in adjoining habitats; in W Africa inhabits mangrove, coastal lagoons, marshlands and ricefields; in S Zaı¨re, ambatch thickets bordering lakes; also gardens and hedges in urban areas. Generally below 1700 m but up to 2350 m in Ruwenzoris. Singly or in pairs or flocks. Forages at all levels in trees. Agile; often hangs upside down when foraging; foot structure shows adaptations to papyrus habitat (Leisler and Winkler 2001). Flock moves rapidly through trees, reminiscent of whiteeyes Zosterops.

south to Bengo and Cuanza Norte. Zambia, L. Mweru, Chisenga I., Kasenga. Common but localised, throughout range. Description. P. p. pelzelni (Hartlaub): NE Zaı¨re, Uganda, Kenya, Rwanda, Burundi and Tanzania. ADULT Y: forehead, crown to just behind eye, lores, cheeks and ear-coverts black; hind-crown and nape golden yellow, passing into olive-green; mantle olivegreen, feathers with faint central streaks; rump greenish yellow; tail olive-green with yellow margins to feathers; chin and throat black, extending onto breast as a bib; sides of breast, flanks, belly, thighs and undertail-coverts golden yellow; wings olive-brown, feathers with yellow margins, broader on coverts and tertials; underwing grey, flecked with yellow. Bill black; eyes brown to yellowish brown; legs blue-grey. No non-breeding dress. ADULT X: lacks any black on head: forehead, crown and nape yellowish green with clear yellow superciliary stripe; cheeks and ear-coverts yellowish green, lores greenish; mantle olive-green with faint central streaks; rump washed with yellow; tail and wings like Y; chin, throat, breast, belly and undertail-coverts golden yellow, duller on flanks; thighs greenish. Bill black; eyes brown; legs blue-grey. SIZE (12 YY, 12 XX): wing, Y 60–66 (623), X 56–62 (595); tail, Y 39–47 (428), X 39–45 (427); bill, Y 160–186 (175), X 154–178 (168); tarsus, Y 170–191 (184), X 176–189 (184). WEIGHT: Y (Kenya, n ¼ 16) 117–151 (142), (Uganda, n ¼ 28) 122–159 (141), (Zaı¨re, n ¼ 9) 10–13 (11); X (Kenya, n ¼ 15) 114–149 (131), (Uganda, n ¼ 18) 118–144 (125), (Zaı¨re, n ¼ 4) 10–11 (108). IMMATURE: resembles X, but upperparts browner, underparts duller yellow with some brown wash on throat and breast; belly white. Upper mandible pale horn, lower mandible yellowish with brown tip; eyes brown. NESTLING: initially covered in moderately long buffy white down; greyish white down, still present when feathers start to emerge. P. p. monacha (Sharpe): W Africa, W Zaı¨re, Angola, Zambia. Bill shorter, less slender. Eyes brown to dull yellow.

Food. Primarily insects; one stomach contained seeds, others only insects, including small caterpillars and ants. In Cameroon, 32 stomachs all contained insects (Serle 1965). Breeding Habits. Colonial nester, in groups generally of 1 year, varying greatly (Hey¨l 1980a). BREEDING SUCCESS/SURVIVAL: over 2 seasons, 35–37% of eggs laid produced young which fledged successfully (Elliott 1973a). Nests robbed by boomslang Dispholidus typus, rats and African Harrier Hawk Polyboroides typus; parasitized by Diederik Cuckoo Chrysococcyx caprius; at one colony in KwaZulu-Natal, 8 of 41 nests were parasitized (Jensen and Vernon 1970). Cuckoos may be attacked (pers. obs.), and African Goshawk Accipiter tachiro was mobbed, but birds remained silent and avoided African Harrier Hawk (Skead 1995). Nests may be heavily infested with mites. YY with larger harems have significantly higher breeding success. Cape Weavers were regular prey of Barn Owls Tyto alba at 2 sites in Western Cape (Grindley et al. 1973). Oldest-known bird 7 years 7 months (Laycock and Hanmer 1981, Craig 1982). Annual ad. mortality estimated at 51%, with no difference between sexes. Of 17 birds ringed as nestlings, 12 recovered in first year, and 2 after 3 years. Using a different technique, Specific Death Rate (reciprocal of longevity) estimated at 0716 (Yom-Tov et al. 1994), and an analysis of both dispersal and recovery of ringed birds showed that this species both moved longer distances and had a shorter survival time than P. cucullatus or P. velatus, with 95% of Cape Weavers recovered within 4 years of ringing (Oatley and Underhill 2001).

1994). Period 13–14 days. Nest temperature differs little from ambient. Egg temperature averages 364
C during incubation; can fall to 284
C.

Key References

Skead, C. J. (1947), Elliott, C. C. H. (1973a).

Ploceus temporalis (Bocage). Bocage’s Weaver. Tisserin de Bocage.

Plate 7

Hyphantornis temporalis Bocage, 1880. J. Lisboa, vii, p. 224; Cacunda, Angola.

(Opp. p. 90)

Range and Status. Endemic resident. NW Zambia near Mwinilunga; Angola, from N Huı´la, Huambo and Bie´ to S Lunda Sul, Moxico and Lunda Norte; S Zaı¨re (Lualaba, Kando); uncommon. Description. ADULT Y (breeding): forehead and crown golden yellow; nape, mantle, rump and tail olive-green; lores and cheeks olive-brown, ear-coverts yellow; chin and throat olive-brown, breast, belly, thighs, undertail-coverts yellow; wings brown, edged with greenish yellow; underwing light olive-green. Bill black; eyes cream; legs brown to flesh-coloured. ADULT Y (nonbreeding): like ad. X but eyes white (Aspinwall and Beel 1998). ADULT X: forehead, crown, nape, mantle, rump and tail dark olive-green; lores olive-green, cheeks yellowish, ear-coverts olivegreen; chin and throat yellowish; breast, flanks and thighs light olive-green; belly, vent and undertail-coverts light grey; wings dark brown with yellowish edges to feathers; underwing grey. Bill brown; eyes greyish; legs brown. SIZE (13 YY, 4 XX): wing, Y 81– 97 (834), X 75–79 (765); tail, Y 47–53 (499), X 44–46 (450); bill, Y 178–194 (186), X 175–183 (179); tarsus, Y 210–238

(225), X 203–214 (209). WEIGHT: unsexed (Angola, n ¼ 3) 344– 370, juv. Y 345. IMMATURE and NESTLING: no information. TAXONOMIC NOTE: formerly treated as a subspecies of P. capensis (Moreau and Greenway 1962); placed in a superspecies with P. capensis by Hall and Moreau (1970), but relationships uncertain.

Field Characters. Length 15–16 cm. Superficially similar to allopatric Cape Weaver P. capensis but bill smaller. Breeding Y has pale eye set in olive-brown face mask, long olive-brown throat patch. X has olive-green lores and hint of light olive-green face patch, brown eye; darker and greener than sympatric weavers. Non-breeding Y like X but eye white. Voice. Tape-recorded (CART). Call, while perched or in flight, a rather soft but grating ‘chyap’, ‘chyip’ or ‘shret’; also a longer, faster series ‘shreti-shreti-shreti shret shret’ and a loud ‘cha’ (Aspinwall and Beel 1998).

Ploceus temporalis

125

DEVELOPMENT AND CARE OF YOUNG: young fed by X, predominantly with insects. Start calling at 4–5 days, older chicks make insistent screeching. Late in season Y feeds nestlings and may feed fledglings; some YY feed young as much as X does. Period: c. 17 days. Y skull pneumatization takes >1 year, varying greatly (Hey¨l 1980a). BREEDING SUCCESS/SURVIVAL: over 2 seasons, 35–37% of eggs laid produced young which fledged successfully (Elliott 1973a). Nests robbed by boomslang Dispholidus typus, rats and African Harrier Hawk Polyboroides typus; parasitized by Diederik Cuckoo Chrysococcyx caprius; at one colony in KwaZulu-Natal, 8 of 41 nests were parasitized (Jensen and Vernon 1970). Cuckoos may be attacked (pers. obs.), and African Goshawk Accipiter tachiro was mobbed, but birds remained silent and avoided African Harrier Hawk (Skead 1995). Nests may be heavily infested with mites. YY with larger harems have significantly higher breeding success. Cape Weavers were regular prey of Barn Owls Tyto alba at 2 sites in Western Cape (Grindley et al. 1973). Oldest-known bird 7 years 7 months (Laycock and Hanmer 1981, Craig 1982). Annual ad. mortality estimated at 51%, with no difference between sexes. Of 17 birds ringed as nestlings, 12 recovered in first year, and 2 after 3 years. Using a different technique, Specific Death Rate (reciprocal of longevity) estimated at 0716 (Yom-Tov et al. 1994), and an analysis of both dispersal and recovery of ringed birds showed that this species both moved longer distances and had a shorter survival time than P. cucullatus or P. velatus, with 95% of Cape Weavers recovered within 4 years of ringing (Oatley and Underhill 2001).

1994). Period 13–14 days. Nest temperature differs little from ambient. Egg temperature averages 364
C during incubation; can fall to 284
C.

Key References

Skead, C. J. (1947), Elliott, C. C. H. (1973a).

Ploceus temporalis (Bocage). Bocage’s Weaver. Tisserin de Bocage.

Plate 7

Hyphantornis temporalis Bocage, 1880. J. Lisboa, vii, p. 224; Cacunda, Angola.

(Opp. p. 90)

Range and Status. Endemic resident. NW Zambia near Mwinilunga; Angola, from N Huı´la, Huambo and Bie´ to S Lunda Sul, Moxico and Lunda Norte; S Zaı¨re (Lualaba, Kando); uncommon. Description. ADULT Y (breeding): forehead and crown golden yellow; nape, mantle, rump and tail olive-green; lores and cheeks olive-brown, ear-coverts yellow; chin and throat olive-brown, breast, belly, thighs, undertail-coverts yellow; wings brown, edged with greenish yellow; underwing light olive-green. Bill black; eyes cream; legs brown to flesh-coloured. ADULT Y (nonbreeding): like ad. X but eyes white (Aspinwall and Beel 1998). ADULT X: forehead, crown, nape, mantle, rump and tail dark olive-green; lores olive-green, cheeks yellowish, ear-coverts olivegreen; chin and throat yellowish; breast, flanks and thighs light olive-green; belly, vent and undertail-coverts light grey; wings dark brown with yellowish edges to feathers; underwing grey. Bill brown; eyes greyish; legs brown. SIZE (13 YY, 4 XX): wing, Y 81– 97 (834), X 75–79 (765); tail, Y 47–53 (499), X 44–46 (450); bill, Y 178–194 (186), X 175–183 (179); tarsus, Y 210–238

(225), X 203–214 (209). WEIGHT: unsexed (Angola, n ¼ 3) 344– 370, juv. Y 345. IMMATURE and NESTLING: no information. TAXONOMIC NOTE: formerly treated as a subspecies of P. capensis (Moreau and Greenway 1962); placed in a superspecies with P. capensis by Hall and Moreau (1970), but relationships uncertain.

Field Characters. Length 15–16 cm. Superficially similar to allopatric Cape Weaver P. capensis but bill smaller. Breeding Y has pale eye set in olive-brown face mask, long olive-brown throat patch. X has olive-green lores and hint of light olive-green face patch, brown eye; darker and greener than sympatric weavers. Non-breeding Y like X but eye white. Voice. Tape-recorded (CART). Call, while perched or in flight, a rather soft but grating ‘chyap’, ‘chyip’ or ‘shret’; also a longer, faster series ‘shreti-shreti-shreti shret shret’ and a loud ‘cha’ (Aspinwall and Beel 1998).

126

PLOCEIDAE

Ploceus temporalis

General Habits. Little-known. In bushes and trees near water, usually in dambos; alongside Holub’s Golden Weaver P. xanthops in Zambia. Rank grassland and tall forbs alongside streams and at the edges of marshes in Angola. In flocks of up to 50. Food. Insects; beetles, grasshoppers and caterpillars in 2 stomachs (H. Penry and P. Bowen, pers. comm.). Breeding Habits. Probably polygynous; colonial with >30 nests together. NEST: ovoid, with short entrance tunnel below, or without tunnel at all (A). Rather loosely woven from grass culms without flowering heads (Angola) or raffia palm leaves or strips of grass leaf blades (Zambia). Roof thick, with some grass heads; interior ceiling made of narrow

pieces of leaf blade and some Pennisetum grass heads. Suspended from branches overhanging water, or from elephant grass along river; 4–20 nests per tree. Length 170, height 140; tunnel 30–60. EGGS: 2–4, av. 23 (Zambia, n ¼ 9). Pyriform, plain skyblue. SIZE: (n ¼ 20, Zambia) av. 237  156. LAYING DATES: Zambia, Aug–Sept, (YY at colony June– Nov); Angola, May, Aug, Oct (juveniles May). INCUBATION: no information. DEVELOPMENT AND CARE OF YOUNG: spiders in a chick’s stomach. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius. Key References Aspinwall, D. R. and Beel, C. (1998), Colebrook-Robjent, J. F. R. (1980).

Plate 7

Ploceus subaureus A. Smith. Eastern Golden Weaver; Yellow Weaver. Tisserin jaune.

(Opp. p. 90)

Ploceus subaureus A. Smith, 1839. Ill. Zool. S. Afr., Aves, pl. 30; Algoa Bay, South Africa. Forms a superspecies with P. xanthops. Range and Status. Endemic resident. Somalia, 1, Balcad Nature Res., 1987. Kenya, coast south of Tana R. mouth, inland to Tsavo, Meru, Embu, Barsalinga, Kilocho. Tanzania, coast south to Mikindani, Zanzibar, inland to ˆ i, Shire valley, northward Songea, Njombe, Moshi. Malaw along lake shore to Mkanjila, Nkata Bay. Mozambique, Mopeia, Mocuba, Netia, Lurio Sul, Namapa, Beira, Vunduzi R., Buzi R., Bazaruto I., Inhambane, Incomati R., Manhica, Maputo, Inhaca I., Bela Vista. South Africa, Mpumalanga, in 2 summers (1974–1975) in Kruger Nat.

Park (Tarboton et al. 1987); KwaZulu-Natal, throughout coastal belt; E Cape, on coast south to Port Elizabeth, up to 50 km inland along river valleys. Swaziland, occasional summer visitor to lowveld, population estimate of 20 birds (Parker 1994). Locally common to abundant; population in S Mozambique estimated at >20,000 birds (Parker 1999). Description. P. s. subaureus A. Smith (includes ‘tongensis’): Mozambique south of Save R. to E Cape. ADULT Y (breeding): forehead and crown golden yellow; lores blackish, cheeks and

126

PLOCEIDAE

Ploceus temporalis

General Habits. Little-known. In bushes and trees near water, usually in dambos; alongside Holub’s Golden Weaver P. xanthops in Zambia. Rank grassland and tall forbs alongside streams and at the edges of marshes in Angola. In flocks of up to 50. Food. Insects; beetles, grasshoppers and caterpillars in 2 stomachs (H. Penry and P. Bowen, pers. comm.). Breeding Habits. Probably polygynous; colonial with >30 nests together. NEST: ovoid, with short entrance tunnel below, or without tunnel at all (A). Rather loosely woven from grass culms without flowering heads (Angola) or raffia palm leaves or strips of grass leaf blades (Zambia). Roof thick, with some grass heads; interior ceiling made of narrow

pieces of leaf blade and some Pennisetum grass heads. Suspended from branches overhanging water, or from elephant grass along river; 4–20 nests per tree. Length 170, height 140; tunnel 30–60. EGGS: 2–4, av. 23 (Zambia, n ¼ 9). Pyriform, plain skyblue. SIZE: (n ¼ 20, Zambia) av. 237  156. LAYING DATES: Zambia, Aug–Sept, (YY at colony June– Nov); Angola, May, Aug, Oct (juveniles May). INCUBATION: no information. DEVELOPMENT AND CARE OF YOUNG: spiders in a chick’s stomach. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius. Key References Aspinwall, D. R. and Beel, C. (1998), Colebrook-Robjent, J. F. R. (1980).

Plate 7

Ploceus subaureus A. Smith. Eastern Golden Weaver; Yellow Weaver. Tisserin jaune.

(Opp. p. 90)

Ploceus subaureus A. Smith, 1839. Ill. Zool. S. Afr., Aves, pl. 30; Algoa Bay, South Africa. Forms a superspecies with P. xanthops. Range and Status. Endemic resident. Somalia, 1, Balcad Nature Res., 1987. Kenya, coast south of Tana R. mouth, inland to Tsavo, Meru, Embu, Barsalinga, Kilocho. Tanzania, coast south to Mikindani, Zanzibar, inland to ˆ i, Shire valley, northward Songea, Njombe, Moshi. Malaw along lake shore to Mkanjila, Nkata Bay. Mozambique, Mopeia, Mocuba, Netia, Lurio Sul, Namapa, Beira, Vunduzi R., Buzi R., Bazaruto I., Inhambane, Incomati R., Manhica, Maputo, Inhaca I., Bela Vista. South Africa, Mpumalanga, in 2 summers (1974–1975) in Kruger Nat.

Park (Tarboton et al. 1987); KwaZulu-Natal, throughout coastal belt; E Cape, on coast south to Port Elizabeth, up to 50 km inland along river valleys. Swaziland, occasional summer visitor to lowveld, population estimate of 20 birds (Parker 1994). Locally common to abundant; population in S Mozambique estimated at >20,000 birds (Parker 1999). Description. P. s. subaureus A. Smith (includes ‘tongensis’): Mozambique south of Save R. to E Cape. ADULT Y (breeding): forehead and crown golden yellow; lores blackish, cheeks and

Ploceus subaureus Voice. Tape-recorded (99, B, McVIC). Shrill repeated ‘tsip’ or ‘tsyip’ accelerates into high-pitched weaver swizzles and dry trills. Calls include grating ‘zweep’, triple ‘chussychussy-chussy’ sharp ‘tzit’ of alarm; shrill chatter from group in colony.

Ploceus subaureus

General Habits. Inhabits coastal plains, river floodplains and lowland river valleys; below 1300 m in E Africa. Forms large flocks when not breeding; forages in trees away from nesting areas, and roosts with other Ploceus species. In Kenya, regularly joins mixed-species flocks foraging in bushed woodland (Pomeroy and Tengecho 1982). Forages on ground, alongside Spectacled Weaver P. ocularis and Southern Yellow-billed Hornbill Tockus flavirostris (KwaZulu-Natal).

?

ear-coverts golden yellow; nape, mantle and rump greenish yellow; tail greenish yellow, outer feathers with yellow inner webs; chin, throat and breast golden yellow, often with some greenish wash; flanks, belly, thighs and undertail-coverts golden yellow; wing-coverts greenish yellow, remiges light brown with narrow yellow outer margins, broad yellow inner web. Bill black; eyes red; legs dark brown. ADULT Y (non-breeding): forehead and crown greenish yellow; faint dark central streaks on mantle; yellow superciliary stripe; underparts dull yellow, sometimes grey flecked with yellow feathers; thighs greyish. Bill horn-brown; eyes red; legs dark brown. ADULT X (breeding): like nonbreeding Y, but eyes dark brown. ADULT X (non-breeding): upperparts uniform olive-green with dark central streaks on feathers of mantle, faint streaks on nape and crown; chin, throat and breast dull buffy; belly, thighs and undertail-coverts greyish white. Bill pale brown; eyes brown; legs brown. SIZE (13 YY, 11 XX): wing, Y 82–86 (843), X 74–79 (771); tail, Y 57–64 (595), X 51–60 (549); bill, Y 206–216 (210), X 184–209 (196); tarsus, Y 223–246 (231), X 202–218 (211). WEIGHT: (South Africa) Y (n ¼ 22) 295–390 (350); X (n ¼ 24) 215–310 (264). IMMATURE: like non-breeding X; but paler, especially on underparts, with variable amount of white. NESTLING: not described. ˆ i, N P. s. aureoflavus A. Smith: Kenya, Tanzania, Malaw Mozambique. Y in breeding dress with orange wash on forehead, crown, sides of head, chin and throat. Smaller. WEIGHT: Y (Kenya, n ¼ 13) 204–275 (251), (Tanzania, n ¼ 37) 20–29 (263); X (Kenya, n ¼ 4) 200–210 (205); (Tanzania, n ¼ 12) 18–23 ˆ i) 1 Y 262, 1 X 28. (208); (Malaw

Field Characters. Length 12–16 cm. Bright yellow, with rounded head, no black or chestnut mask; breeding Y in E Africa has strong rufous-orange wash on head. Smaller and shorter-billed than Holub’s Golden Weaver P. xanthops, eye red, orange or pink, not pale yellow. X green above with some streaking (heaviest on non-breeding birds), belly white; juv. like non-breeding X but eye dark. Does not overlap Orange Weaver P. aurantius; for differences from Golden Palm Weaver P. bojeri, see that species.

Food. Seeds, including rice and millet (a pest of these crops in Zanzibar – Pakenham 1979); also insects. Young fed largely on insects; feeds regularly on termite alates (Brooke et al. 1972); anthers of Nicotiana glauca flowers, nectar of Erythrina caffra and Aloe bainesii (Skead 1997). Breeding Habits. Colonial, with small territories; probably polygynous. 150 nests in large tamarind tree (Zanzibar: Vaughan 1930). Y displays hanging below nest entrance, fluttering wings; when X enters nest, Y perches nearby with drooped wings, quivering tail. NEST: oval to spherical, with entrance on underside (A); compact and tightly-woven of grass or strips from reeds, often attached to a single reed stem at the side, sometimes

127

128

PLOCEIDAE

supported by reed on either side (B, an unfinished nest). Lined with soft Eragrostris inflorescences, rarely a few feathers. Sited usually only 1–2 m above water or ground, often suspended from low Acacia bushes; reeds or branches above nest are stripped of leaves. On Zanzibar, in tall trees such as tamarinds and coconut palms, or in low shrubs and rank vegetation; nests in trees and palms up to 9 m above ground (Moreau and Moreau 1937). May nest alongside Golden Palm Weaver P. bojeri in E Africa, Southern Brown-throated Weaver P. xanthopterus and Thick-billed Weaver Amblyospiza albifrons in South Africa. EGGS: 2–4 (24, n ¼ 147). White, cream or pale blue, plain, or variably marked with dark brown, violet or black spots. SIZE: (n ¼ 53, southern Africa) 210–260  141–163 (230  151). LAYING DATES: Kenya, Jan–Mar, May, July–Dec; Tanzania, (nesting July), Zanzibar, Apr–June (breeding ˆ i, Oct–Feb; Mozambique, Sept–Feb, condition, Oct); Malaw Apr; South Africa, KwaZulu-Natal and E Cape, Sept–Feb. INCUBATION: by X only, period not known. DEVELOPMENT AND CARE OF YOUNG: nestling period (in captivity): 19–22 days. BREEDINGSUCCESS/SURVIVAL: African Fish Eagle Haliaeetus ˆ i, vocifer tore nests off reeds and consumed contents (Malaw Tweddle and Tweddle 1984). Occasional host of Diederik Cuckoo Chrysococcyx caprius (KwaZulu-Natal, Jensen and Vernon 1970). Oldest bird recaptured after >8 years ˆ i, Hanmer 1989); of 13 birds recovered, 1 had (Malaw moved 78 km, the others were within 10 km of ringing site (Craig 1982). Key References

Pakenham, R. H. W. (1979), Skead, C. J. (1995).

Plate 7

Ploceus xanthops (Hartlaub). Holub’s Golden Weaver. Tisserin safran.

(Opp. p. 90)

Hyphantornis xanthops Hartlaub, 1862. Ibis, p. 342; Angola. Forms a superspecies with P. subaureus. Range and Status. Endemic resident. Congo, in SW, north to Mindouli. Zaı¨re, in W, north to Bululu, Katanga, Marungu, Kivu, L. Albert. Uganda, L. Victoria Basin, in W, north to Bunyoro and Kabalega Falls. Kenya, Mt Elgon, Saiwa and Kerio Valley, south and east to Loita and Nguruman hills, Nairobi, Kitui and Embu. Rwanda. Burundi. Tanzania, east to Songea, Njombe, Mahenge and Babati. Angola, Cabinda southwards, except SW ˆ i, Namibe. Zambia, throughout except extreme SW. Malaw throughout. Mozambique, Vila Coutinho, Furancungo, Tete region, Vial Paiva de Andrada, Gorongosa Nat. Park, Vila de Manica, south of Save R. (old record at Conguno, recent sighting at Mawayela). Namibia, Caprivi region, along Cunene River. Botswana, Okavango, Chobe and Linyati R. systems. Zimbabwe, central plateau south to

Bikita and Great Zimbabwe, west to Bulawayo and Matobos Nat. Park, middle Zambezi valley and N sector of Hwange Nat. Park, general in E highlands, uncommon in Sabi valley and absent from S lowveld. Swaziland, local in centre and W; estimated 4000 birds (Parker 1994). South Africa, Limpopo R. south along escarpment in Northern Prov. and Mpumulanga, coast and midlands in KwaZulu-Natal. Local and sparsely distributed. Description. ADULT Y: forehead and crown golden yellow; lores, cheeks and ear-coverts yellow; nape golden yellow, merging into olive-green on mantle; mantle, rump and tail olive-green; chin and throat golden yellow with some chestnut-brown wash; breast, flanks, belly, thighs and undertail-coverts golden yellow; wing feathers brown with dull yellow margins, wing-coverts greenish with yellow margins; underwing yellow. No seasonal change in

128

PLOCEIDAE

supported by reed on either side (B, an unfinished nest). Lined with soft Eragrostris inflorescences, rarely a few feathers. Sited usually only 1–2 m above water or ground, often suspended from low Acacia bushes; reeds or branches above nest are stripped of leaves. On Zanzibar, in tall trees such as tamarinds and coconut palms, or in low shrubs and rank vegetation; nests in trees and palms up to 9 m above ground (Moreau and Moreau 1937). May nest alongside Golden Palm Weaver P. bojeri in E Africa, Southern Brown-throated Weaver P. xanthopterus and Thick-billed Weaver Amblyospiza albifrons in South Africa. EGGS: 2–4 (24, n ¼ 147). White, cream or pale blue, plain, or variably marked with dark brown, violet or black spots. SIZE: (n ¼ 53, southern Africa) 210–260  141–163 (230  151). LAYING DATES: Kenya, Jan–Mar, May, July–Dec; Tanzania, (nesting July), Zanzibar, Apr–June (breeding ˆ i, Oct–Feb; Mozambique, Sept–Feb, condition, Oct); Malaw Apr; South Africa, KwaZulu-Natal and E Cape, Sept–Feb. INCUBATION: by X only, period not known. DEVELOPMENT AND CARE OF YOUNG: nestling period (in captivity): 19–22 days. BREEDINGSUCCESS/SURVIVAL: African Fish Eagle Haliaeetus ˆ i, vocifer tore nests off reeds and consumed contents (Malaw Tweddle and Tweddle 1984). Occasional host of Diederik Cuckoo Chrysococcyx caprius (KwaZulu-Natal, Jensen and Vernon 1970). Oldest bird recaptured after >8 years ˆ i, Hanmer 1989); of 13 birds recovered, 1 had (Malaw moved 78 km, the others were within 10 km of ringing site (Craig 1982). Key References

Pakenham, R. H. W. (1979), Skead, C. J. (1995).

Plate 7

Ploceus xanthops (Hartlaub). Holub’s Golden Weaver. Tisserin safran.

(Opp. p. 90)

Hyphantornis xanthops Hartlaub, 1862. Ibis, p. 342; Angola. Forms a superspecies with P. subaureus. Range and Status. Endemic resident. Congo, in SW, north to Mindouli. Zaı¨re, in W, north to Bululu, Katanga, Marungu, Kivu, L. Albert. Uganda, L. Victoria Basin, in W, north to Bunyoro and Kabalega Falls. Kenya, Mt Elgon, Saiwa and Kerio Valley, south and east to Loita and Nguruman hills, Nairobi, Kitui and Embu. Rwanda. Burundi. Tanzania, east to Songea, Njombe, Mahenge and Babati. Angola, Cabinda southwards, except SW ˆ i, Namibe. Zambia, throughout except extreme SW. Malaw throughout. Mozambique, Vila Coutinho, Furancungo, Tete region, Vial Paiva de Andrada, Gorongosa Nat. Park, Vila de Manica, south of Save R. (old record at Conguno, recent sighting at Mawayela). Namibia, Caprivi region, along Cunene River. Botswana, Okavango, Chobe and Linyati R. systems. Zimbabwe, central plateau south to

Bikita and Great Zimbabwe, west to Bulawayo and Matobos Nat. Park, middle Zambezi valley and N sector of Hwange Nat. Park, general in E highlands, uncommon in Sabi valley and absent from S lowveld. Swaziland, local in centre and W; estimated 4000 birds (Parker 1994). South Africa, Limpopo R. south along escarpment in Northern Prov. and Mpumulanga, coast and midlands in KwaZulu-Natal. Local and sparsely distributed. Description. ADULT Y: forehead and crown golden yellow; lores, cheeks and ear-coverts yellow; nape golden yellow, merging into olive-green on mantle; mantle, rump and tail olive-green; chin and throat golden yellow with some chestnut-brown wash; breast, flanks, belly, thighs and undertail-coverts golden yellow; wing feathers brown with dull yellow margins, wing-coverts greenish with yellow margins; underwing yellow. No seasonal change in

Ploceus xanthops

Ploceus xanthops

Voice. Tape-recorded (86, 88, 99, 104, B, C, F, CART, LEM, PAY). Song of birds in Kenya and Zambia begins with harsh chatter, followed by dry trill, ending with 4–5 shrill notes, e.g. ‘kakakakakaka-grrrrreeee, ka-jee-kalagwee’; varied: central trill may be loud and liquid. Further south, song more a typical weaver swizzle, with accelerating introductory notes that may be dry or liquid, ‘jit, jit, jit-jit-jiji-swizzzzzz-jerjerjerjer’; swizzle song lasts c. 5 s, and may be repeated 4–5 times without a break. Call a dry, grating ‘jip’, ‘jert’ or ‘juck’. General Habits. Inhabits bushy areas interspersed with tall grass: forest margins, woody savanna and dense vegetation along watercourses and stream lines. In E Africa usually at altitudes of 1200–2300 m; up to 2200 m in Inyanga area, Zimbabwe. Occurs mainly singly and in pairs. Nonˆ i forage in forest canopy, well away breeding birds in Malaw from usual nesting areas (Medland 1992b). Territorial pairs dominated Masked Weaver P. velatus at an artificial feeding station in Botswana (Herremans and Herremans-Tonnoeyr 1995). Birds in Kenya ‘ant’ using ants Paratrachina sp. (Cunningham-van Someren 1996). Extract grubs from pods of Burkea africana (Lorber 1982); termite alates caught and eaten on ground, also caught on wing and eaten at perch (Brooke 1970).

plumage. Bill black; eyes pale yellow; legs brown. ADULT X: like Y, but yellow wash on forehead only, upperparts uniform olivegreen; underparts yellow, less richly coloured than Y and no chestnut wash on chin or throat. Bill black; eyes yellow; legs brown. Aberrant grey, olive and buff X described (Hanmer 2002a). SIZE (12 YY, 11 XX): wing, Y 87–97 (922), X 80–91 (852); tail, Y 64–79 (702), X 58–75 (651); bill, Y 217–235 (225), X 205–229 (217); tarsus, Y 246–273 (258), X 230–259 (245). WEIGHT: (Zaı¨re) Y (n ¼ 6) 42–50 (46), X (n ¼ 5) 36–42 (39), (Kenya) Y 45, 50, X (n ¼ 5) 360–425 (389), (Zambia) Y (n ¼ 9) 395–500 (443), X (n ¼ 4) 350–400 (373); unsexed (Tanzania, n ¼ 8) 335–470, (Zimbabwe, n ¼ 3) 350–402; immature (Zambia, n ¼ 6) 360–440 (379). IMMATURE: upperparts uniform dark olive-green, tail feathers with pale greenish yellow margins; wings and coverts dark brown with greenish buff margins; lores, cheeks and ear-coverts olivegreen; chin and throat pale buffy yellow; broad buff-brown band across breast; flanks dark buff; belly, thighs and undertail-coverts pale buff to yellowish. Bill pale brown with darker base; eyes dark brown. NESTLING: not described. TAXONOMIC NOTE: several races have been described (‘camburni’, ‘jamesoni’, ‘manuensis’) based on size and colour differences. All colour variations can be found in Angola, so species best treated as monotypic (Hall 1960, Pinto 1965; see also Dean 2000).

Field Characters. Length 16–18 cm. A golden yellow weaver, larger and heavier-billed than Yellow Weaver P. subaureus, eye pale yellow, not red; Y has orange wash on throat and upper breast only (YY of sympatric golden weavers have whole head orange). X lacks orange and is duller and greener, upperparts unstreaked (streaked in X Eastern Golden Weaver P. subaureus); juv. like X but darker above, buffy yellow below with brownish breastband, eyes dark.

Food. Insects, berries, fruits of Ficus and Rhus pyroides, green and ripe seeds of grasses such as Panicum maximum; seeds of Pinus patula (Zimbabwe: Harwin 1983). Nectar of Ruttya and Erythrina flowers (Zimbabwe), Agave, Prunus, Grevillia robusta, Acrocarpus fraxinifolius and Leonotis nepetifolia (E Africa). Insects include grubs and termites. Young fed grasshoppers, praying mantises, tabanid flies and spiders (Mattocks 1971). Breeding Habits. Colonial nester; monogamous; territorial. Breeds in loose colony of 2–5 nests; seldom more than 2–3 YY in one area. Y builds several nests; only one normally used for eggs; he perches near nests, calling and displaying to any approaching X (Ruwet 1965). Pair meeting up after separation performs ‘song stretch’ display; Y perches next to X, leans towards her with throat and head feathers ruffled as he sings (Craig 1984). Y on horizontal perch near X crouched, tail high and quivering; pursued X when she flew off (Savalli 1989). May be doublebrooded in Zaı¨re (Vincent 1949). NEST: large, bulky, kidney-shaped structure, with no spout but usually with ‘veranda’ of seed-heads protruding from entrance on underside (A). Woven of coarse, broadbladed grass, lined with grass heads and stems and sometimes a few feathers; often a pad of plant down in egg chamber. Usually suspended at its roof; rarely, slung between reeds or supported at one side by reed or grass stalk; placed in tall bushes amongst elephant grass, on drooping tips overhanging water, and in reedbeds; usually 2–25 m above ground or water. Of 97 nests in Zimbabwe, 54 in reeds, 43 in trees. SIZE: 170–190 long, 120–140 high, with entrance 60–65  65–70. Built by Y; X does not bring material except seed heads for ceiling and lining, but may do some weaving alongside Y. Leaves stripped from vegetation surrounding nest.

129

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PLOCEIDAE

spotted with red and lilac. In Zimbabwe, plain blue or spotted white are commonest. SIZE: (n ¼ 29, Zaire) 227– 267  155–169 (243  162, (n ¼ 8, Zimbabwe) 239– 254  160–173 (245  166), (n ¼ 55, South Africa) 222–255  153–173 (241  162). LAYING DATES: Zaı ¨re, Jan–Apr and Sept–Nov in S, July, Nov and Jan–Apr in Ruwenzori region and NE; Rwanda, Dec, Mar–May; Uganda, Feb, Apr–June, Oct; Kenya, Jan– May, Aug, Oct–Dec; Angola, Sept–Apr; Zambia, Sept–Feb ˆ i, Oct–Apr; (nest-building Aug, young in nest Apr); Malaw Botswana, Sept–Feb; Zimbabwe, primarily Sept–Feb, also Aug, Mar–Apr; Mozambique, Oct–Nov, Jan–May; South Africa, KwaZulu-Natal, Oct–Feb, Mpumalanga, Sept–Mar. INCUBATION: period, in captivity, c. 14 days. DEVELOPMENT AND CARE OF YOUNG: Young fed by both Y and X. Nestling period: 19–22 days. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius. In South Africa, 6 birds recovered within 10 km of place where ringed, after max. 25 months (Craig 1982); oldest bird recaptured in Zimbabwe after 6 years (Manson 1982). ˆ i). Immaculate greenish EGGS: 1–3 (20, n ¼ 26, Malaw blue, pale blue, white or pinkish; or evenly freckled and

Key References

Mattocks, T. (1971), Vincent, A. W. (1949).

Plate 7

Ploceus aurantius (Vieillot). Orange Weaver. Tisserin orange.

(Opp. p. 90)

Malimbus aurantius Vieillot, 1805. Ois. Chant., p. 73; Malimbe, Portuguese Congo. Forms a superspecies with P. bojeri and P. castaneiceps. Range and Status. Endemic resident. Sierra Leone, coastal in SE. Liberia, common, all along coast and on rocky islets up to 15 km offshore (and unconfirmed sight record in Mt Nimba area). Ivory Coast, along coast and inland north to 6
N along larger rivers. Ghana, coast from Axim to Cape Coast. Togo, uncommon at L. Togo. Nigeria, coast from Badagri to Bonny, inland to Warri and Okitipupa. Cameroon, Korup Nat. Park, in Sanaga and Mbam R. basins, inland in gallery forests as at Lobe´ke´ Nat. Park. Central African Republic, Lobaye and Haute Sangha regions, and record at Orkam. Mbini, Mont Alen Nat. Park. Gabon, localized at Ogooue´, Fougamou, Mouila, Impfondo and on rivers of Ivindo basin. Congo, coastal and along R. Congo. Zaı¨re, from lower Congo R. and central Kasai region along major rivers to upper Uele and Semliki Valley; also Okapi Faunal Res., Itombwe. Uganda, shores of L. Victoria, Busoga to Masaka, Sese Islands. Kenya, occasional in L. Victoria basin, as at Kisumu. Tanzania, W shores of L. Victoria south to Bukoba. Angola, Cabinda southward along coastal plain to Luanda. Localized, often uncommon. Description. P. a. aurantius (Vieillot): Sierra Leone to Zaı¨re. ADULT Y: forehead, crown and nape orange; mantle olive-green; rump orange; tail olive-brown; wings dark brown with narrow orange edges to remiges, broader edges to coverts, and orange tips to upper primary coverts; lores black to a variable extent, cheeks and ear-coverts orange; underparts uniform orange. Bill blackish to brown; legs brown; eyes pale grey to white. No seasonal change

Ploceus aurantius

in plumage. ADULT X: forehead, crown and nape yellowish green; mantle duller olive-green and unstreaked; rump yellowish; tail and wings as in Y; faint yellow superciliary stripe; lores, cheeks

130

PLOCEIDAE

spotted with red and lilac. In Zimbabwe, plain blue or spotted white are commonest. SIZE: (n ¼ 29, Zaire) 227– 267  155–169 (243  162, (n ¼ 8, Zimbabwe) 239– 254  160–173 (245  166), (n ¼ 55, South Africa) 222–255  153–173 (241  162). LAYING DATES: Zaı ¨re, Jan–Apr and Sept–Nov in S, July, Nov and Jan–Apr in Ruwenzori region and NE; Rwanda, Dec, Mar–May; Uganda, Feb, Apr–June, Oct; Kenya, Jan– May, Aug, Oct–Dec; Angola, Sept–Apr; Zambia, Sept–Feb ˆ i, Oct–Apr; (nest-building Aug, young in nest Apr); Malaw Botswana, Sept–Feb; Zimbabwe, primarily Sept–Feb, also Aug, Mar–Apr; Mozambique, Oct–Nov, Jan–May; South Africa, KwaZulu-Natal, Oct–Feb, Mpumalanga, Sept–Mar. INCUBATION: period, in captivity, c. 14 days. DEVELOPMENT AND CARE OF YOUNG: Young fed by both Y and X. Nestling period: 19–22 days. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius. In South Africa, 6 birds recovered within 10 km of place where ringed, after max. 25 months (Craig 1982); oldest bird recaptured in Zimbabwe after 6 years (Manson 1982). ˆ i). Immaculate greenish EGGS: 1–3 (20, n ¼ 26, Malaw blue, pale blue, white or pinkish; or evenly freckled and

Key References

Mattocks, T. (1971), Vincent, A. W. (1949).

Plate 7

Ploceus aurantius (Vieillot). Orange Weaver. Tisserin orange.

(Opp. p. 90)

Malimbus aurantius Vieillot, 1805. Ois. Chant., p. 73; Malimbe, Portuguese Congo. Forms a superspecies with P. bojeri and P. castaneiceps. Range and Status. Endemic resident. Sierra Leone, coastal in SE. Liberia, common, all along coast and on rocky islets up to 15 km offshore (and unconfirmed sight record in Mt Nimba area). Ivory Coast, along coast and inland north to 6
N along larger rivers. Ghana, coast from Axim to Cape Coast. Togo, uncommon at L. Togo. Nigeria, coast from Badagri to Bonny, inland to Warri and Okitipupa. Cameroon, Korup Nat. Park, in Sanaga and Mbam R. basins, inland in gallery forests as at Lobe´ke´ Nat. Park. Central African Republic, Lobaye and Haute Sangha regions, and record at Orkam. Mbini, Mont Alen Nat. Park. Gabon, localized at Ogooue´, Fougamou, Mouila, Impfondo and on rivers of Ivindo basin. Congo, coastal and along R. Congo. Zaı¨re, from lower Congo R. and central Kasai region along major rivers to upper Uele and Semliki Valley; also Okapi Faunal Res., Itombwe. Uganda, shores of L. Victoria, Busoga to Masaka, Sese Islands. Kenya, occasional in L. Victoria basin, as at Kisumu. Tanzania, W shores of L. Victoria south to Bukoba. Angola, Cabinda southward along coastal plain to Luanda. Localized, often uncommon. Description. P. a. aurantius (Vieillot): Sierra Leone to Zaı¨re. ADULT Y: forehead, crown and nape orange; mantle olive-green; rump orange; tail olive-brown; wings dark brown with narrow orange edges to remiges, broader edges to coverts, and orange tips to upper primary coverts; lores black to a variable extent, cheeks and ear-coverts orange; underparts uniform orange. Bill blackish to brown; legs brown; eyes pale grey to white. No seasonal change

Ploceus aurantius

in plumage. ADULT X: forehead, crown and nape yellowish green; mantle duller olive-green and unstreaked; rump yellowish; tail and wings as in Y; faint yellow superciliary stripe; lores, cheeks

Ploceus aurantius and ear-coverts dull yellow; chin, throat and breast white washed with yellow; belly and undertail-coverts off-white; flanks and thighs grey. Bill grey to brown, lower mandible paler; eyes brown to whitish; legs brown. SIZE (12 YY, 12 XX): wing, Y 69–75 (715), X 63–68 (658); tail, Y 40–46 (438), X 37–42 (396); bill, Y 165–191 (177), X 158–175 (166); tarsus, Y 172–195 (180), X 163–179 (169). WEIGHT: Y (Uganda, n ¼ 6) 245–290 (265), (Zaı¨re, n ¼ 10) 21–26 (237); X (n ¼ 3) 223–249 (243), (Zaı¨re) 25; juv. Y (Zaı¨re) 202, 21, 23; unsexed (n ¼ 5) 212–289 (247). IMMATURE: upperparts olive-green; wings dark brown with greenish yellow margins, inner webs yellow; cheeks and earcoverts olive-green; yellow superciliary stripe; underparts white, washed with yellow in throat region; bill horn-brown. Sub-ad. Y more yellow on underparts than X. NESTLING: not described. P. a. rex Neumann: Uganda, Kenya, Tanzania. Breeding Y has black spot in front of eye, sometimes extending to lores.

Field Characters. Length 14 cm. A weaver of swampy vegetation, with pale slender bill and pale grey eye; Y has head and body uniform orange, including rump, back olive, wings with broad yellow edgings; small black spot in front of eye sometimes present. Voice. Tape-recorded (104, B). Call a loud ‘kew-kewp’ or ‘kew-kew-kewp’, sometimes 7–8 notes; incorporated into conversational chuckling song of grating notes and liquid whistles, including down-slurred ‘kweeyu-kweeyu-kweeyu’; also has a dry, nasal accelerating trill, the equivalent of a swizzle, preceded by high thin whistles. Y has both short and long songs; long song used in nest advertisement display and when X enters nest (Crook 1969). General Habits. In W Africa, primarily coastal in vegetation and mangroves fringing lagoons; coastal thickets, trees along banks of rivers near their mouths, small creeks. Always in small groups close to water. Appears to move away from colony site after breeding. Birds on offshore islets commute to coast to feed (Liberia). In central Africa along major rivers; around L. Victoria, in papyrus swamps. Food. Insects and fruit (berries, pulp and fruit seed; locust eggs, beetles, caterpillars). Young fed caterpillars and grasshoppers. Breeding Habits. Colonial; colonies of 12 to several hundred nests; nests often not close together; sometimes isolated nests in Gabon, on river banks or islands, also in primary forest and villages. Colony of 20 YY in trees fringing a pond near Cape Coast, Ghana. In Cameroon, all nests solitary except 1 on fringe of Vieillot’s Black Weaver P. nigerrimus colony; in Zaı¨re also nests near P. nigerrimus and Village Weaver P. cucullatus in palms and larger trees, and in Uganda often associated with colonies of Goldenbacked Weaver P. jacksoni and Northern Brown-throated Weaver P. castanops. NEST: compact ovoid or globular structure, tightly woven of grass or strips from palm fronds (A), no entrance spout (Liberia, Zaı¨re) or short tubular tunnel (Uganda). Sited in reeds or other vegetation, 15–25 m over water; and up to 48 m over ground; on offshore islets, nests low down in bush or shrub on leeward side (Gatter and Hodgson 1987);

favours Nauclea pobeguini trees (Gabon); uses mangrove ferns and Pandanus trees. SIZE: 130 long, 90–115 high; entrance oval, 40  475. EGGS: 2, pale blue, green, brown or sometimes reddish, with fine or bold lilac and pale brown spots, denser at thick end. SIZE: 20  14–15 (Liberia), 194–213  145–151 (Zaı¨re), 20–22  14–15 (Uganda). LAYING DATES: Liberia, Oct–Apr; Ghana, June–July (copulation, Apr); Togo, (nesting June); Nigeria, Feb, June, Sept, Nov–Dec; Cameroon, (nesting June, Dec; juv. Jan); Gabon, Oct–Mar (nesting Sept); Gabon, primarily Dec–Mar, also Sept–Nov; Congo, Dec, Feb; Zaı¨re, Apr– Sept in forest belt, Mar–Sept in Ituri, Itombwe, Jan (possibly all year near equator: Chapin 1954); Uganda, Jan–May, Nov, mainly Mar–Apr; Angola, (juv. July). INCUBATION: by X only. Period: 13–14 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: nestling period: 13– 16 days (in captivity).

Key Reference Brosset, A. and Erard, C. (1986).

131

132

PLOCEIDAE

Plate 7

Ploceus bojeri (Cabanis). Golden Palm Weaver. Tisserin palmiste.

(Opp. p. 90)

Hyphantornis bojeri Cabanis, 1869. Von der Decken’s Reise, 3, p. 32; Mombasa, Kenya. Forms a superspecies with P. aurantius and P. castaneiceps. Range and Status. Endemic resident. Ethiopia, old records from upper Jubba R. at Bia, Sarrgo. Somalia, south of 5
N on Jubba and Shabeele Rivers and in Boni Forest. Kenya, coast from Lamu south to Moa, along Tana R. inland to Meru, Northern Uaso Nyiro, Nzio, Embu, Kiboko, Simba, Shaba and Samburu Game Res., Athi R., Kibwezi. Tanzania, coastal belt south to Tanga. Common on coast, localized inland. Description. ADULT Y: forehead, crown, nape, lores, cheeks and ear-coverts orange, ending in sharp line on nape; mantle greenish yellow with no streaking; rump more yellowish, slightly contrasting with mantle; tail greenish with yellow fringe; wings olive-green with yellow edges and vivid yellow inner webs to remiges, broader yellow edges on coverts, and yellow tips to upper coverts; chin and throat orange, passing into golden yellow, outlined by a chestnut-brown bib, better defined in some individuals, from corners of mouth to breast; breast, except for collar, golden yellow; belly, thighs, flanks and undertail-coverts golden yellow. Bill black; eyes brown; legs flesh-coloured. No seasonal change in plumage. ADULT X: forehead, crown, nape, cheeks and ear-coverts greenish yellow, with yellow superciliary stripe; mantle more greenish with faint streaking; rump more yellowish; tail and wings as in Y; underparts uniform yellow. Bill dark brown above, lighter brown lower mandible; eyes brown; legs flesh-coloured. SIZE (12 YY, 12 XX): wing, Y 71–80 (761), X 64–70 (658); tail, Y 48–57 (531), X 41–59 (444); bill, Y 169– 187 (178), X 158–185 (171); tarsus, Y 206–233 (220), X 176– 199 (186). WEIGHT: (Kenya) Y (n ¼ 37) 220–292 (255); X (n ¼ 14) 170–247 (197); (Tanzania) X (n ¼ 6) 190–240 (207). IMMATURE: like X, but underparts paler yellow and mantle streakier. NESTLING: not described.

Field Characters. Length 14 cm. A weaver of coastal palms in E Africa. Black eye in striking contrast to orange or yellow head. Head of Y uniform orange, shading to rufouschestnut patch on lower throat (in Y Eastern Golden Weaver P. subaureus orange-rufous face pales onto earcoverts and sides of neck). Yellow-headed subad. Y developing orange on nape and showing some orangerufous on lower throat easily mistaken for Taveta Golden Weaver P. castaneiceps, but lacks well-defined occipital crescent and rufous on upper breast (Zimmerman et al. 1996), and they are allopatric. X entirely yellow below (X Yellow has white belly, pink eye), back indistinctly streaked (heavily streaked in X Taveta Golden Weaver). Does not overlap Orange Weaver P. aurantius. Voice. Tape-recorded (B, GREG, McVIC, ZIM). In colonies, high-pitched chippering and chattering notes accelerating into swizzles. Variety of calls include grating ‘jyip’, ‘jeep’, ‘jut’, ‘jyert’ and ‘ti-jaa’, low nasal ‘jer’, higher, liquid ‘tyip’, ‘tyer’ or ‘ti-tyer’, and hissing ‘tssssss’; aggressive ‘kik! kik!’ when one Y chasing another. Song described as a creaking ‘eee-urrr-twa-twee-twu’ (Moreau and Moreau 1937).

Ploceus bojeri

General Habits. Inhabits palm savanna on coast, and riverine habitats below 1200 m and with >500 mm annual rainfall inland. Wing-moult mostly Apr-Aug, although some wing-moult in all months: (n ¼ 61, Mombasa, Kenya, Britton and Britton 1986); one case of interrupted moult. Food. Seeds and insects. Breeding Habits. Colonial, probably polygynous (single Y displaying at 4 nests: Collias and Collias 1964). Occurs in colonies alongside P. subaureus and with Village Weaver P. cucullatus. Y displays whilst hanging below nest entrance, with wings spread vertically, but wings usually move very little; head may be bowed slowly. NEST: spherical with no entrance tube. Outer shell woven by Y of grass strips or strips from palm fronds 3 mm wide, 250–300 long; complete inner shell made of short grass strips, 3–9 wide, 50–100 long, and lined with leaf fragments and fine grass heads suspended under palm frond or over water in thorn tree, or attached to tall grass and bushes; 15–4 m (occasionally >10 m) above ground as low as 03–18 m above water. Nests in Tanzania in bulrushes Typha or less often in reeds Phragmites, always attached to 3–5 leaves, which may meet above nest. Nest in bamboos (Mombasa). SIZE: 120–125 long, 90–100 high, entrance diam. 35. EGGS: 2. Sea-green, mottled all over with grey or reddish markings; also white, overlaid with fawn. SIZE: (Kenya,

Ploceus castaneiceps n ¼ 12) 195–222  135–155 (211  143); (Tanzania, n ¼ 4) 195–220  132–150 (212  143). LAYING DATES: Kenya, all months, with peaks in Oct and Jan; at Mombasa, nests May–Oct; Somalia, May–July; Tanzania, June, Dec–Apr.

133

BREEDING SUCCESS/SURVIVAL: House Crows Corvus splendens took eggs, young and adults; after repeated raids, a colony was abandoned (Mombasa, Ryall 1992).

Key Reference Collias, N. E. and Collias, E. C. (1964).

Ploceus castaneiceps (Sharpe). Taveta Golden Weaver. Tisserin de Taveta.

Plate 7

Hyphantornis castaneiceps Sharpe, 1890. Cat. Birds Brit. Mus., 13, p. 448; Taveta, Kenya.

(Opp. p. 90)

Forms a superspecies with P. aurantius and P. bojeri. Range and Status. Endemic resident. Kenya, Amboseli and Taveta area, L. Jipe. Tanzania, along Pangani R., south and south west of Mt Meru, Kilimanjaro; at 250–1500 m; common but very local. Description. ADULT Y: forehead, lores, cheeks and ear-coverts golden yellow; crown pale chestnut-brown, darker towards rear, with sharply-defined chestnut-brown collar on nape which links to bib; hind nape golden yellow, passing into greenish yellow on mantle, which is unstreaked; rump pale yellow; tail greenish yellow; wing olive-green with narrow yellow rims to flight feathers and broader yellow margins to tertials and wing-coverts, with upper primary coverts tipped with yellow; chin, throat and breast golden yellow, except for narrow band of chestnut-brown linked to nape, forming bib on breast, and enclosing some areas washed with chestnut; flanks, belly, thighs and undertail-coverts golden yellow. Bill black; eyes brown; legs brown. No seasonal change in plumage reported. ADULT X: forehead, crown and nape greenish yellow; mantle greener with dark central streaks; rump yellowish, unstreaked; tail greenish yellow; wing as in Y; yellow superciliary stripe, lores, cheeks and ear-coverts buffy yellow; chin, throat and breast pale yellow; belly off-white with some yellow streaks; flanks with buffy grey wash; thighs and undertailcoverts off-white washed with yellow. Bill brown, upper mandible darker; eyes brown; legs brown. SIZE (8 YY, 9 XX): wing, Y 75– 78 (764), X 66–73 (697); tail, Y 51–55 (524), X 44–51 (470); bill, Y 186–192 (189), X 165–186 (173); tarsus, Y 210–233 (221), X 177–212 (202). WEIGHT: X (n ¼ 3) 178, 187, 19. IMMATURE: pattern like X, but browner on upperparts, buffy on breast and flanks, bill dark brown above, yellowish white on lower mandible. NESTLING: not described.

Field Characters. Length 14–145 cm. Golden yellow, with chestnut crown ending in sharply defined crescent on nape, and chestnut band on breast. X and juv differ from Eastern Golden Weaver P. subaureus in dark eye, yellow supercilium, heavily streaked back and 2-tone bill (dark above, yellowish below); X yellow below (no white on belly), juv. browner above, buffy below. Very restricted range in S Kenya/N Tanzania; does not overlap Orange Weaver P. aurantius or Golden Palm Weaver P. bojeri. Voice. Tape-recorded (B, GREG, McVIC, PAR). Calls, harsh grating ‘jet’ or ‘jick’, buzzy scold, ‘dzzzzzzz’ or ‘jaaaaaaa’, and a swizzle. Chattering is low-pitched. Song a twittering ‘creee-er-curee-twee-twee’ (Zimmerman et al. 1996).

Ploceus castaneiceps

General Habits. Inhabits woodland and dry bush country; moves into swamps to breed. Food. Grass seeds, also maize; ants (in 1 stomach). Breeding Habits. Colonial, in swampy or flooded areas. Breeding areas deserted abruptly and completely, for weeks or months at a time; not clear if this follows a regular periodicity in Tanzania (Sclater and Moreau 1932–1933). NEST: spherical or ovoid with no entrance tube but a slight porch; woven of strips of reed leaf-blades. Entrance diam. 35. Generally suspended over water in reedbeds or bulrushes Typha; sometimes in low trees overhanging water. EGGS: 3, olive-green with fine darker flecks. SIZE: (n ¼ 1) 23  16. LAYING DATES: Kenya, Oct–May; Tanzania, Sept–Jan, Apr–May. INCUBATION: period:13 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: nestling period: 16 days (in captivity).

Ploceus castaneiceps n ¼ 12) 195–222  135–155 (211  143); (Tanzania, n ¼ 4) 195–220  132–150 (212  143). LAYING DATES: Kenya, all months, with peaks in Oct and Jan; at Mombasa, nests May–Oct; Somalia, May–July; Tanzania, June, Dec–Apr.

133

BREEDING SUCCESS/SURVIVAL: House Crows Corvus splendens took eggs, young and adults; after repeated raids, a colony was abandoned (Mombasa, Ryall 1992).

Key Reference Collias, N. E. and Collias, E. C. (1964).

Ploceus castaneiceps (Sharpe). Taveta Golden Weaver. Tisserin de Taveta.

Plate 7

Hyphantornis castaneiceps Sharpe, 1890. Cat. Birds Brit. Mus., 13, p. 448; Taveta, Kenya.

(Opp. p. 90)

Forms a superspecies with P. aurantius and P. bojeri. Range and Status. Endemic resident. Kenya, Amboseli and Taveta area, L. Jipe. Tanzania, along Pangani R., south and south west of Mt Meru, Kilimanjaro; at 250–1500 m; common but very local. Description. ADULT Y: forehead, lores, cheeks and ear-coverts golden yellow; crown pale chestnut-brown, darker towards rear, with sharply-defined chestnut-brown collar on nape which links to bib; hind nape golden yellow, passing into greenish yellow on mantle, which is unstreaked; rump pale yellow; tail greenish yellow; wing olive-green with narrow yellow rims to flight feathers and broader yellow margins to tertials and wing-coverts, with upper primary coverts tipped with yellow; chin, throat and breast golden yellow, except for narrow band of chestnut-brown linked to nape, forming bib on breast, and enclosing some areas washed with chestnut; flanks, belly, thighs and undertail-coverts golden yellow. Bill black; eyes brown; legs brown. No seasonal change in plumage reported. ADULT X: forehead, crown and nape greenish yellow; mantle greener with dark central streaks; rump yellowish, unstreaked; tail greenish yellow; wing as in Y; yellow superciliary stripe, lores, cheeks and ear-coverts buffy yellow; chin, throat and breast pale yellow; belly off-white with some yellow streaks; flanks with buffy grey wash; thighs and undertailcoverts off-white washed with yellow. Bill brown, upper mandible darker; eyes brown; legs brown. SIZE (8 YY, 9 XX): wing, Y 75– 78 (764), X 66–73 (697); tail, Y 51–55 (524), X 44–51 (470); bill, Y 186–192 (189), X 165–186 (173); tarsus, Y 210–233 (221), X 177–212 (202). WEIGHT: X (n ¼ 3) 178, 187, 19. IMMATURE: pattern like X, but browner on upperparts, buffy on breast and flanks, bill dark brown above, yellowish white on lower mandible. NESTLING: not described.

Field Characters. Length 14–145 cm. Golden yellow, with chestnut crown ending in sharply defined crescent on nape, and chestnut band on breast. X and juv differ from Eastern Golden Weaver P. subaureus in dark eye, yellow supercilium, heavily streaked back and 2-tone bill (dark above, yellowish below); X yellow below (no white on belly), juv. browner above, buffy below. Very restricted range in S Kenya/N Tanzania; does not overlap Orange Weaver P. aurantius or Golden Palm Weaver P. bojeri. Voice. Tape-recorded (B, GREG, McVIC, PAR). Calls, harsh grating ‘jet’ or ‘jick’, buzzy scold, ‘dzzzzzzz’ or ‘jaaaaaaa’, and a swizzle. Chattering is low-pitched. Song a twittering ‘creee-er-curee-twee-twee’ (Zimmerman et al. 1996).

Ploceus castaneiceps

General Habits. Inhabits woodland and dry bush country; moves into swamps to breed. Food. Grass seeds, also maize; ants (in 1 stomach). Breeding Habits. Colonial, in swampy or flooded areas. Breeding areas deserted abruptly and completely, for weeks or months at a time; not clear if this follows a regular periodicity in Tanzania (Sclater and Moreau 1932–1933). NEST: spherical or ovoid with no entrance tube but a slight porch; woven of strips of reed leaf-blades. Entrance diam. 35. Generally suspended over water in reedbeds or bulrushes Typha; sometimes in low trees overhanging water. EGGS: 3, olive-green with fine darker flecks. SIZE: (n ¼ 1) 23  16. LAYING DATES: Kenya, Oct–May; Tanzania, Sept–Jan, Apr–May. INCUBATION: period:13 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: nestling period: 16 days (in captivity).

134

PLOCEIDAE

Plate 7

Ploceus princeps (Bonaparte). Prı´ncipe Golden Weaver. Tisserin de Prı´ncipe.

(Opp. p. 90)

Symplectes princeps Bonaparte, 1850. Consp. Gen. Av. imm, i, p. 439; Principe Island. Range and Status. Endemic resident, restricted to Prı´ncipe Island, where it is one of the commonest birds.

Ploceus princeps

Description. ADULT Y: forehead, crown and nape chestnutbrown; mantle and rump dull yellowish green; tail olive-green; lores, cheeks and ear-coverts chestnut-brown; chin and throat yellow; breast yellow with chestnut wash, individually variable in extent; flanks yellow with some chestnut wash; belly, thighs and undertail-coverts yellow; wing dark brown, with yellow margins to remiges, broad yellowish edges to coverts. Bill black; eyes white, tinged with yellow; legs brown. No seasonal change in plumage. ADULT X: forehead, crown, and nape yellowish green; mantle olive-green with faint central streaks on feathers; rump olive-green, unstreaked; tail olive-green; clear yellow superciliary stripe; lores and cheeks yellow; some chestnut-brown wash on ear-coverts; chin, throat and breast yellow; flanks and thighs greyish; belly and undertail-coverts white; wings dark brown. Bill brown; eyes pale greyish yellow; legs brown. SIZE (10YY, 2XX): wing, Y 84–90 (873), X 81; tail, Y 52–58 (555), X 51; bill, Y 231–253 (245), X 236, 237; tarsus, Y 221–237 (230), X 214, 215. WEIGHT: 1, unsexed, 312 (P. Jones, pers. comm.). IMMATURE: young Y like X, but more yellowish both on upperparts and underparts. Eyes brown. NESTLING: not described.

Field Characters. Length 18 cm. A large, heavy-billed weaver, mainly yellow, Y with orange-chestnut head from crown to cheeks; yellow eye conspicuous. The only weaver on Prı´ncipe I. Voice. Tape-recorded (104, B, JO PJ, TYE, CHR). Song in 3 parts: 4–6 loud whistles, 1–4 softer 2-note whistles, and set phrase of scratchy notes, ‘wee-wee-wee-wee, werdiwerdi, chuckji-jerji-chickwoyjer’. Also has wheezing song, ‘pzeep-pzeep-pzzzzzzzzzzrrrrrr’ (Borrow and Demey 2001). Contact call a loud ‘zeep’ (Snow 1950), probably the same as the short, sharp ‘ksuit’ of Christy and Clarke (1998), who also list repeated ‘tzic’ and liquid ‘ksueep’. General Habits. Absence of other weavers on Prı´ncipe has allowed it to occupy all available habitats and all vegetation levels, from grass and thick undergrowth in fields and banana plantations to canopy of tallest trees (Christy and Clarke 1998); occurs in primary and old secondary forest and forest edge, coffee plantations shaded by Erythrina trees, mixed vegetation on poorly-maintained plantations. Common to abundant in all degraded and secondary habitats; probably the first endemic species seen by visitors to the island, around the airport and in the gardens of San Antonio. Occurs in pairs which remain together all year, sometimes singly; often in groups of up to 30 birds. Foraging bands include many young and are very noisy; may join mixed-species flocks with e.g. Dorhn’s ThrushBabbler Horizorhinus dohrni and Prı´ncipe Glossy Starling Lamprotornis ornatus. Feeds on ground in taro fields, disappearing among large leaves and jumping from stem to stem. In tall vegetation beside R. Papagaio acts like a granivore, whereas in secondary forest and in undergrowth

at edges of primary forest acts more like an insectivore; gleans bark and searches through leaves and flowers in canopy of tall erythrinas like a typical insect-eater, joining

Ploceus xanthopterus

sunbirds and white-eyes Speirops at flowers (Christy and Clarke 1998); makes short sallies after flying insects.

Breeding Habits. Apparently monogamous, nesting in small groups in adjacent trees, with up to 6 nests in 1 tree. Y builds nest, then displays near it or hanging under it: beats wings and sings with head in nest entrance, then in most intense display leans back and points head downwards; then perches near nest, quivering wings and singing. NEST: large mass of green stalks, round or oval, with entrance below, without a tunnel (A, B – two views of same nest). Made from strips torn from leaves of palms or bananas, lined with fine grass. Suspended from tip of branch or palm leaf, usually >10 m above ground. Trees used include Erythrina, Fagara, Ceiba, Xylopia, Musanga, coconut palms. EGGS: 1–2, plain blue. LAYING DATES: May–June, Aug–Sept, perhaps Feb (nestbuilding Jan, Sept, juvs Feb–Mar). INCUBATION: tail of sitting bird protrudes from nest entrance, visible from below. DEVELOPMENT AND CARE OF YOUNG: fed by X; Y also feeds young once they are begging in nest entrance. BREEDING SUCCESS/SURVIVAL: nests may be robbed by Blue-breasted Kingfishers Halcyon malimbica and by Prı´ncipe Glossy Starling Lamprotornis ornatus.

Food. Fruit such as bananas, berries, chilis, oil palm fruit; seeds; insects including beetles; nectar of Erythrina trees.

Key References Christy, P. and Clark, W. V. (1998), Jones, P. J. and Tye, A. (1988).

135

Ploceus xanthopterus (Finsch and Hartlaub). Southern Brown-throated Weaver. Tisserin a` gorge Plate 7 brune. (Opp. p. 90) Hyphantornis xanthopterus Finsch and Hartlaub, 1870. Vo ¨g. Ost-Afr., p. 399; Shire Valley. Forms a superspecies with P. castanops.

Ploceus xanthopterus Range and Status. Endemic resident. Zaı¨re, unconfirmed records from L. Lufira (Ruwet 1965). Zambia, upper Zambezi R. and its tributaries. Zimbabwe, west of Victoria Falls in Zambezi valley. Namibia, Caprivi region. Botswana, Okavango,LinyantiandChobeR.systems.Tanzania,localinSW, ˆ i; old from Mufindi to Mwaya, and Songea region to L. Malaw ˆ i, lake littoral from Karonga record from Nachingwea. Malaw southwards, Mbenje Is., Shire R. valley. Mozambique, Mocuba, Mopeia, Zambezi R., Save R., Mapinhane, but no recent records from Save R. or north of 24
N, only from Inharrime south to Maputa. Zimbabwe, Mazoe R., SabiLundi confluence. South Africa, coastal lowlands of KwaZulu-Natal from Durban northwards, old record from Kangwane in Mpumalanga (Tarboton et al. 1987). Generally uncommon though local density can be high. In S Mozambique, population of c. 1000 birds; may be declining here and in Botswana as result of disturbance of wetlands. ˆ i, Mozambique Description. P. x. xanthopterus Tanzania, Malaw south to Zambezi R., NE Zimbabwe. ADULT Y (breeding): forehead and crown, cheeks and ear-coverts golden yellow, passing into yellowish green on nape; mantle greenish yellow; rump bright yellow; tail greenish with broad yellow inner webs

Ploceus xanthopterus

sunbirds and white-eyes Speirops at flowers (Christy and Clarke 1998); makes short sallies after flying insects.

Breeding Habits. Apparently monogamous, nesting in small groups in adjacent trees, with up to 6 nests in 1 tree. Y builds nest, then displays near it or hanging under it: beats wings and sings with head in nest entrance, then in most intense display leans back and points head downwards; then perches near nest, quivering wings and singing. NEST: large mass of green stalks, round or oval, with entrance below, without a tunnel (A, B – two views of same nest). Made from strips torn from leaves of palms or bananas, lined with fine grass. Suspended from tip of branch or palm leaf, usually >10 m above ground. Trees used include Erythrina, Fagara, Ceiba, Xylopia, Musanga, coconut palms. EGGS: 1–2, plain blue. LAYING DATES: May–June, Aug–Sept, perhaps Feb (nestbuilding Jan, Sept, juvs Feb–Mar). INCUBATION: tail of sitting bird protrudes from nest entrance, visible from below. DEVELOPMENT AND CARE OF YOUNG: fed by X; Y also feeds young once they are begging in nest entrance. BREEDING SUCCESS/SURVIVAL: nests may be robbed by Blue-breasted Kingfishers Halcyon malimbica and by Prı´ncipe Glossy Starling Lamprotornis ornatus.

Food. Fruit such as bananas, berries, chilis, oil palm fruit; seeds; insects including beetles; nectar of Erythrina trees.

Key References Christy, P. and Clark, W. V. (1998), Jones, P. J. and Tye, A. (1988).

135

Ploceus xanthopterus (Finsch and Hartlaub). Southern Brown-throated Weaver. Tisserin a` gorge Plate 7 brune. (Opp. p. 90) Hyphantornis xanthopterus Finsch and Hartlaub, 1870. Vo ¨g. Ost-Afr., p. 399; Shire Valley. Forms a superspecies with P. castanops.

Ploceus xanthopterus Range and Status. Endemic resident. Zaı¨re, unconfirmed records from L. Lufira (Ruwet 1965). Zambia, upper Zambezi R. and its tributaries. Zimbabwe, west of Victoria Falls in Zambezi valley. Namibia, Caprivi region. Botswana, Okavango,LinyantiandChobeR.systems.Tanzania,localinSW, ˆ i; old from Mufindi to Mwaya, and Songea region to L. Malaw ˆ i, lake littoral from Karonga record from Nachingwea. Malaw southwards, Mbenje Is., Shire R. valley. Mozambique, Mocuba, Mopeia, Zambezi R., Save R., Mapinhane, but no recent records from Save R. or north of 24
N, only from Inharrime south to Maputa. Zimbabwe, Mazoe R., SabiLundi confluence. South Africa, coastal lowlands of KwaZulu-Natal from Durban northwards, old record from Kangwane in Mpumalanga (Tarboton et al. 1987). Generally uncommon though local density can be high. In S Mozambique, population of c. 1000 birds; may be declining here and in Botswana as result of disturbance of wetlands. ˆ i, Mozambique Description. P. x. xanthopterus Tanzania, Malaw south to Zambezi R., NE Zimbabwe. ADULT Y (breeding): forehead and crown, cheeks and ear-coverts golden yellow, passing into yellowish green on nape; mantle greenish yellow; rump bright yellow; tail greenish with broad yellow inner webs

136

PLOCEIDAE

on outer feathers; chin and throat chestnut-brown, extending onto breast to some extent, variable individually; breast otherwise golden yellow; belly, flanks, thighs and undertail-coverts golden yellow; wings olive-brown with narrow yellow outer margins, and broad yellow; inner webs (so that in birds >3 years old brown is restricted to tip and band along rachis); wing-coverts olive-brown with broad yellow margins. Bill black; eyes bright brownish red; legs pinkish. ADULT Y (non-breeding): forehead, crown and nape olive-green with paler margins to feathers; mantle pale brown with blackish central streaks on feathers; rump buffy brown, unstreaked; tail olive-green with yellow edges and inner webs; lores, cheeks and ear-coverts olive-green; chin, throat and breast buffy; belly, flanks, thighs and undertail-coverts dull white, with buffy wash on flanks; wing-coverts olive-brown with narrow yellowish margins. Amount of yellow on flight feathers decreases with age, enabling birds to be placed in age classes of 15–25 years, 25–45 years and >45 years (Hanmer 1984). ADULT X: like non-breeding Y, but yellow on inner webs of wing feathers restricted to base, no change after first year. Upper mandible dark brown, lower mandible pale brown; eyes brown; legs brown. When breeding, distinct yellow wash on throat and breast, extending onto belly in midline; legs pinkish. SIZE: wing, Y (n ¼ 384) 69–77 (728), X (n ¼ 322) 60–68 (649); tail, Y (n ¼ 150) 44–53 (475), X (n ¼ 142) 39–46 (429); bill, Y (n ¼ 150) 175–195 (183), X (n ¼ 142) 160–180 (170); tarsus, Y (n ¼ 10) 214–236 ˆ i) Y (n ¼ 394) (221), X (n ¼ 4) 189–197 (193). WEIGHT: (Malaw 22–31 (251), X (n ¼ 342) 16–24 (192), imm. Y (n ¼ 376) 21–28 (238), imm X (n ¼ 365) 15–23 (184); (Mozambique) Y (n ¼ 119) 23–28 (249), X (n ¼ 103) 16–24 (191), imm. Y (n ¼ 126) 20–27 (233), imm X (n ¼ 123) 15–22 (182). Adult YY heaviest Sept– Dec, lose weight during breeding and moult period; ad. XX also heaviest Sept–Dec (excluding gravid ones). IMMATURE: like X, but underparts dull white with less buffy wash; broad buffy margins to wing-coverts. Legs grey until 1 year, thereafter brownish or pinkish; eyes dull brown, brighter after 1 year. Yellow on flight feathers and tail pale until first moult, then deeper yellow, but area expands progressively with age. NESTLING: not described. P. x. castaneigula (Cabanis): Namibia, Botswana and upper Zambezi River. Crown orange-yellow, in sharp contrast to green mantle. Larger than nominate race. P. x. marleyi (Roberts): Mozambique from Save R. southwards, SE Zimbabwe, South Africa. Crown bright yellow, little contrast with mantle; throat patch tawny rather than chestnut. Wing, Y (n ¼ 3, KwaZulu-Natal) 73–80; WEIGHT: Y 285, 305, 305; sub-ad. 26.

Field Characters. Length 15 cm. Small, with short tail and rather square head. Breeding Y is the only weaver in its range with brown face and throat sharply demarcated from yellow head and neck (does not meet Northern Brownthroated Weaver P. castanops); not likely to be found alongside much larger Cape Weaver P. capensis, which has orange-brown wash on face but white eye, dark lores and longer, more tapered bill. X and non-breeding Y have crown and upperparts olive-green; told from Holub’s and Eastern Golden Weavers P. xanthops and P. subaureus by dark eye. Breeding X otherwise like breeding X Eastern Golden Weaver, with yellow underparts, but non-breeding birds and buffy below, without yellow. Voice. Tape-recorded (88, 99, 103, B, HOR). Song, introductory thin, down-slurred whistles accelerating into swizzle, ending with 2 nasal buzzy notes, ‘zaaa-zaaaaaa’; instead of whistles song may begin with hard nasal calls: ‘zwaay, zwaay, zit zit zit-zit-zizizizizizzzzzzzzz, zaaazaaaaaa’; see sonagrams in Maclean 1993. Calls include

harsh grating ‘jeck’, ‘jit’, ‘cujecku’ or ‘jaaa’, double ‘bew-jit’ and thin hissing notes. General Habits. Inhabits reedbeds and tall aquatic vegetation in wetlands. Forages in adjacent riparian forest and other dense vegetation, but seldom far from water; at ˆ i, in non-breeding season, higher altitudes in Malaw forages in forest canopy well away from breeding areas (Medland 1992b). Usually in flocks of 2000 birds (Parker 1999), Swaziland about 600 (Parker 1994). Description. P. i. intermedius Ru ¨ ppell: Ethiopia and Somalia to Uganda, Rwanda, Kenya and W and central Tanzania (south to Rukwa). ADULT Y (breeding): forehead and forecrown black, merging with dark brown hindcrown; nape, hindneck and sides of neck golden yellow, tinged brown; mantle, scapulars and back yellowish olive-green with fine dusky shaft streaks; rump and short uppertail-coverts plain greenish yellow; long uppertailcoverts yellowish olive-green. Tail feathers dusky olive, edged yellowish green. Lores to forecrown, supercilium, ear-coverts, cheeks, chin and throat black, forming large face mask. Rest of underparts bright yellow, washed brown across upper breast, paler on undertail-coverts. Upperwing feathers dark brown; primaries and secondaries edged pale greenish yellow, primary coverts edged olive-green, broad edges and tips of tertials and greater coverts and broad tips of median coverts pale greenish yellow; lesser coverts broadly fringed yellowish olive-green. Underwing-coverts and axillaries yellow; undersides of inner borders of flight feathers greyish white. Bill black; eyes white or pale straw; legs grey. ADULT Y (non-breeding): like breeding ad. X but top of head and mantle yellowish olive-green, underparts more strongly tinged yellow. ADULT X (breeding): forehead to hindneck and sides of neck olive-green with faint dusky streaks; mantle, back and scapulars olive-green with narrow blackish brown streaks, faint on back; rump and short uppertail-coverts yellowish olive-green, long uppertail-coverts olive-green with dusky centres. Tail feathers dusky olive, edged olive-green. Poorly marked yellowish superciliary stripe, and below this a dusky olive stripe through lores and behind eye; cheeks and earcoverts olive-yellow marked with dusky olive-green. Chin and throat pale olive-yellow; breast olive-yellow, grading to white on lower belly, with brown tinge across upper breast; flanks brownish yellow; undertail-coverts pale yellow. Upperwing as ad. Y. Bill dark slate grey above, pale grey below; eyes pale clay yellow or yellowish white; legs dull greyish. ADULT X (nonbreeding): like breeding X, but breast to belly and undertailcoverts whiter. SIZE (10 YY, 10 XX): wing, Y 71–77 (733), X 67– 71 (692); tail, Y 44–49 (464), X 38–46 (433); bill, Y 17–185 (180), X 165–18 (170); tarsus, Y 205–22 (212), X 195–205 (202). WEIGHT: Kenya, Y (n ¼ 50) 173–268 (233), X (n ¼ 42) 175–254 (220); Uganda, Y (n ¼ 9) 199–259 (237), X (n ¼ 12) 20–258 (230). IMMATURE: juvenile similar to ad. X but upperparts browner, underparts off-white, washed buffy yellow on breast and flanks; eyes dark brown. NESTLING: not described. It has more natal down than other Ploceus species. P. i. cabanisii (Peters): Coastal Tanzania, south to Natal, west to N Namibia and interior S Angola. Y with hind-crown and breast brighter yellow, less washed with saffron. SIZE: wing, Y (n ¼ 16) 71–75 (738), X (n ¼ 17) 67–73 (69); tail, (n ¼ 29) 46–56; bill, (n ¼ 29) 14–17; tarsus, (n ¼ 29) 18–22. WEIGHT: (Gauteng, South Africa) Y (n ¼ 6) 21–242 (230); X (n ¼ 8) 179–222 (200). P. i. beattyi Traylor: W Angola (north of Benguela). Adult Y has chestnut wash on hind-crown darker, extending over hindneck; chestnut wash on breast darker, extending over flanks.

Field Characters. Length 125–14 cm. Adults in all plumages identified by pale yellow eye, blue-grey legs and feet. Black mask on breeding Y extends to top of crown, and is rounded on upper breast (pointed in Southern Masked Weaver P. velatus); Vitelline Masked Weaver P. vitellinus has red eye, saffron crown, reduced black mask barely reaching upper breast. Nape and

143

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PLOCEIDAE

underparts pure bright yellow from Tanzania south, but elsewhere in E Africa suffused with saffron. Speke’s P. spekei and Heuglin’s Masked Weavers P. heuglini have black masks and yellow eyes but both have yellow foreheads. X and non-breeding Y have yellow face (dark like through eye) and underparts, white only on belly. Immature with dark eye and brown legs probably not distinguishable. Voice. Tape-recorded (86, 88, 99, 104, B, F, LEM, PAY). ˆ i (104) typical weaver swizzles, Song at colony in Malaw ‘swizzzzy . . . jubjub . . . swizzurrrrrrrrr’ or descending ‘squeeerrrrrrrrr’, interspersed with short ‘chap’ or ‘chupchup’, high-pitched sharp ‘tsip’ or ‘tseep’, low ‘zwerrr’, metallic ‘chickle’ and fast, dry, buzzy chatter. In South Africa said to give ‘rasping swizzle’ (Maclean 1993), but songs on tapes 88 and 99 are a rather liquid chatter, some with a hollow, gobbling quality, with call notes ‘tyewp’, ‘pew-pew-pew’ and ‘tsop-tsop’. General Habits. Occurs in acacia savanna, bushveld, open woodland and riverine trees, preferring areas close to water and also human habitation. Arid grass and thorn-scrub country (Sudan). Bushland and open woodland, not necessarily associated with water (Somalia). Below 2000, or more usually 1500 m, in wooded and bushed grassland, cultivated areas, and pastures (Uganda). Open bushed and wooded habitats, up to 2000 m, but mainly below 1500 m (Kenya). Dry, open woodland, especially acacia, though usually not far from water (Zambia). Relatively dry areas; ˆ i). most typically acacia savanna near water (Malaw Riverine woodland (Angola). Tree and bush savanna, edges of deciduous woodland, riparian acacia; usually in river valleys but also in areas near water of ponds, dams, reservoirs and pools; nests in colonies in trees and bushes, sometimes over water (Botswana). Woodland and savanna (S Mozambique). Swamps, sometimes in bush vegetation, mangroves (Inhaca I., Mozambique). Open woodland or thornveld; favours areas close to water more than Southern Masked Weaver, but like it will nest in both trees and reeds; often nests away from open water in ornamental gardens (southern Africa). Gregarious in small flocks or solitary when not breeding. Joins large flocks of other weavers in winter in Botswana. Forages in canopy of trees, especially Acacia tortilis and A. albida, for insects, probing under leaves. Probes flowers for insects and nectar, face may be coloured orange or yellow with pollen. May roost with other weavers, including Red-billed Quelea Q. quelea. Extent of movements unknown, although colonies deserted in non-breeding season. Brief showers cause start of nest-building, but if no further rain, birds move on. Rainy season visitor to acacia savanna in SE Sudan. In Kenya usually resident, but also wanders and regularly moves into arid Tsavo Nat. Park in rains. In Botswana birds disappear in some years. 1 bird killed in Northern Province, 74 km from place of ringing. Food. Insects, especially caterpillars, also termites, caterpillars, and grasshoppers. Nectar (mainly of aloes), flowers

of tondo tree Albizia sp., mulberries, small seeds. Maize porridge in gardens. Chicks fed on insects. Breeding Habits. Polygynous; Y has 2–3 XX simultaneously, and probably several more in a season. Breeds in colonies of 3 to c. 60 nests. Nests usually very close together, sometimes suspended from other nests. Y displays from nest by fanning wings very fast, spreading tail, and slightly swaying head while singing. Old nests used by Cut-throat Finch Amadina fasciata for breeding. Nests may be associated with paper wasps Belonogaster. Colonies may be monospecific or mixed with other weavers, especially Village P. cucullatus, Viellot’s Black P. nigerrimus and Speke’s; also Southern Masked, Eastern Golden P. subaureus, and Thick-billed Amblyospiza albifrons in reeds, Red-billed Quelea, or Ru ¨ ppell’s Weaver P. galbula. NEST: kidney-shaped, thin-walled but strong, with no ceiling; constructed with narrow strips of grass, reed or palm leaves and fresh green fine grass heads (A); 1 built of pine needles. In Kenya nest material consists mainly of grass heads of Cynodon dactylon, Panicum maximum, Bothriochloa radicans, Digitaria argyrotricha, Digitaria scalarum, Pennisetum mezianum, Eragrostris aethiopicum, and Leptochloa chinensis; also acacia leaves; in Zimbabwe more strips of grass leaves, fewer grass heads; in Ethiopia grasses Echinochloa, Eragrostris, and Panicum. Short lengths of material stick out untidily. Ext. length 130–140, depth 80–95, entrance diam. 30–35, entrance length 25–120 (n ¼ 10).

Ploceus velatus

Built by Y (B, first stage of ring construction); X lines accepted nests, but often with little or no material (silky grey fibres, sometimes a few grass heads). Situated at end of branch on outside or inside of tree, also in bush or palm, often over water. Nest trees include gall Acacia Acacia drepanolobium, A. erubescens, fever tree A. xanthophloea, A. albida, Ficus, mangroves, flamboyant Delonix regia, pepper tree, Syringa, ambatch, Borassus and coconut palms, eucalyptus and pawpaw. Also sited in Phragmites and Typha reeds, papyrus and bamboo, and hung from roof edge of building, or from telephone wire. ˆi EGGS: usually 2–4; E Africa (n ¼ 40) 1–5 (23); Malaw ˆ i (n ¼ 123) (n ¼ 12) 2–3 (23); Zimbabwe, Zambia, Malaw

145

1–4 (22); Angola (n ¼ 7) 1–4 (21); southern Africa (n ¼ 35) 2–4 (25). Short to long ovals; little or no gloss; white or pale bluish white, coarse-grained with obvious pores, unmarked. SIZE: (E Africa) av. c. 23  15; (n ¼ 50, Uganda) 200–244  138–161 (221  147); (n ¼ 13, Zimbabwe) 189–219  139–155 (201  146); (n ¼ 47, southern Africa) 185–235  138–156 (213  146). LAYING DATES: Somalia, May, June, Aug, Nov; Ethiopia, Mar, May–July, Sept; E Africa: Tanzania, Feb–Apr; Region B, Mar–June; Region C, Apr; Region D, Jan– Aug; Rwanda, Oct–Nov, Mar–May; Zaı¨re, Jan–Apr; Congo, Jan; Angola, rains; Zambia, Aug–Apr (mainly Sept and ˆ i, Nov–Mar (peak Dec); Botswana, Oct– Jan–Feb); Malaw Feb; Zimbabwe, Aug–Mar (mainly Oct–Feb); southern Mozambique, Nov–Feb; Namibia, Jan–Mar; South Africa, Oct–Feb, Natal, Oct–Feb. INCUBATION: by X; 48% of day spent incubating. Period: unknown. DEVELOPMENT AND CARE OF YOUNG: young fed by both parents. BREEDING SUCCESS/SURVIVAL: occasional host to Diederik Cuckoo Chysococcyx caprius (24% overall rate in southern Africa, 66% in Zambia); rarely, of Klaas’s Cuckoo C. klaas. Eggs and nestlings taken by snakes, particularly boomslang Dispholidus typus. Nests of a mixed colony of Lesser Masked and Eastern Golden Weavers taken by Fish Eagle Haliaeetus vocifer and raided by Vervet Monkey Cercopithecus aethiops. Adult taken by Gabar Goshawk Micronisus gabar, Adult died after being strangled by fishline incorporated in nest. Infected with Haemoproteus and Leucocytozoon blood parasites (1 each of 2 adults). Greatest elapsed time of ringing recoveries, Y 8 years, X. May desert colony with nests containing eggs and chicks.

Key Reference Bruggers et al. (1985).

Ploceus velatus Vieillot. Southern Masked Weaver. Tisserin masque´.

Plate 8

Ploceus velatus Vieillot, 1819. Nouv. Dict. d’Hist. Nat., vol. xxxiv, p. 132; Namaqualand, restricted to Graaff-Reinet, E Cape, By Brooke, 1985, Ostrich 56, 214–215.

(Opp. p. 91)

Forms a superspecies with P. vitellinus, P. reichardi, P. katangae and P. ruweti. Range and Status. Endemic resident. In Angola common along coastal plain from Luanda and Bengo to Namibe, and east inland to W Benguela, S and W Huı´la and Cunene; isolated population at L. Dilolo and L. Cameia, Moxico (Dean 2000); breeding colony on Luachimo R. near Dundo in Lunda Norte (Gu ¨ nther and Feiler 1986) needs confirmation (Dean 1996). Common in S Zambia, Kafue drainage to Luswishi confluence, and throughout Zambezi drainage, north through Luangwa Valley. Unconfirmed sight records from Copperbelt may be P. katangae (q.v.). No records between L. Tanganyika and upper Luangwa valley. In ˆ i ranges higher than Lesser Masked Weaver Malaw P. intermedius, thus uncommon at Dedza and Mzimba.

Locally common in central and S Mozambique, but not in parts of littoral. In Namibia, absent from treeless parts of Namib. Common in Botswana, with isolated nesting colonies in semi-desert areas. Locally common in Zimbabwe. Most widespread weaver in South Africa, but uncommon in tropical lowlands of Natal, Swaziland and Transvaal lowveld; absent from Transkei; expanded its range in SW Cape in 20th century. Common in lowland Lesotho and in river valleys in highlands. Acceptance of fences as nest sites has allowed expansion into treeless areas. Sa˜o Tome´, common in N. Population in S Mozambique probably >5000 birds (Parker 1999). Density in Acacia savannas, Swaziland, 5, 8

Ploceus velatus

Built by Y (B, first stage of ring construction); X lines accepted nests, but often with little or no material (silky grey fibres, sometimes a few grass heads). Situated at end of branch on outside or inside of tree, also in bush or palm, often over water. Nest trees include gall Acacia Acacia drepanolobium, A. erubescens, fever tree A. xanthophloea, A. albida, Ficus, mangroves, flamboyant Delonix regia, pepper tree, Syringa, ambatch, Borassus and coconut palms, eucalyptus and pawpaw. Also sited in Phragmites and Typha reeds, papyrus and bamboo, and hung from roof edge of building, or from telephone wire. ˆi EGGS: usually 2–4; E Africa (n ¼ 40) 1–5 (23); Malaw ˆ i (n ¼ 123) (n ¼ 12) 2–3 (23); Zimbabwe, Zambia, Malaw

145

1–4 (22); Angola (n ¼ 7) 1–4 (21); southern Africa (n ¼ 35) 2–4 (25). Short to long ovals; little or no gloss; white or pale bluish white, coarse-grained with obvious pores, unmarked. SIZE: (E Africa) av. c. 23  15; (n ¼ 50, Uganda) 200–244  138–161 (221  147); (n ¼ 13, Zimbabwe) 189–219  139–155 (201  146); (n ¼ 47, southern Africa) 185–235  138–156 (213  146). LAYING DATES: Somalia, May, June, Aug, Nov; Ethiopia, Mar, May–July, Sept; E Africa: Tanzania, Feb–Apr; Region B, Mar–June; Region C, Apr; Region D, Jan– Aug; Rwanda, Oct–Nov, Mar–May; Zaı¨re, Jan–Apr; Congo, Jan; Angola, rains; Zambia, Aug–Apr (mainly Sept and ˆ i, Nov–Mar (peak Dec); Botswana, Oct– Jan–Feb); Malaw Feb; Zimbabwe, Aug–Mar (mainly Oct–Feb); southern Mozambique, Nov–Feb; Namibia, Jan–Mar; South Africa, Oct–Feb, Natal, Oct–Feb. INCUBATION: by X; 48% of day spent incubating. Period: unknown. DEVELOPMENT AND CARE OF YOUNG: young fed by both parents. BREEDING SUCCESS/SURVIVAL: occasional host to Diederik Cuckoo Chysococcyx caprius (24% overall rate in southern Africa, 66% in Zambia); rarely, of Klaas’s Cuckoo C. klaas. Eggs and nestlings taken by snakes, particularly boomslang Dispholidus typus. Nests of a mixed colony of Lesser Masked and Eastern Golden Weavers taken by Fish Eagle Haliaeetus vocifer and raided by Vervet Monkey Cercopithecus aethiops. Adult taken by Gabar Goshawk Micronisus gabar, Adult died after being strangled by fishline incorporated in nest. Infected with Haemoproteus and Leucocytozoon blood parasites (1 each of 2 adults). Greatest elapsed time of ringing recoveries, Y 8 years, X. May desert colony with nests containing eggs and chicks.

Key Reference Bruggers et al. (1985).

Ploceus velatus Vieillot. Southern Masked Weaver. Tisserin masque´.

Plate 8

Ploceus velatus Vieillot, 1819. Nouv. Dict. d’Hist. Nat., vol. xxxiv, p. 132; Namaqualand, restricted to Graaff-Reinet, E Cape, By Brooke, 1985, Ostrich 56, 214–215.

(Opp. p. 91)

Forms a superspecies with P. vitellinus, P. reichardi, P. katangae and P. ruweti. Range and Status. Endemic resident. In Angola common along coastal plain from Luanda and Bengo to Namibe, and east inland to W Benguela, S and W Huı´la and Cunene; isolated population at L. Dilolo and L. Cameia, Moxico (Dean 2000); breeding colony on Luachimo R. near Dundo in Lunda Norte (Gu ¨ nther and Feiler 1986) needs confirmation (Dean 1996). Common in S Zambia, Kafue drainage to Luswishi confluence, and throughout Zambezi drainage, north through Luangwa Valley. Unconfirmed sight records from Copperbelt may be P. katangae (q.v.). No records between L. Tanganyika and upper Luangwa valley. In ˆ i ranges higher than Lesser Masked Weaver Malaw P. intermedius, thus uncommon at Dedza and Mzimba.

Locally common in central and S Mozambique, but not in parts of littoral. In Namibia, absent from treeless parts of Namib. Common in Botswana, with isolated nesting colonies in semi-desert areas. Locally common in Zimbabwe. Most widespread weaver in South Africa, but uncommon in tropical lowlands of Natal, Swaziland and Transvaal lowveld; absent from Transkei; expanded its range in SW Cape in 20th century. Common in lowland Lesotho and in river valleys in highlands. Acceptance of fences as nest sites has allowed expansion into treeless areas. Sa˜o Tome´, common in N. Population in S Mozambique probably >5000 birds (Parker 1999). Density in Acacia savannas, Swaziland, 5, 8

146

PLOCEIDAE

Ploceus velatus

and 10 birds per 100 ha (Monadjem 2002); in Moremi Wildlife Reserve, Botswana, 014 birds per ha (Milewski and Campbell 1976). Description. P. v. velatus Vieillot: W Cape, N Cape, Free State. ADULT Y (breeding): narrow black band on forehead; face, ear-coverts, chin and throat black, ending in a point on lower neck; crown yellow washed with saffron; nape golden yellow; mantle, back and tail yellowish green, faintly streaked dusky; upper tail-coverts and rump yellow green; breast, belly, and flanks golden yellow, chest slightly washed with saffron; wingcoverts and flight feathers dusky with yellow edges. Bill black; eyes red to orange-red; legs and feet brownish; ADULT Y (nonbreeding): head, face and mantle olive-yellow, mantle streaked dusky; wing-coverts and flight feathers with pale yellow edges; rump, tail and chin to chest pale yellow; belly white. Bill horn; eyes reddish to brown; ADULT X (breeding): yellow-green above, mantle and back streaked dusky; narrow yellow superciliary stripe; breast bright yellow; belly white; bill horn. ADULT X (non-breeding): similar to non-breeding Y, but more olive above and paler yellow from chin to chest. SIZE (Western Cape): wing, Y (n ¼ 15) 78–89 (847), X (n ¼ 7) 76–84 (800); tail, Y (n ¼ 4) 515– 575 (535); bill, Y (n ¼ 4) 175–21 (189), X 14–19; tarsus, Y (n ¼ 4) 23–26 (245), X 19–25. WEIGHT: (Western Cape) Y (n ¼ 17) 31–455 (373), X (n ¼ 6) 285–33 (304). IMMATURE: similar to ad. X in non-breeding dress. NESTLING: hatches naked, skin orange-pink, sparse white down on feather tracts; bill pinkish horn, gape flange whitish yellow; later more dark reddish. P. v. nigrifrons Cabanis: E Cape, SE Mpumalanga, N and W Natal, and W Swaziland. Y differs from nominate race in having deeper saffron wash around mask; X darker than nominate. SIZE: wing, Y (n ¼ 9) 79–845 (825), X (n ¼ 4) 725–76 (750); tail, Y (n ¼ 3) 48–515 (498); bill, Y (n ¼ 3) 155–17 (162); tarsus, Y (n ¼ 2) 25. WEIGHT: (Natal) Y (n ¼ 11) 275–335 (307), X (n ¼ 3) 24–26 (246). P. v. tahatali Smith (includes ‘peixotoi’, introduced to Sa˜o Tome´): Zimbabwe, SE Botswana, SW Mozambique, E Swaziland, NE Natal and Northern Province. Y differs from nominate race

in having hind-crown, nape and mantle paler yellow, rump bright yellow; X paler than nominate. SIZE: wing, Y (n ¼ 26) 71–845 (797), X (n ¼ 26) 62–81 (727); tail, (n ¼ 2) Y 47, 505, X 465, 485; bill, (n ¼ 2) Y 145, 16, X 15, 155; tarsus, (n ¼ 2) Y 225, 25, X 22, 235. WEIGHT: Y (n ¼ 73) 205–35 (281), X (n ¼ 35) 196– 315 (245). ˆ i, Mozambique. Similar to P. v. shelleyi Sharpe: Zambia, Malaw tahatali but more golden yellow; differs from all races in having greater contrast between black and yellow of wing coverts. SIZE: wing, Y (n ¼ 12) 74–785 (760), X 68–73; tail, Y 45; bill, Y 175; tarsus, Y 24. WEIGHT: 1 Y 26. P. v. caurinus Clancey: S Angola, Namibia, Botswana and N Cape (Gordonia and Kuruman districts). Differs from tahatali in having narrower frontal black, mantle and scapulars paler and greener, rump yellow. SIZE: wing, Y (n ¼ 12) 76–81 (782), X (n ¼ 12) 70–75 (718); tail, Y (n ¼ 5) 45–51 (49), X (n ¼ 1) 45; bill, Y (n ¼ 3) 145–16, X (n ¼ 1) 145. WEIGHT: Y (n ¼ 2) 28, 28. P. v. finschi Reichenow: coastal Angola. Y paler than caurinus, mantle and scapulars greyer. SIZE: wing, Y (n ¼ 3) 75–765 (758); tail, Y (n ¼ 3) 45–49 (471). TAXONOMIC NOTE: see under P. vitellinus.

Field Characters. Length 125–14 cm. Distinguished from Lesser Masked Weaver P. intermedius by red eye and pinkish brown legs. Y has mainly yellow crown (black confined to narrow band on forehead), black mask ends in point on throat (rounded in Lesser Masked). Smaller than Village Weaver P. cucullatus, with unspotted greenish back; Y of southern race spilonotus of Village Weaver has yellow forehead. X and non-breeding Y have olive-brown back with dark streaks, red-brown eye, yellow edges to wing and tail feathers; face and throat pale yellow, breast buffy, belly white. Voice. Tape-recorded (22, 72, 86, 88, 99, 104, B, F, LEM, PAY). Song a series of buzzy swizzling and churring phrases, often introduced by chattering and chuckling notes, ‘chop-chop-cha-cha-cha-cukcukcukcuk zwurrrrrrrrswizzzzzz-zweeeeee’, or ‘chucky-chucky-chucky-zeeeeeeetrrrrrrrr-zurrrrrrr’, interspersed with harsh ‘dzit’, forceful ‘tsee-tsee-tsee’ and low chuckling; lacks wheezy final note of Village Weaver. Alarm and contact calls, harsh ‘chik’; X has soft contact note when arriving at or leaving nest; young beg with high-pitched ‘chi’. General Habits. Inhabits open savanna, semi-arid scrub, riverine thickets, woodland edges, farmland with clumps of trees, plantations of exotics, urban and rural gardens; in ^ Zaı¨re, arid areas with thorn-bushes and trees; in Malawi, clearings around human habitation; in Mozambique, woodland, savanna, clearings and cultivated land; riverine woodland in arid and semi-arid savanna (Angola). Avoids evergreen forest and coastal bush. Common throughout arid regions where water is available. Altitudinal range of ˆ i, to 2600 m, 330–1300 m in Zambia, up to 1600 m in Malaw along rivers, in Lesotho. Gregarious; forages in small flocks, taking seeds from ground and grass stems, gleaning insects from leaves and bark of trees, extracting beetle larvae from seeds of Acacia caffra; in Botswana, large flocks of non-breeding birds forage in semi-arid savanna. Roosts in reeds or trees; gathers in nearby trees up to 45 min before sunset, singing and chattering; most fly into roost 10 min before sunset.

Ploceus velatus Leaves roost c. 15 min before sunrise, hopping to tops of reed or trees and fanning out in different directions in small groups. Roosts large (1000–2000 birds) or small. In SW Cape (winter rainfall) breeds and moults 1 month earlier than in Gauteng, formerly S Transvaal, (summer rainfall), indicating annual cycle has advanced during the 50 years of range expansion into SW Cape. In Botswana a partial migrant – ad. YY sedentary, while juveniles move away in winter, as do many XX during drought conditions; large-scale dry-season influx into Okavango basin, while in Kalahari sandveld species appears to be a wet-season migrant. In the moister east it is probably mostly sedentary. Bird ringed in Johannesburg was shot near Kimberley, 466 km away. Food. Seeds, especially of grasses, also of elm, cosmos, Aleppo pine Pinus halepensis. Flower parts, especially ovaries and buds, e.g. of peach and apricot Prunus and driedoring Rhigozum trichotomum. Stamens of African marigold Tagetes erecta. Nectar of flat-flowered aloe Aloe marlotthi and many other aloe spp., silken oak Grevillea robusta, Cape honeysuckle Tecomaria capensis, Schotia brachypetala, Eucalyptus sideroxylon and Mexican blood trumpet Phaedranthus buccinatorius. Fruits of common taaibos Rhus pyroides, satsuma plum Viscum rotundifolium and puzzle bush Ehretia rigida. Insects, including beetles, termites, mayflies, larvae of lawn caterpillars Spodoptera capicola. Table scraps, including porridge and bread. Young fed caterpillars, termites and other insects. Breeding Habits. Polygynous, in colonies of 1 or more YY. One Y may attract 2–3 XX simultaneously and in one breeding season up to 12 XX successively. In urban areas colonies usually have 1 Y, in rural areas 1 or more. Y displays from nest by fanning the wings and spreading tail slightly while singing. Probably multi-brooded. NEST: kidney-shaped (A), constructed with strips from leaves of grass reed, palm, vaderlandsche bamboo or velvety heads of vasey grass Paspalum urvillei; Sanseviera fibre strips; spikes of grass Lophacme; once of pine needles. Grass blades 160–350 long, width 1–4. Ceiling of leaves, including eucalyptus or acacia, grass seed heads; large entrance on underside. Length 145–150, height 100, width 50, entrance diam. 30  40; dry weight 89. Rarely, double nests. Built by Y; if a X accepts nest, he adds a short entrance tube of c. 20 mm. X lines accepted nest with grass inflorescenes, leaves and feathers. Twigs holding nests are often stripped of their leaves; in reeds nests are attached at the side to 1 reed or slung between 2 reeds. Sited at end of tree branch, in bush, reeds or palm, sometimes over water. Sites provided by Acacia albida, A. erubescens, A. karroo, Terminalia sericea, Rhus quartiniana, palm Phoenix reclina; exotic mesquite Prosopis, Pinus, Eucalyptus, Salix, reeds Typha and Phragmites, and bamboo. Nests also suspended from barbed wire fence, telephone wire or telephone strut line; sometimes placed near nests of wasp Polistes and Icaria, and in trees with eagle nests, especially Wahlberg’s Eagles Aquila wahlbergi (Ewbank 2002). Y demolishes old nests to make space for new ones. At end of breeding season nests are often not destroyed, and may be used for breeding by other birds, especially Red-headed Finch

Amadina erythrocephala, also Zebra Waxbill Sporaeginthus subflavus, Blue Waxbill Uraeginthus angolensis, Cape Sparrow Passer melanurus, Scaly-fronted Weaver Sporopipes squamifrons, and climbing mice Dendromys. EGGS: 1–6, usually 2–4, laid at daily intervals; Angola ˆ i (n ¼ 20) 1–4 (195); Zimbabwe, (n ¼ 13) 1–4 (22); Malaw ˆ i (n ¼ 560) 1–5 (24); Zimbabwe (n ¼ 28) Zambia, Malaw 2–4 (29); Free State (n ¼ 15) 2–3 (25); Cape Province (n ¼ 37) 1–4 (29); southern Africa (n ¼ 196) 2–4 (26). Normal to elongated oval; white, pink, pale green or blue, plain, or spotted, speckled or blotched with grey, brown, red or purple. SIZE (n ¼ 82, Zimbabwe) 175–244  124– 155 (202  140); (n ¼ 72, Mpumulanga Prov.) 186– 212  132–151 (198  143); (n ¼ 418, southern Africa) 175–266  124–196 (212  145). WEIGHT: 26 g. ˆ i, LAYING DATES: Angola, Nov; Zambia, Sept–Feb; Malaw Jan–Apr (mainly Feb–Mar); Botswana, Sept–Mar (peak Nov); Zimbabwe, July–Apr (mainly Oct–Feb); southern Mozambique, Dec; South Africa, July–Apr (mainly Sept–Feb); Karoo, May, Aug, Oct, Dec (depending on rains); W Cape, Aug–Oct. INCUBATION: by X 59% of day spent incubating. Bouts average 70 min. in nest, 48 min. out of nest (n ¼ 306). Period: 12–13 days (Oschadleus 1996); 14 days (Schoeman 1995). DEVELOPMENT AND CARE OF YOUNG: downy feathers lost after 6 days. Feathers protrude from pins on day 10 (Schoeman 1995). Fed by X; rarely, Y feeds chicks in nest or fledged young. Young in nest c. 16–17 days. X breeds from 1 year; Y from 2nd year.

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PLOCEIDAE

BREEDING SUCCESS/SURVIVAL: regular host to Diederik Cuckoo Chysococcyx caprius (66% of nests, South Africa; 63%, Zambia). 63% and 55% fledging success in 2 seasons (41 and 67 nests respectively: Hunter 1961); 50% fledging success in 30 nests (Schoeman 1995); 7 of 8 clutches hatched (875% success: Oschadleus 1991). Predators of eggs and nestlings include egg-eating snake Dasypeltis scabra, boomslang Dispholidus typus, Grey Hornbill Tockus nasutus, Gabar Goshawk Micronisus gabar, African Harrier Hawk Polyboroides typus, Yellow-billed Kite Milvus migrans parasitus and vervet monkey Cercopithecus aethiops. Fiscal Shrike Lanius collaris took fledged chicks. Predators of adults include domestic cats, ringhals Sepedon hemachates, African Marsh Harrier Circus ranivorous, Little Sparrowhawk Accipiter minullus, Lanner Falco biarmicus, Pearlspotted Owlet Glaucidium perlatum, Spotted Eagle Owl Bubo africanus, Barn Owl Tyto alba and Cattle Egret Bubulcus ibis. Adult X died after being suspended by

strand of grass of nest. May drown when perching on algae on dams to feed on insects. Infected with blood parasites Haemoproteus (6 of 61 birds), Leucocytozoon (4 of 61) and Trypanosoma (1 of 61); microfilaria also recorded. Mite Pellonyssus reedi commonly in nests (mostly when chicks ready to fledge). Tick Hyalomma rufipes recorded. Larvae of fly Passeromyia heterochaeta suck blood of nestlings. YY needs hundreds of grass stems 20–30 cm long to build a multiple nest, and nesting may be constrained when such stems used up (Ewbank 2002). Greatest time between ringing and recovery, 12 years 2 months; median time, 10 months; a captive bird lived >23 years. Annual survival rate estimated at 506% in Gauteng, 57% for southern Africa. Key References Howman, H. R. G. and Begg, G. W. (1987, 1995), Hunter, H. C. (1961), Schoeman, S. (1995).

Plate 8

Ploceus vitellinus (Lichtenstein). Vitelline Masked Weaver. Tisserin vitellin.

(Opp. p. 91)

Fringilla vitellina Lichtenstein, 1823. Verz. Doubl., p. 23, Senegambia. Forms a superspecies with P. reichardi, P. katangae, P. ruweti and P. velatus. Range and Status. Endemic resident. Mauritania, common along Senegal R., breeding visitor to coast near Nouakchott, occasionally north to Tagant. Gambia, uncommon at coast, widespread inland. Senegal, common south of 17
N, absent from Casamance area. Mali, general south of 17
N: Boucle du Baoule´, Bani, Mopti, Gourma, Sankarani, Bourem, Timbuktu. Burkina Faso, widespread to northern edge of savanna as at Ouahigouya. Ghana, northern savanna as at Tumu, Mole, also Accra plains. Togo, northern savannas south to Okpahoue´. Benin, northern savannas; Nigeria, Sokoto to L. Chad, south to Borgu Game Res. in W. Niger, Parc du W, Niamey, Niger R., Tahoua, Maradi, Agades, Monts Bagzans, Dabaga, Kori Tarare, Trabelladen, Tin Telloust, Timia. Cameroon, Poli, Touroua northwards. Chad, throughout S. Central African Republic, in N sector. Zaı¨re, in NE (Ishwa, Mahagi, Niarembe). Sudan, very common in Kordofan and Darfur; isolated population in Atbara; also in S (Bahr al Ghazal, El Buheyrat, W Equatoria) and east of Nile R., Blue Nile Prov., Sobat, W Jonglei and western E Equatoria. Ethiopia, in SE and S regions, and Rift Valley. Somalia, local in NW, and in S associated with major rivers. Uganda, West Nile, Moroto, east to Kidepo Nat. Park. Kenya, primarily in central region, north to L. Turkana; absent from E, L. Victoria basin and from coastal belt except for winter vagrants. Tanzania, Serengeti Nat. Park, Crater and Mbulu highlands, L. Manyara, Tarangire, lowlands around Arusha, Masai Steppe, Mkomazi Game Reserve. Common. Description. P. v. vitellinus (Lichtenstein): Senegal to W Sudan. ADULT Y (breeding): forehead with narrow black band; lores, cheeks and ear-coverts black; crown chestnut-brown passing into

golden yellow on nape and into greenish yellow on mantle, with faint dark central streaks; rump dull yellow; tail olive-green with pale yellow edges; chin and throat black; breast adjoining throat has chestnut-brown margin, sides golden yellow; belly, flanks, thighs and undertail-coverts golden yellow. Bill black; eyes red; legs pale brown, feet pinkish. ADULT Y (non-breeding): forehead and crown yellowish; nape olive-brown with faint streaks; mantle olive-brown with heavier streaking; rump dull yellow; tail

Ploceus vitellinus

148

PLOCEIDAE

BREEDING SUCCESS/SURVIVAL: regular host to Diederik Cuckoo Chysococcyx caprius (66% of nests, South Africa; 63%, Zambia). 63% and 55% fledging success in 2 seasons (41 and 67 nests respectively: Hunter 1961); 50% fledging success in 30 nests (Schoeman 1995); 7 of 8 clutches hatched (875% success: Oschadleus 1991). Predators of eggs and nestlings include egg-eating snake Dasypeltis scabra, boomslang Dispholidus typus, Grey Hornbill Tockus nasutus, Gabar Goshawk Micronisus gabar, African Harrier Hawk Polyboroides typus, Yellow-billed Kite Milvus migrans parasitus and vervet monkey Cercopithecus aethiops. Fiscal Shrike Lanius collaris took fledged chicks. Predators of adults include domestic cats, ringhals Sepedon hemachates, African Marsh Harrier Circus ranivorous, Little Sparrowhawk Accipiter minullus, Lanner Falco biarmicus, Pearlspotted Owlet Glaucidium perlatum, Spotted Eagle Owl Bubo africanus, Barn Owl Tyto alba and Cattle Egret Bubulcus ibis. Adult X died after being suspended by

strand of grass of nest. May drown when perching on algae on dams to feed on insects. Infected with blood parasites Haemoproteus (6 of 61 birds), Leucocytozoon (4 of 61) and Trypanosoma (1 of 61); microfilaria also recorded. Mite Pellonyssus reedi commonly in nests (mostly when chicks ready to fledge). Tick Hyalomma rufipes recorded. Larvae of fly Passeromyia heterochaeta suck blood of nestlings. YY needs hundreds of grass stems 20–30 cm long to build a multiple nest, and nesting may be constrained when such stems used up (Ewbank 2002). Greatest time between ringing and recovery, 12 years 2 months; median time, 10 months; a captive bird lived >23 years. Annual survival rate estimated at 506% in Gauteng, 57% for southern Africa. Key References Howman, H. R. G. and Begg, G. W. (1987, 1995), Hunter, H. C. (1961), Schoeman, S. (1995).

Plate 8

Ploceus vitellinus (Lichtenstein). Vitelline Masked Weaver. Tisserin vitellin.

(Opp. p. 91)

Fringilla vitellina Lichtenstein, 1823. Verz. Doubl., p. 23, Senegambia. Forms a superspecies with P. reichardi, P. katangae, P. ruweti and P. velatus. Range and Status. Endemic resident. Mauritania, common along Senegal R., breeding visitor to coast near Nouakchott, occasionally north to Tagant. Gambia, uncommon at coast, widespread inland. Senegal, common south of 17
N, absent from Casamance area. Mali, general south of 17
N: Boucle du Baoule´, Bani, Mopti, Gourma, Sankarani, Bourem, Timbuktu. Burkina Faso, widespread to northern edge of savanna as at Ouahigouya. Ghana, northern savanna as at Tumu, Mole, also Accra plains. Togo, northern savannas south to Okpahoue´. Benin, northern savannas; Nigeria, Sokoto to L. Chad, south to Borgu Game Res. in W. Niger, Parc du W, Niamey, Niger R., Tahoua, Maradi, Agades, Monts Bagzans, Dabaga, Kori Tarare, Trabelladen, Tin Telloust, Timia. Cameroon, Poli, Touroua northwards. Chad, throughout S. Central African Republic, in N sector. Zaı¨re, in NE (Ishwa, Mahagi, Niarembe). Sudan, very common in Kordofan and Darfur; isolated population in Atbara; also in S (Bahr al Ghazal, El Buheyrat, W Equatoria) and east of Nile R., Blue Nile Prov., Sobat, W Jonglei and western E Equatoria. Ethiopia, in SE and S regions, and Rift Valley. Somalia, local in NW, and in S associated with major rivers. Uganda, West Nile, Moroto, east to Kidepo Nat. Park. Kenya, primarily in central region, north to L. Turkana; absent from E, L. Victoria basin and from coastal belt except for winter vagrants. Tanzania, Serengeti Nat. Park, Crater and Mbulu highlands, L. Manyara, Tarangire, lowlands around Arusha, Masai Steppe, Mkomazi Game Reserve. Common. Description. P. v. vitellinus (Lichtenstein): Senegal to W Sudan. ADULT Y (breeding): forehead with narrow black band; lores, cheeks and ear-coverts black; crown chestnut-brown passing into

golden yellow on nape and into greenish yellow on mantle, with faint dark central streaks; rump dull yellow; tail olive-green with pale yellow edges; chin and throat black; breast adjoining throat has chestnut-brown margin, sides golden yellow; belly, flanks, thighs and undertail-coverts golden yellow. Bill black; eyes red; legs pale brown, feet pinkish. ADULT Y (non-breeding): forehead and crown yellowish; nape olive-brown with faint streaks; mantle olive-brown with heavier streaking; rump dull yellow; tail

Ploceus vitellinus

Ploceus vitellinus olive-brown with faint yellow edges; wings brown with yellow edges to remiges, coverts tipped with buff; faint yellow supercilium; cheeks olive-brown washed with yellow; chin and throat pale yellow; breast and flanks buff washed yellow; belly, thighs and undertail-coverts whitish; wings dark brown with narrow yellow margins on remiges, broad yellow edges on tertials and coverts, yellow tips to upperwing coverts; underwing cream with yellowish tinge. Bill brown, darker on upper mandible; eyes orange to red; legs brown, feet pinkish. ADULT X: in breeding plumage, like non-breeding Y; non-breeding X has forehead, crown and nape light olive-brown with faint central streaks; mantle olive-brown with broad, dark central streaks on feathers; rump dull buffy, unstreaked; tail olive-green; no obvious supercilium; cheeks olive-brown; chin and throat dull white with yellow wash; breast buffy; flanks grey; belly dull white; thighs buffy grey; undertail-coverts white; wings brown with narrow yellow edges to remiges, broader edges on coverts. Bill brown, upper mandible darker; eye red-brown to red; legs brown with pinkish feet. SIZE (7YY, 5XX): wing, Y 68–73 (701), X 63–67 (656); tail, Y 41–45 (436), X 40–45 (423); bill, Y 166–176 (172), X 156–174 (164); tarsus, Y 192–207 (198), X 172–186 (180). WEIGHT: 1 Y (Cameroon) 20. IMMATURE: like non-breeding X, but upperparts more olivegreen, with rump also olive-green, unstreaked; faint supercilium behind the eye only; underparts with yellowish buff wash. Bill horn-brown; eyes brown; legs light brown. NESTLING: not described. P. v. uluensis (Neumann): Ethiopia south to N Tanzania. Breeding Y has broader black on forehead, less extensive chestnut on crown. However, there is considerable individual variation, and validity of this race has been questioned (cf. Lynes 1924–1925, Granvik 1934). TAXONOMIC NOTE: generally treated as conspecific with P. velatus (Moreau 1960, White 1963, Hall and Moreau 1970, Dowsett and Dowsett-Lemaire 1993a, Borrow and Demey 2001, Stevenson and Fanshawe 2002), but separated by Wolters (1977– 1982), Sibley and Monroe (1990) and others. Clearly, the vitellinus and velatus groups are closely allied, with each other and with reichardi, katangae and ruweti. They are allopatric and may all be conspecific; reichardi and ruweti are particularly close (as are katangae and upembae which were considered to be separate species, P. katangae and P. upembae, by Louette 1987, but not by other authorities nor by ourselves), and indeed so are velatus and vitellinus. Within this complex, Dowsett and Dowsett-Lemaire (1993a) recognized 2 species (P. vitellinus/katangae/velatus and P. reichardi/ruweti) without linking them as a superspecies, and Sibley and Monroe (1990) recognized 5 species in 2 superspecies (P. velatus/P. vitellinus; and P. reichardi/P. katangae/P. ruweti). We treat these masked weavers as a superspecies of 5 species, less out of conviction, more as a means of drawing attention to the need for further research on vocal and other recognition characters and on mate selection, nest-building and other reproductive characters.

Field Characters. Length 14 cm. Y has smallest mask of any ‘masked’ weaver, black chopped off cleanly on lower throat, not extending onto breast, and only narrow band on forehead; also distinguished by red eye, chestnut crown paling to yellow nape. X and non-breeding Y told by red eye, indistinct supercilium, lack of yellow on underparts except as wash on throat and breast; bill smaller than other ‘masked’ weavers. Brown-eyed juv. probably not distinguishable from juv. Lesser Masked Weaver P. intermedius except by slightly thicker bill, and in older birds by grey feet (Zimmerman et al. 1996).

Voice. Tape-recorded (104, B, MOR). Song a hurried shrill chatter followed by a drier chatter, also weaker scrambled phrases of grating and whistled notes, e.g. ‘chick-chikleewer–chi’. Calls include grating ‘chick’, ‘chi-dit’ and double ‘wee-jer’, and sharp ‘pink’ or ‘zik’ in contact. Also has a swizzle song with rasps, whistles and chirps, see renditions in Barlow et al. (1997) and Zimmerman et al. (1996). General Habits. Inhabits dry savanna woodland and scrubland, generally below 1800 m, in the sahel belt and similar semi-arid regions. On ground moves primarily by hopping; can move rapidly sideways along a branch. Feeds on nectar by biting flower off at base, mandibulating it to extract nectar, and then discarding it (Short and Horne 1978). In Somalia, roosts in Acacia trees along with Ru ¨ ppell’s Weaver P. galbula, Lesser Masked Weaver P. intermedius, Chestnut Weaver P. rubiginosus and Red-billed Quelea Quelea quelea (>300 birds altogether: Clarke 1985). Resident; but forms nomadic flocks in non-breeding season and may be partially migratory (Chapin 1954); in Sudan, moves north in rainy season, whilst acquiring breeding plumage. Food. Seeds and insects; also nectar of Leonotis nepetifolia. Breeding Habits. Colonial nester but never in large colony, typically small groups of nests with single YY in attendance. Polygynous. Y sings long song during

149

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PLOCEIDAE

territorial encounters; he also sings in courtship, far from the nest; short song during nest advertisement display (Crook 1969); displays by waving spread wings whilst hanging below nest. NEST: oval to pear-shaped, with entrance below and a high lip inside, tightly woven, ‘like a new ball of string’ (Mundy and Cook 1974); very short entrance tunnel or none at all (A); ceiling made of grass heads of 2–3 different kinds such as Setaria, Pennisetum; floor lined with softer grass heads of Panicum, Chloris. Suspended from single central point of attachment; sited 2–5 m above ground along or near tip of pendulous branch; often far from water. Bird removes nearly all leaves around nest. In dry country, often nests in trees, e.g. Acacia, over pool in garden or desert oasis. EGGS: 2–4 (av. 24, n ¼ 29, Nigeria). Blue-white, white (sometimes tinged with pink) or greenish, with large and sparse, or fine and dense, reddish or violet flecks; occasional white eggs have fine black spotting. More than one X may lay in same nest (based on egg coloration: Granvik 1934). SIZE: (Nigeria, n ¼ 17) 185–216  120–145

(195  132); (Ghana) 198–205  133–140; (Mali) 203– 206  135; (Zaı¨re) 185–216  132–138; (n ¼ 58, Uganda) 205  137. LAYING DATES: Mauritania, Nouakchott, Mar, along Senegal R., July–Oct; Senegal and Gambia, July–Oct (breeding plumage, Dec); Mali, July–Oct; Ghana, Apr– Sept, also Dec, Mar; Burkina Faso, June–Sept; Togo, Aug (nesting June, Aug, Oct); Nigeria, June–Sept; Niger, June– July, Sept; Zaı¨re, June–Sept (post-nuptial moult Nov); Sudan, Aug–Oct (feeding young, Nov); Ethiopia, Feb–May (nesting, Mar); Uganda, June–Aug; Somalia, June–July, Sept, Nov–Dec; Kenya, primarily March–May, also July, Nov–Feb; Tanzania, Jan–June. INCUBATION: period 12–13 days. DEVELOPMENT AND CARE OF YOUNG: nestling period: 14–16 days. BREEDING SUCCESS/SURVIVAL: at 29 Nigerian nests, 25% of eggs produced fledged young. Key References (1943b).

Mundy, P. J. and Cook, A. W. (1974), Serle, W.

Plate 8

Ploceus reichardi Reichenow. Tanganyika Masked Weaver. Tisserin de Reichard.

(Opp. p. 91)

Ploceus reichardi Reichenow, 1886. Zool. Jahrb. 1, p. 150; Karema [east shore of L. Tanganyika]. Forms a superspecies with P. vitellinus, P. katangae, P. ruweti and P. velatus. Range and Status. Endemic resident. Tanzania, locally common in Karema, Namanyere and L. Rukwa; reported occurrence in Ruaha Nat. Park an error (N. E. Baker, pers. comm). Zambia, common along Saisi R. in NE (Leonard and Beel 1999).

Ploceus reichardi

Description. ADULT Y (breeding): forehead black, crown dark chestnut-brown to behind the eyes, hindcrown and nape golden yellow, merging into greenish yellow on mantle; rump golden yellow; tail yellowish green; chin, lores, cheeks and ear-coverts black; throat black, extending to a point on breast; breast chestnut-brown, colour extending onto flanks and front of belly; thighs, centre of belly, and undertail-coverts ochre-yellow. Wing feathers brown, primaries with narrow yellow edges, coverts with broad yellow edges and yellow tips. Bill black; eyes red; legs brown. ADULT Y (non-breeding): not known. ADULT X: upperparts greenish yellow with light streaking; underparts yellow, with orange wash on breast (Aspinwall and Beel 1998). Bill brown; eyes brown; legs brown. SIZE (13 YY, 1 X): wing, Y 67–72 (696), X 64; tail, Y 42–47 (448), X 40; bill, Y 16–18 (167), X 155; tarsus, Y 20–22 (210), X 19. IMMATURE: similar to ad. X. NESTLING: unknown. TAXONOMIC NOTE: see under P. vitellinus.

Field Characters. Length 13–15 cm. Breeding Y distinguished from Lesser, Southern and Katanga Masked Weavers P. intermedius, P. velatus and P. katangae by strong chestnut wash on breast and flanks. Similar to Lake Lufira Weaver P. ruweti but black on top of head confined to forehead, not extending to hind-crown. X and nonbreeding Y dull olive above, breast orange, rest of

underparts dull yellow, belly white, flanks with dusky brown wash. Voice. Not tape-recorded. Song is a typical weaver-like swizzling, similar to Southern Masked Weaver but with

150

PLOCEIDAE

territorial encounters; he also sings in courtship, far from the nest; short song during nest advertisement display (Crook 1969); displays by waving spread wings whilst hanging below nest. NEST: oval to pear-shaped, with entrance below and a high lip inside, tightly woven, ‘like a new ball of string’ (Mundy and Cook 1974); very short entrance tunnel or none at all (A); ceiling made of grass heads of 2–3 different kinds such as Setaria, Pennisetum; floor lined with softer grass heads of Panicum, Chloris. Suspended from single central point of attachment; sited 2–5 m above ground along or near tip of pendulous branch; often far from water. Bird removes nearly all leaves around nest. In dry country, often nests in trees, e.g. Acacia, over pool in garden or desert oasis. EGGS: 2–4 (av. 24, n ¼ 29, Nigeria). Blue-white, white (sometimes tinged with pink) or greenish, with large and sparse, or fine and dense, reddish or violet flecks; occasional white eggs have fine black spotting. More than one X may lay in same nest (based on egg coloration: Granvik 1934). SIZE: (Nigeria, n ¼ 17) 185–216  120–145

(195  132); (Ghana) 198–205  133–140; (Mali) 203– 206  135; (Zaı¨re) 185–216  132–138; (n ¼ 58, Uganda) 205  137. LAYING DATES: Mauritania, Nouakchott, Mar, along Senegal R., July–Oct; Senegal and Gambia, July–Oct (breeding plumage, Dec); Mali, July–Oct; Ghana, Apr– Sept, also Dec, Mar; Burkina Faso, June–Sept; Togo, Aug (nesting June, Aug, Oct); Nigeria, June–Sept; Niger, June– July, Sept; Zaı¨re, June–Sept (post-nuptial moult Nov); Sudan, Aug–Oct (feeding young, Nov); Ethiopia, Feb–May (nesting, Mar); Uganda, June–Aug; Somalia, June–July, Sept, Nov–Dec; Kenya, primarily March–May, also July, Nov–Feb; Tanzania, Jan–June. INCUBATION: period 12–13 days. DEVELOPMENT AND CARE OF YOUNG: nestling period: 14–16 days. BREEDING SUCCESS/SURVIVAL: at 29 Nigerian nests, 25% of eggs produced fledged young. Key References (1943b).

Mundy, P. J. and Cook, A. W. (1974), Serle, W.

Plate 8

Ploceus reichardi Reichenow. Tanganyika Masked Weaver. Tisserin de Reichard.

(Opp. p. 91)

Ploceus reichardi Reichenow, 1886. Zool. Jahrb. 1, p. 150; Karema [east shore of L. Tanganyika]. Forms a superspecies with P. vitellinus, P. katangae, P. ruweti and P. velatus. Range and Status. Endemic resident. Tanzania, locally common in Karema, Namanyere and L. Rukwa; reported occurrence in Ruaha Nat. Park an error (N. E. Baker, pers. comm). Zambia, common along Saisi R. in NE (Leonard and Beel 1999).

Ploceus reichardi

Description. ADULT Y (breeding): forehead black, crown dark chestnut-brown to behind the eyes, hindcrown and nape golden yellow, merging into greenish yellow on mantle; rump golden yellow; tail yellowish green; chin, lores, cheeks and ear-coverts black; throat black, extending to a point on breast; breast chestnut-brown, colour extending onto flanks and front of belly; thighs, centre of belly, and undertail-coverts ochre-yellow. Wing feathers brown, primaries with narrow yellow edges, coverts with broad yellow edges and yellow tips. Bill black; eyes red; legs brown. ADULT Y (non-breeding): not known. ADULT X: upperparts greenish yellow with light streaking; underparts yellow, with orange wash on breast (Aspinwall and Beel 1998). Bill brown; eyes brown; legs brown. SIZE (13 YY, 1 X): wing, Y 67–72 (696), X 64; tail, Y 42–47 (448), X 40; bill, Y 16–18 (167), X 155; tarsus, Y 20–22 (210), X 19. IMMATURE: similar to ad. X. NESTLING: unknown. TAXONOMIC NOTE: see under P. vitellinus.

Field Characters. Length 13–15 cm. Breeding Y distinguished from Lesser, Southern and Katanga Masked Weavers P. intermedius, P. velatus and P. katangae by strong chestnut wash on breast and flanks. Similar to Lake Lufira Weaver P. ruweti but black on top of head confined to forehead, not extending to hind-crown. X and nonbreeding Y dull olive above, breast orange, rest of

underparts dull yellow, belly white, flanks with dusky brown wash. Voice. Not tape-recorded. Song is a typical weaver-like swizzling, similar to Southern Masked Weaver but with

Ploceus katangae some of the longer wheezy notes at a higher pitch (Leonard and Beel 1996); also described as an unmusical mixture of chirps, twitterings and chattering, with occasionally a coarse high trill or a grasshopper-like churring, lasting 1–2 s (Beesley 1956). Calls, ‘chuck’, warbler-like ‘click’ or rapid ‘chut-chut’, slightly harsher than Southern Masked Weaver. General Habits. Poorly known. Inhabits papyrus swamps, reedbeds of Phragmites mauritianus and bulrushes mixed with bushes and Sesbania leptocarpa; sometimes occurs far from dry land. Forages in nearby woodland with termitaria, singly or in small groups. Actively hops along acacia branches; hangs upside down; on floodplains feeds in long grass under acacias. Chatters at nesting sites and when feeding in grass. Roosts in flocks in papyrus swamps. Moults in Dec and July. Food. Grass seeds. Breeding Habits. Colonial, with 4–30 nests, occasionally up to 150; territorial, probably polygynous. Y sings from tops of reeds; in courtship perches with plumage fluffed, wings partly opened and quivered, while facing X and singing (‘ruweti’, Ruwet 1965). NEST: retort-shaped, strongly and closely woven with thick walls, neat and firm. Built of 3 mm strips of grass

151

blades c. 70 mm long, with a thin ceiling of transverse strips of grass, bowl lined with stalks and panicles of fine grasses, and a few feathers; without any entrance tunnel, semicircular entrance 40  30, facing downwards; size 110 long, 90 high. Suspended from bushes standing in shallow water, also in Pennisetum purpureum on river bank in flooded grassland, attached to single grass stalk along one side, 15–24 m up; or suspended from ambatch Sesbania bushes, 18–28 m above ground level, and 20 nests. Y sings from tops of reeds; in courtship he perches

Key Reference Louette, M. and Benson, C. W. (1982).

General Habits. Occurs in swamps with reedbeds (Phragmites and Typha), interspersed with bushes including ambatch and Sesbania leptocarpa.

153

Ploceus heuglini Reichenow. Heuglin’s Masked Weaver. Tisserin masque´.

Plate 8

Ploceus heuglini Reichenow, 1886. Zool. Jahrb., 1, p. 147; Bahr el Ghazal.

(Opp. p. 91)

Range and Status. Endemic resident; perhaps nomadic in N. Senegal, Nioro-du-Rip, Tambacounda, Ferlo North, once at Ziguinchor. Gambia, near coast; scarce inland but probably occurs throughout (Barlow et al. 1997). Mali, southwards from 14
N, primarily in woodlands along the rivers (Lamarche 1981). Liberia, unconfirmed sight records at Wologozi, Monrovia, Harper and Buchanan (Gatter 1997). Ivory Coast, widespread in northern savanna (e.g. in Comoe´ Nat. Park), south to Lamto. Burkina Faso, in SW and SE regions. Ghana, uncommon but widespread: at Tamale, Vea dam, Mole Nat. Park, Wenchi, Kumasi, Kete Kratchi, suburbs of Accra, Legon and inselbergs of Accra Plains. Togo, Ebe´va north to Namoundjoga, commonest in Kara-Naboulgou area, uncommon nearer coast. Benin, northern savanna from Be´te´rou north to Parc du ‘W’. Niger, rare: Ayorou, Parc du ‘W’. Nigeria, widespread and not uncommon north of Benue R. and Niger R. as far north as Sokoto, Kano region and HadejiaNguru wetlands; south to Ilorin, Enugu and Serti. Cameroon, from Benue Plain northwards. Chad, from L. Chad southwards and in E at Zakouma Nat. Park. Central African Republic, Manova-Gounda-St Floris Nat. Park. Sudan, west of Nile R. at Kajo Kaji, north to Bendingilo, west to Southern Nat. Park. Zaı¨re, only in NE as Garamba Nat. Park. N Uganda, from West Nile to Teso district (and east to Kenya border – Stevenson and Fanshawe 2002). Kenya, regular at Kiminini, otherwise rare (Lokitaung, Trans Nzoia, Soy, Bungoma, L. Baringo, Kerio R., Kapenguria). At Lamto, Ivory Coast, density of 4 pairs per 50 ha in dry season, 7 pairs per 50 ha in rainy season (Thiollay 1971). Uncommon; locally fairly common; but distribution markedly patchy. Description. ADULT Y (breeding): forehead and crown golden yellow; nape greenish yellow; mantle olive-green with faint, narrow central streaks; rump greenish yellow; tail olive-green with narrow yellow margins; chin, throat, lores, cheeks and earcoverts black; breast yellow, with chestnut-brown wash at edges of black mask (central line extends down onto breast); flanks, thighs, belly and undertail-coverts yellow; wings olive-green with narrow yellow outer margins, broader yellow edges on coverts and yellow tips to upperwing-coverts. Bill black; eyes pale yellow; legs brown. ADULT Y (non-breeding): forehead, crown, nape and mantle greenish; rump greenish yellow; lores, cheeks and ear-coverts greenish; underparts yellow; tail and wings as in breeding Y. Bill brown, darker on upper mandible. ADULT X: forehead, crown,

Ploceus heuglini

? ? ?

nape and mantle dull olive-green, with some faint central streaks on feathers of mantle; rump paler green; tail dark olive-green; supercilium pale yellow; cheeks and ear-coverts pale greenish brown; chin and throat pale yellow; breast pale yellow with buffy wash; flanks buffy; belly grey; thighs and undertail-coverts yellowish; wings dark olive-green with narrow yellow edges to remiges, broader yellow margins to coverts. Bill with dark brown upper mandible, light brown lower mandible; eyes brown; legs brown. Non-breeding X has duller upperparts and paler underparts with ochre wash on breast; bill pale brown. SIZE (11 YY 2 XX): wing, Y 72–76 (741), X 67, 70; tail, Y 41–47 (435), X 41, 41; bill, Y 183–197 (190), X 160, 191; tarsus, Y 190–204 (199), X 189, 191. WEIGHT: Y (Nigeria, May–June, n ¼ 5) 223–265 (239); 1X (Mali) 35; unsexed (Nigeria, n ¼ 12) 212–265 (232), (Ghana) 230, 254; immature (Nigeria) 206, 250. IMMATURE: like non-breeding X, except for buff rather than yellow edges to wing feathers and wing-coverts. NESTLING: not described.

Field Characters. Length 12–14 cm. Breeding Y has pale eye, black mask extending to narrow point on breast, crown entirely golden yellow (no black on forehead), plain

156

PLOCEIDAE

green back. Does not meet similar but much larger Speke’s Weaver P. spekei, which has streaked black and yellow back; Lesser Masked Weaver P. intermedius has pale eye but head black from forehead to mid-crown. Somewhat chunkier and shorter-tailed than widely-overlapping Vitelline Masked Weaver P. vitellinus (Barlow et al. 1997); further separated by pale eye (both sexes), lack of chestnut wash on crown (Y), plainer, more olive upperparts and yellower underparts (X and non-breeding Y). X very like X Lesser Masked but has pink, not grey legs. Voice. Tape-recorded (104, B, BRU, CHA, GRI). Chattering song of dry and some whistled notes lasting 4–6 s, accelerating into central nasal trill, ending with more chatters and dry trills ‘chukakakakaka-kew-kew-kewkwee-kwee-kway-kway-jarrrrrrrrrrr-chatatatatatatturrrrrr’; repertoire very varied. X also sings, ‘seet seet seet sueet sureet chree chree chree chroo’ (Zimmerman et al. 1996). Calls include hard, grating ‘juck’ and soft ‘chaar’. General Habits. Occurs in scrub around cultivation, light woodland edges and dense savanna woodland (Gambia), coastal thickets and inselbergs (Ghana), and around farms and villages (Nigeria). Generally in drier and taller woodland than Vitelline Masked Weaver P. vitellinus, but in Nigeria locally sympatric with it and with Village Weaver P. cucullatus in soudanian zone from Sokoto east to L. Chad. Solitary or in pairs. Once in mixed-species flock foraging in woodland (Ghana: Greig-Smith 1978). Examines leaves and flowers, apparently searching for insects. Moves locally, away from breeding areas (Benin); in Nigeria widespread in N during rains but apparently moves south in dry season. In Zaı¨re, evidently a dry-season visitor, present in Uele district only to breed. Food. Insects and seeds. Diet at Lamto, Ivory Coast, estimated at 70% arthropods, 10% fruit and 10% seeds by weight (Thiollay 1971). 4 of 7 birds in Zaı¨re had eaten caterpillars; 3, other insects, 1 a spider, and 1 some seeds (Chapin 1954). In Mali, mainly insects (beetles, grasshoppers, caterpillars and dragonflies) in both adults and nestlings, also seeds (Pennisetum spp.) and grit. Breeding Habits. Solitary or colonial nester, most often 2–15 nests, widely spaced; forms mixed colonies with Vitelline Masked Weaver P. vitellinus (Gambia); polygynous. Y displays with tail quivering and wings trembling, hanging below nest, beating wings; copulates on branch near nest. Seldom near water. Returns to same nest site each year. NEST: kidney-shaped, untidy, small version of nest of Village Weaver P. cucullatus, with wide opening into tunnel (A), sometimes up to 150–200 long. Normally suspended at 2 points. Woven of wiry strips of leaves of grasses Schizachyrium exile, Andropogon, Chloris and Pennisetum; lined with downy flower heads of grasses, e.g. Imperata cylindrica, Andropogon, Chloris and Pennisetum. 1 nest built in a single day. Does not strip leaves from vegetation around nest. SIZE: 130 long, 110 high, entrance diam. 35 (Collias and Collias 1964); weight (n ¼ 3, Mali) av. 39.7.

Nests near Bamako, Mali, attached to telephone lines: single nests immediately next to support pole, or colonies of up to 35 within 1–2 m of pole, 5–6 m above ground; similar colonies in Senegal and Gambia. Nest woven throughout its width to wires; >80% of colonies close to nests of mud wasps Megachile spp. and paper wasps Polistes fastidiosus (which are placed in hollow insulators on the poles). Colonies of 33% of nests contained deserted eggs; 80% of single eggs were infertile, but only 40% of other clutches. Dead chicks suggest that colony may be abandoned when birds leave as a group. Despite defence of nest by ads, Black Crake Amaurornis flavirostra flew up to a nest on papyrus stalk and removed contents (Wooton 1990). Predators include Augur Buzzard Buteo augur, Yellow-billed Kite Milvus migrans and Gabar Goshawk Melierax gabar. Newly-built nests may be taken over by Northern Grey-headed Sparrows Passer diffusus. Empty nests in Nairobi used for roosting by Superb Starlings Spreo superbus and Willow Warblers Phylloscopus trochilus.

Key References Archer, G. and Godman, E. M. (1961), Gichuki, C. M. (1988).

Plate 9

Ploceus spekeoides Grant and Mackworth-Praed. Fox’s Weaver. Tisserin de Fox.

(Opp. p. 138)

Ploceus spekeoides Grant and Mackworth-Praed, 1947. Bull. Br. Orn. Club 68, p. 7; Ngariam, Teso, Uganda. Range and Status. Endemic resident, Uganda. Recently known only from interior of Lango (Lira, north of L. Kyoga) and Teso (Soroti, 1
430 N, 33
370 E) and considered Near-threatened. Formerly common at Aketa, Usuku (2
010 N, 33
580 E), Katakwi (1
550 N, 33
570 E) and Nariam (1
530 N, 34
080 E), north of L. Bisina (Mann 1976).

Ploceus spekeoides

Description. ADULT Y: forehead, crown and nape golden yellow with some black speckles on crown; mantle black, outer feathers with yellow fringes; rump yellow; tail light brown; chin, lores, side of head, ear-coverts and throat black; breast has pointed black bib in centre surround by chestnut-brown wash, otherwise golden yellow; flanks, belly, thighs and undertail-coverts golden yellow; wings brown, with yellow edges to remiges, broad yellow tips to upperwing-coverts and tertials. Bill black; eyes orange-red; legs brown. Not clear if Y has non-breeding dress. ADULT X: forehead, crown, sides of head, nape and mantle olive-green with dark central streaks; rump greenish yellow; tail olive-green; chin and throat pale yellow; breast with buffy wash; flanks, belly, thighs and undertail-coverts whitish washed with yellow; wings brown with pale yellow edges to remiges, broader pale margins on wingcoverts but less vivid than in Y. Bill blackish brown; eyes brown; legs brown. SIZE: (9 YY, 7 XX): wing, Y 73–77 (743), X 69–74 (715); tail, Y 38–40 (387), X 36–38 (370); bill, Y 206–224 (215), X 205–220 (210); tarsus, Y 204–219 (211), X 194–206 (202). IMMATURE: upperparts less heavily marked than in X; bill brown. NESTLING: undescribed.

Field Characters. Length 14 cm. A rare weaver with large head, heavy bill and short tail. Y has red eye, black mask ending in point on breast; back black with yellow feather edgings, rump yellow; resembles Speke’s Weaver P. spekei but confined to Uganda. Heuglin’s Weaver has small bill, pale eye, plain green back; Vitelline Masked Weaver P. vitellinus has chestnut crown, black mask cut off at throat; Lesser Masked Weaver has white eye, black forecrown. X

has crown and back with heavy dark steaks, rump and underparts yellow. Voice. Unknown. General Habits. Inhabits bushed and wooded grassland in swampy areas; in dry season apparently leaves breeding

160

PLOCEIDAE

nest). Ceiling of grass heads and some acacia leaves, floor lined with grass heads and an occasional feather. Outer shell mainly Setaria spp., inner lining of different grasses, particularly Rhynchelytrum and Panicum. Unused or incomplete nests torn down, fall to the ground below nesting tree. Y builds nest in 8–10 days, X adds lining once nest and Y accepted. SIZE: 180 long, 140 high, entrance diam. 50. Sometimes builds in Eucalyptus trees, most often in thorny acacias; 2–7 m above ground. EGGS: 1–4 (av. 21, n ¼ 110, Kenya). Bright blue, darker at blunt end, forming a cap; rarely with a few black spots. SIZE: (Somalia) 220–235  150–160 (227  157) (n ¼ 9, Kenya) 228–258  146–163 (232  156). LAYING DATES: Ethiopia, Apr–Sept, possibly also Dec– Feb; Somalia, May–June; Uganda, May–June; Kenya, Feb–July, Oct–Dec, peak months Mar–Apr; Tanzania, May, at Arusha, Feb–Apr. INCUBATION: by X only, period 11 days. DEVELOPMENT AND CARE OF YOUNG: fed by both Y and X, period 16 days.

BREEDING SUCCESS/SURVIVAL: urban colonies in Nairobi had av. brood size of 18 (n ¼ 53); nestling mortality 30– 35%, with larvae of fly Passeromyia heterochaeta a major factor; also mites and fleas in nests. At colonies in Kenya, >33% of nests contained deserted eggs; 80% of single eggs were infertile, but only 40% of other clutches. Dead chicks suggest that colony may be abandoned when birds leave as a group. Despite defence of nest by ads, Black Crake Amaurornis flavirostra flew up to a nest on papyrus stalk and removed contents (Wooton 1990). Predators include Augur Buzzard Buteo augur, Yellow-billed Kite Milvus migrans and Gabar Goshawk Melierax gabar. Newly-built nests may be taken over by Northern Grey-headed Sparrows Passer diffusus. Empty nests in Nairobi used for roosting by Superb Starlings Spreo superbus and Willow Warblers Phylloscopus trochilus.

Key References Archer, G. and Godman, E. M. (1961), Gichuki, C. M. (1988).

Plate 9

Ploceus spekeoides Grant and Mackworth-Praed. Fox’s Weaver. Tisserin de Fox.

(Opp. p. 138)

Ploceus spekeoides Grant and Mackworth-Praed, 1947. Bull. Br. Orn. Club 68, p. 7; Ngariam, Teso, Uganda. Range and Status. Endemic resident, Uganda. Recently known only from interior of Lango (Lira, north of L. Kyoga) and Teso (Soroti, 1
430 N, 33
370 E) and considered Near-threatened. Formerly common at Aketa, Usuku (2
010 N, 33
580 E), Katakwi (1
550 N, 33
570 E) and Nariam (1
530 N, 34
080 E), north of L. Bisina (Mann 1976).

Ploceus spekeoides

Description. ADULT Y: forehead, crown and nape golden yellow with some black speckles on crown; mantle black, outer feathers with yellow fringes; rump yellow; tail light brown; chin, lores, side of head, ear-coverts and throat black; breast has pointed black bib in centre surround by chestnut-brown wash, otherwise golden yellow; flanks, belly, thighs and undertail-coverts golden yellow; wings brown, with yellow edges to remiges, broad yellow tips to upperwing-coverts and tertials. Bill black; eyes orange-red; legs brown. Not clear if Y has non-breeding dress. ADULT X: forehead, crown, sides of head, nape and mantle olive-green with dark central streaks; rump greenish yellow; tail olive-green; chin and throat pale yellow; breast with buffy wash; flanks, belly, thighs and undertail-coverts whitish washed with yellow; wings brown with pale yellow edges to remiges, broader pale margins on wingcoverts but less vivid than in Y. Bill blackish brown; eyes brown; legs brown. SIZE: (9 YY, 7 XX): wing, Y 73–77 (743), X 69–74 (715); tail, Y 38–40 (387), X 36–38 (370); bill, Y 206–224 (215), X 205–220 (210); tarsus, Y 204–219 (211), X 194–206 (202). IMMATURE: upperparts less heavily marked than in X; bill brown. NESTLING: undescribed.

Field Characters. Length 14 cm. A rare weaver with large head, heavy bill and short tail. Y has red eye, black mask ending in point on breast; back black with yellow feather edgings, rump yellow; resembles Speke’s Weaver P. spekei but confined to Uganda. Heuglin’s Weaver has small bill, pale eye, plain green back; Vitelline Masked Weaver P. vitellinus has chestnut crown, black mask cut off at throat; Lesser Masked Weaver has white eye, black forecrown. X

has crown and back with heavy dark steaks, rump and underparts yellow. Voice. Unknown. General Habits. Inhabits bushed and wooded grassland in swampy areas; in dry season apparently leaves breeding

Ploceus nigerrimus areas. Little-known; a lack of recent data (Owiunzi and Arinaitwe 2001).

161

EGGS:

undescribed. Uganda, June–July. No other information. LAYING DATES:

Food. Seeds, probably also insects. Breeding Habits. Colonial. NEST: oval, roughly woven with no entrance tunnel. Outer shell and ceiling made of same selection of grasses. SIZE: 140 long, 80 high, entrance 45  50.

Key References Collar, N. J. and Stuart, S. N. (1985), Collias, N. E. and Collias, E. C. (1964).

Ploceus nigerrimus Vieillot. Vieillot’s Black Weaver. Tisserin de Vieillot.

Plate 10

Ploceus nigerrimus Vieillot, 1819. Nouv. Dict. Nist. Nat., nouv. ´ed., 34, p. 130; ‘Kingdom of Congo’.

(Opp. p. 139)

Range and Status. Endemic resident. Gambia, LaminPirang, Basse (breeding 1979). Guinea, Fore´cariah, Kindia, Macenta, Mafou. Guinea Bissau, R. Tombali. Sierra Leone, Freetown, Rokupr, Bo, Bintumane, Tembikunda, Gola Mts, Tingi Hills, Sefadu. Liberia, throughout, below 800 m. Ivory Coast, common along coast as at Yapo, Azagny Nat. Park, inland to Mont Sangle´ in W and Bossematie´ in E, local near water further north in Comoe´ Nat. Park and at Korhogo. Ghana, throughout forest zone, to edge of plains at Abokobi, near Dodowa and Odumasi Krobo. Togo, locally common in S guinean savanna from Tse´vie´ to Atakpame´. Benin, coastal region as at Lama Forest, L. Nokoue´. Nigeria, throughout coastal belt, inland to Oyo, Akure, Enugu, north to Serti on Mambilla plateau, Pandam Wildlife Park, north of Benue R. Cameroon, forest zone, north to Adamawa Plateau at Ngoundala (>150 localities as mapped: M. Languy, pers. comm. 2002). Central African Republic, Bangui, Dzanga, Berbe´rati, north to Marali. Mbini, Mont Alen Nat. Park, Gabon, coastal areas as Akanda, Ogooue´ delta, Gamba, inland to Lope´, Ipassa. Congo, Nouabale´-Ndoki Nat. Park, Odzala Nat. Park, Dimonika Res., Lower Kouilou basin and Conkouati Nat. Park. Zaı¨re, through central basin to Lendu Plateau in NE, south to L. Edward, Itombwe plateau; absent from SE Katanga, Maniema and W shores of L. Tanganyika. Rwanda, Akanyaru wetlands. Burundi, along L. Tanganyika, Kibira Nat. Park, L. Rwihinda, Ruvubu Nat. Park. Uganda, in S and W, north to Kabalega Falls, Bombo, Elgon, Sudan, Zande district and Aloma Plateau in SW near border with Zaı¨re. Kenya, local at Kakamega, Kapsabet, Nandi, wandering to Nambale, Ng’iya and Kisumu districts. Tanzania, isolated population at Mahali on L. Tanganyika, abundant on W shore of L. Victoria, Bukoba, Keragwe, old records from Gombe. Angola, Cabinda, south through Uige and Cuanza Norte to Gabela in Cuanza Sul, and in N Lunda Norte. Common to abundant, closely associated with human activity. Description. P. n. nigerrimus Vieillot: extreme SE Nigeria (Oban, Ikom, Serti, Butatong, Mambilla Plateau) to Uganda and W Kenya, south to Angola and Zaı¨re. ADULT Y: plumage wholly black; bill black, eyes golden yellow, legs dark brown; no eclipse plumage. ADULT X: forehead, crown and nape dull olive-green, feathers with narrow dark central streaks; mantle olive-green with broad central streaks; rump olive-green, unstreaked; tail

Ploceus nigerrimus

dark brown; lores, cheeks and ear-coverts buff; chin and throat pale yellow; broad buffy band across breast (in some birds tone more olive, which may extend to throat); flanks, thighs and undertail-coverts buff (sometimes olive wash on flanks); wings dark brown with narrow olive margins to remiges, pale buffy edges to wing-coverts; underwing greenish, mottled with black; Upper mandible dark brown, lower mandible horn-coloured; eyes pale yellow; legs brown. SIZE (12 YY, 12 XX): wing; Y 77–92 (833), X 75–82 (775); tail, Y 50–58 (536), X 47–55 (508); bill, Y 200–230 (213), X 191–210 (202); tarsus, Y 203–220 (213), X 190–210 (203). WEIGHT: Y (Zaı¨re, n ¼ 13) 31–42 (375), 1 X (Zaı¨re) 44; unsexed (Cameroon, n ¼ 7) 21–35 (308), Kenya, n ¼ 12) 270–330 (302), (Uganda, n ¼ 7) 248–352 (292). IMMATURE: like X, but forehead and crown unstreaked, underparts buffier with only central part of belly yellow, margins to wing-coverts buffy. Eyes dark grey initially, then brown in X, yellow in Y, before ad. plumage attained. NESTLING: corners of mouth bright yellow; bill pinkish grey, eyes dark grey, feet pinkish grey. P. n. castaneofuscus Lesson: Sierra Leone to E Nigeria (Calabar; Mambilla Plateau). ADULT Y: like nominate race but mantle,

Ploceus nigerrimus areas. Little-known; a lack of recent data (Owiunzi and Arinaitwe 2001).

161

EGGS:

undescribed. Uganda, June–July. No other information. LAYING DATES:

Food. Seeds, probably also insects. Breeding Habits. Colonial. NEST: oval, roughly woven with no entrance tunnel. Outer shell and ceiling made of same selection of grasses. SIZE: 140 long, 80 high, entrance 45  50.

Key References Collar, N. J. and Stuart, S. N. (1985), Collias, N. E. and Collias, E. C. (1964).

Ploceus nigerrimus Vieillot. Vieillot’s Black Weaver. Tisserin de Vieillot.

Plate 10

Ploceus nigerrimus Vieillot, 1819. Nouv. Dict. Nist. Nat., nouv. ´ed., 34, p. 130; ‘Kingdom of Congo’.

(Opp. p. 139)

Range and Status. Endemic resident. Gambia, LaminPirang, Basse (breeding 1979). Guinea, Fore´cariah, Kindia, Macenta, Mafou. Guinea Bissau, R. Tombali. Sierra Leone, Freetown, Rokupr, Bo, Bintumane, Tembikunda, Gola Mts, Tingi Hills, Sefadu. Liberia, throughout, below 800 m. Ivory Coast, common along coast as at Yapo, Azagny Nat. Park, inland to Mont Sangle´ in W and Bossematie´ in E, local near water further north in Comoe´ Nat. Park and at Korhogo. Ghana, throughout forest zone, to edge of plains at Abokobi, near Dodowa and Odumasi Krobo. Togo, locally common in S guinean savanna from Tse´vie´ to Atakpame´. Benin, coastal region as at Lama Forest, L. Nokoue´. Nigeria, throughout coastal belt, inland to Oyo, Akure, Enugu, north to Serti on Mambilla plateau, Pandam Wildlife Park, north of Benue R. Cameroon, forest zone, north to Adamawa Plateau at Ngoundala (>150 localities as mapped: M. Languy, pers. comm. 2002). Central African Republic, Bangui, Dzanga, Berbe´rati, north to Marali. Mbini, Mont Alen Nat. Park, Gabon, coastal areas as Akanda, Ogooue´ delta, Gamba, inland to Lope´, Ipassa. Congo, Nouabale´-Ndoki Nat. Park, Odzala Nat. Park, Dimonika Res., Lower Kouilou basin and Conkouati Nat. Park. Zaı¨re, through central basin to Lendu Plateau in NE, south to L. Edward, Itombwe plateau; absent from SE Katanga, Maniema and W shores of L. Tanganyika. Rwanda, Akanyaru wetlands. Burundi, along L. Tanganyika, Kibira Nat. Park, L. Rwihinda, Ruvubu Nat. Park. Uganda, in S and W, north to Kabalega Falls, Bombo, Elgon, Sudan, Zande district and Aloma Plateau in SW near border with Zaı¨re. Kenya, local at Kakamega, Kapsabet, Nandi, wandering to Nambale, Ng’iya and Kisumu districts. Tanzania, isolated population at Mahali on L. Tanganyika, abundant on W shore of L. Victoria, Bukoba, Keragwe, old records from Gombe. Angola, Cabinda, south through Uige and Cuanza Norte to Gabela in Cuanza Sul, and in N Lunda Norte. Common to abundant, closely associated with human activity. Description. P. n. nigerrimus Vieillot: extreme SE Nigeria (Oban, Ikom, Serti, Butatong, Mambilla Plateau) to Uganda and W Kenya, south to Angola and Zaı¨re. ADULT Y: plumage wholly black; bill black, eyes golden yellow, legs dark brown; no eclipse plumage. ADULT X: forehead, crown and nape dull olive-green, feathers with narrow dark central streaks; mantle olive-green with broad central streaks; rump olive-green, unstreaked; tail

Ploceus nigerrimus

dark brown; lores, cheeks and ear-coverts buff; chin and throat pale yellow; broad buffy band across breast (in some birds tone more olive, which may extend to throat); flanks, thighs and undertail-coverts buff (sometimes olive wash on flanks); wings dark brown with narrow olive margins to remiges, pale buffy edges to wing-coverts; underwing greenish, mottled with black; Upper mandible dark brown, lower mandible horn-coloured; eyes pale yellow; legs brown. SIZE (12 YY, 12 XX): wing; Y 77–92 (833), X 75–82 (775); tail, Y 50–58 (536), X 47–55 (508); bill, Y 200–230 (213), X 191–210 (202); tarsus, Y 203–220 (213), X 190–210 (203). WEIGHT: Y (Zaı¨re, n ¼ 13) 31–42 (375), 1 X (Zaı¨re) 44; unsexed (Cameroon, n ¼ 7) 21–35 (308), Kenya, n ¼ 12) 270–330 (302), (Uganda, n ¼ 7) 248–352 (292). IMMATURE: like X, but forehead and crown unstreaked, underparts buffier with only central part of belly yellow, margins to wing-coverts buffy. Eyes dark grey initially, then brown in X, yellow in Y, before ad. plumage attained. NESTLING: corners of mouth bright yellow; bill pinkish grey, eyes dark grey, feet pinkish grey. P. n. castaneofuscus Lesson: Sierra Leone to E Nigeria (Calabar; Mambilla Plateau). ADULT Y: like nominate race but mantle,

162

PLOCEIDAE

scapulars, rump, belly and undertail-coverts rich chestnut-brown. ADULT X: like nominate race, but upperparts brownish rather than olive-green, rump rufous, wings with buffy margins, underparts buffy yellow with rufous wash on flanks. Intergrades with nominate race in E Nigeria (Serle 1957). WEIGHT: (Liberia) Y (n ¼ 40) 27–39 (332); X (n ¼ 36) 24–32 (282). TAXONOMIC NOTE: castaneofuscus and nigerrimus were originally described as distinct species but have long been regarded as conspecific (Bannerman 1949, White 1963, Hall and Moreau 1970, Sibley and Monroe 1990, Dowsett and Forbes-Watson 1993, Elgood et al. 1994, Borrow and Demey 2001). Hybrids and intermediates are known in SE Nigeria, from Enugu to near Cameroon border (P. nigerrimus also hybridizes with P. cucullatus, with which it has a close, almost symbiotic, nesting association). However, voice, and some reproductive parameters, reported below, may differ between castaneofuscus and nigerrimus; moreover, both breed in one locality (Mambilla Plateau, SE Nigeria: Elgood et al. 1994). The possibility of their being separate species after all needs to be revisited.

Field Characters. Length 15–17 cm. Y wholly black (nominate race) or black with chestnut back, rump and belly (castaneofuscus). All-black Y extremely similar to Maxwell’s Black Weaver P. albinucha but larger, best distinguished by yellow, not whitish eye; told from X Cassin’s Malimbe M. cassini, which has dark eye, by longer, thicker bill. X and juv. Y are dingy olive except for a little yellow on throat and belly and streaks on back, eye pale; juv. X has dark eye. Most X malimbes, which forage in same trees, have some red on them, all have dark eyes. X Village Weaver P. cucullatus has bright yellow supercilium, breast and wing-edgings. Voice. Tape-recorded (P. n. nigerrimus, 104, B, C, CHA, ERA, GREG, McVIC, ZIM; P. n. castaneofuscus 104, B, CHA, GREG, PAY). In P. n. nigerrimus, sputtering and chattering at breeding colonies includes sharp ‘tsik’ or ‘tsuk’, singly or in irregular phrases, ‘tsuk-way-tsik’, ‘waytsik-tsuk-tsik’, and whistled notes, ‘sweet’, up-slurred ‘way-yee’ and phrase ‘tyo-tyo-tyo-tyeew’. Short song ‘chrrr-szee-zuit’ given during territorial encounters, during courtship away from nest, and when X enters nest. Distinct call used by Y during nest advertisement display. Sonagrams of song and aggressive ‘chak’ calls in Crook (1969). Voice of P. n. castaneofuscus similar, short ‘tyup’, ‘jup’, ‘pik’ and ‘zeet’ interspersed with rapid thin screechy chatters, also some sweet whistled notes, ‘weeyoo’, ‘woo’ and ‘wee-weeyoo’. General Habits. Inhabits forest clearings, often in villages, wooded areas and high grass savannas; although not found within forest; distribution in Uganda coincides with areas in which forest habitat occurs (Hall 1968). In villages up to 1300 m (Cameroon) and 2000 m (Zaı¨re); generally in high rainfall regions of Kenya, between 1000 and 2000 m. Gregarious at all times. Moves far from colony to forage in savanna and gallery forest; regularly joins mixed-species flocks of insectivores foraging in more open areas of forest; hawks insects. Foraging parties in coffee plantations associate with Dark-backed Weaver P. bicolor (Angola: Heinrich 1958). Nigerian birds search areas where squirrels Funisciurus anerythrus have been feeding, collect outer fibrous layers of oil palm nuts which squirrels

discard; also exploit nuts dislodged by Red-headed Malimbe Malimbus rubricollis while feeding (Pettet 1969). During dry season, plumage coloured by pollen of plants at which the birds feed (Ouagadougou, Burkina Faso: Thonne´rieux et al. 1989). Roosts in tall elephant grass, even while actively building nests at a nearby colony. Often roosts with Village Weavers. Wet-season roosts are generally in broad-leaved trees and dry-season ones in high grass in swampy areas. Moults in Liberia in Nov–Jan; occasional XX moult while breeding. Y birds lightest in weight at end of moult (Chapman 1995). Anting occurs in captive birds. Food. Mainly insects (grasshoppers, winged ants including Oecophylla, winged termites, caterpillars and butterflies); also seeds, fruit, berries (e.g. Rauwolfia) and oil-palm Elaeis guineensis fruits; nectar. In Nigeria, during dry season eats oil palm fruits, wild figs, seeds and maize; insects (mantids, grasshoppers and caterpillars) taken are primarily for feeding chicks (Din 1991). Seeds of rice and maize. Breeding Habits. Colonial, gregarious, territorial, polygynous, with up to 5 XX in 1 Y territory. Breeding activity at colonies appears to coincide with periods of rainfall. Some XX in a Ugandan colony synchronised nesting, others did not (Hall 1970a). Forms mixed colonies with Village Weaver in tall trees in settlements and villages; the 2 occasionally hybridize. In Gabon, Village Weaver appears to select site and start colony, and is then joined by Vieillot’s Black Weaver (Brosset and Erard 1986). Sometimes forms mixed colonies with Orange Weaver P. aurantius in Nigeria; in Owerri province, almost all Village Weaver colonies have at least some Vieillot’s Black Weaver nests in them, perhaps as few as 2% (Marchant 1942). On Ife university campus, all 34 Vieillot’s Black Weaver colonies were mixed with Village Weavers; at 16, Village Weaver was more abundant, at 9, Vieillot’s Black Weaver; Village Weaver occupied middle and upper branches, Vieillot’s Black Weaver the lower branches (Din 1991). In Gabon, such mixed colonies often in parasol trees Musanga (Brosset 1974), in Cameroon in mangos Mangifera indica or sago-palms Pachylobus edulis (Germain et al. 1973). Mixed colonies in Uganda are common, but many suitable sites unoccupied. Y Vieillot’s Black Weaver does not respond to bursts of activity by Y Village Weavers in same colony. Throughout forest zone, colonies are generally in villages and clearings, often over water or close to swampy areas. Pure colonies occur along rivers, in reeds, bulrushes, bamboos or tall grasses. May be double-brooded. Territory size varies from 02 to 1 m2; each territory contains 1–6 nests, 27–52 cm apart. Upsurge in territorial defence when Y returns to territory after rain. Territory is defended by Y using threat posture with head feathers ruffled, bill slightly open, wings quivering, and tail alternately fanned and closed; this often develops into an ‘aggressive dance’ or lunging match. In mixed colonies, Y defends territory against neighbouring Village Weavers, who have a much more static display. Other YY flying through territory are not chased, but a perched intruder is attacked. Fighting YY may fall to ground or into water. X

Ploceus nigerrimus

defends only her nest, and may vigorously repulse Y or intruding X at nest entrance. Most active colonies seem to attract roving parties of XX. Every arrival of a X is greeted by bursts of song from attendant Y. In advertisement display, Y hangs below nest, quivers rather than flaps wings below the body axis, with tail turned away from nest (A); wings are raised higher and move faster than in Village Weaver. Similar display by bird hanging under a branch indicates a future nest-building site. Y also displays on a branch, drooping and fluttering wings with head raised and breast feathers fluffed, giving short chirps. If X present, Y may move rapidly from 1 branch to another, or fly to nest to hang below it. Short-distance pursuits of X seem to be special feature of the courtship of castaneofuscus (Crook 1963a). X soliciting copulation moves head, makes short sideto-side hops, quivers wings and tail, raises bill and gives a distinctive call; finally she crouches, fans tail, lifts head up and lowers wings. Y approaches with back feathers ruffled, head lowered, and gives hissing call. He flaps wings whilst mounting, and touches X’s head with bill. Y may mount 3–4 times in succession; some copulations are disrupted by neighbouring birds. After copulation, X typically enters nest and Y hangs at entrance and displays. In general, building activity peaks before 11h00 and colony is almost deserted from 12h00 to 15h00; activity resumes in late afternoon, when most copulations seem to take place.

NEST: oval, rough exterior, very like that of Village Weaver but lacks spout at entrance, which is large and semicircular. Woven from grass or strips of palm leaves; lined with Panicum and Sporobolus grasses, also cotton (Nigeria). 1 colony of c. 100 nests was in maize field; nests at top of stalks recently stripped of seeds. SIZE: (P. n. castaneofuscus): 125 long, 95 high, entrance diam. 40. Nest of P. n. nigerrimus may be larger: 130–150 long, 115 high, entrance diam. 45 (Collias and Collias 1964). Nests in Liberia 1–2 m above water, also 6–8 m up in forest tree. Sited also in palms, bamboos, trees in swamps, tall elephant grass, and bulrushes. Nests in trees often up to 25 m above ground. X sometimes carries nesting material, but Y does all building of outer shell; ceiling inside nest chamber usually made of leaves, of e.g. Eucalyptus, Azadirachta, Delonix and bamboo. Y may complete nest in 6–11 h (n ¼ 5, av. 9 h), with 3-day interval between construction of successive nests. X rarely visits nest under construction; she usually begins lining immediately after copulation, then she starts to roost in nest. Most material collected within 50 m; Y may steal material from neighbours’ nests. One Y stole nest material from, and ultimately drove off, pair of Spectacled Weaver P. ocularis building in same palm; nest construction much faster than by P. ocularis (Diesselhorst 1971). Branch on which nest is built is stripped of leaves. Old nests may be demolished by Y c. 7–19 days after construction; demolition takes no more than 30 min. and appears to free space for new construction, so that no more than 5 nests in a territory at a time. At a mixed colony in Nigeria, 20 YY took over and modified some nests in a nearby colony of Red-headed Quelea Quelea erythrops but later abandoned them (Din 1994). EGGS: 2, plain blue; sometimes different parts shaded darker and lighter; rarely with a few purplish spots or blotches at thick end. SIZE: (Cameroon, n ¼ 40) 211–254  150–174 (239  161); (Nigeria, n ¼ 37) 183– 261  111–170 (229  157); (Zaı¨re) 220–262  154– 170. Re-lays if eggs removed from nest (Eaglesfield 1969). WEIGHT: 24–35 (29, n ¼ 32, Nigeria). LAYING DATES: Gambia, Jan; Guinea, Feb; Liberia, Feb– Apr, Aug–Nov (in breeding condition all year); Ghana, Jan–July; Togo, Feb, Sept–Oct; Benin, (colony active Apr–May); Nigeria, May–Aug (castaneofuscus), Feb (nigerrimus) (nest-building throughout year); Cameroon, all months except Aug, peak activity in lowlands in June– July; Gabon, Dec–Feb, Apr, July; Central African Republic, July–Aug; Congo, Jan–Feb, Sept, Nov–Dec; Zaı¨re, Nov in S, April and Aug–Sept in savanna; Lukolela, July–Feb, Itombwe, Feb, July, Aug, Nov (probably all year); Sudan, Aug, Nov; Uganda, all months except Aug and Nov, with peaks in Feb–Apr and Sept; Kenya, Kakamega, July–Sept; Angola, Oct–Mar. INCUBATION: by X only; some eggs ejected from nests during incubation. Period unknown. DEVELOPMENT AND CARE OF YOUNG: fed primarily by X, but Y may contribute, particularly late in season as nesting activity declines. Nestling period unknown. Once young leave nest, they accompany X while foraging and do not return to colony.

163

164

PLOCEIDAE

BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius (Ghana: Macdonald 1980). Colony in Uganda destroyed by party of Black-and-white Casqued Hornbills Bycanistes subcylindricus (the site not re-occupied: Hall 1970b). Nests robbed by Pied Crow Corvus alba, Senegal Coucal Centropus senegalensis and African

Plate 9 (Opp. p. 138)

Harrier Hawk Polyboroides typus (Macdonald 1980, Verheyen 1955); also by boomslang Dispholidus typus (Elgood and Ward 1963) and egg-eating snake Dasypeltis (Pitman 1958). Key References Crook, J. H. (1963), Din, N. A. (1992a, b, c, d), Hall, J. R. (1970a, b, c), Verheyen, R. (1955).

Ploceus cucullatus (Mu ¨ ller). Village Weaver; Black-headed Weaver; Spotted-backed Weaver. Tisserin gendarme. Oriolus cucullatus P. L. S. Mu ¨ ller, 1776. Natursyst., Suppl., p. 87; Senegal. Range and Status. Endemic resident and local intraAfrican migrant. Introduced in Mauritius, Re´union and West Indies (Hispaniola, Martinique). Widespread in subsaharan Africa except arid SW and NE, especially along watercourses, lake shores and coast lines. Generally abundant, but local and seasonal in dry areas on periphery of range. Centres of abundance are clearings in tropical rain forests in central and W Africa, and moist open country. Northern limits: Mauritania (Tagant), 18
N; Mali, delta region around Tombouctou, c. 17
N; Niger, (Tahoua, Tanout), 15
N; Chad, (Ouadi Haddat), 14
320 N; Sudan (where uncommon in E), 14
N; Eritrea, c. 16
N. Absent from desert regions of E Ethiopia and from Somalia north of 5
N; abundant along Jubba and Shabeelle river systems in S Somalia, and reappears in extreme S near Kenya border. Extreme N Kenya at Moyale and Daua R. on Ethiopian border, otherwise absent from arid regions north of Mt Elgon and Kapenguria and east of Tana R. drainage (except on coast). Southern limits: Angola (except central plateau) south to Mossamedes and Huı´la; Namibia, mainly in Caprivi, extending west along Okavango R.; in Botswana restricted to Okavango region and Limpopo drainage; Transvaal except SW, absent from western Highveld and Bushveld; Free State, mainly in moist E, scattered records elsewhere; rare in lowlands of Lesotho (ButheButhe, Leribe, Maseru, Mohale’s Hoek); E Cape Prov. west to Port Elizabeth and lower Gamtoos R. Occurs on Bioko, Sa˜o Tome´ (probably recently introduced: Christy and Clarke 1998); Pemba, Zanzibar, Mafia. Density, Acacia sevannas, Swaziland, 3, 20 and 51 birds per 100 ha. Description. P. c. cucullatus (Mu ¨ ller): Mauritania to Chad, south to Bioko, N Gabon and NW Zaı¨re. ADULT Y (breeding): head black; nape and hind neck chestnut; mantle yellow with 2 lateral black bands converging posteriorly on back to form distinct ‘V’; rump yellow, lightly washed chestnut; tail olive, outer feathers edged with yellow on inner web. Breast and belly yellow, washed with chestnut which grades into black of throat; undertail-coverts yellow. Remiges dark brownish olive narrowly edged yellow on outer web and broadly on proximal portion of inner web; lesser coverts dark grey edged with olive-green; median coverts dark grey broadly tipped yellow, forming wing bar; greater and primary coverts dark brownish olive edged pale yellow; underwing-coverts bright yellow. Bill black; eyes orange-red to bright red; legs and feet light brown. ADULT Y (non-breeding): like non-breeding X but larger. When testes regress bill loses much of black colour which depends on testosterone (Collias et al. 1961). Y may have no off-season dress in equatorial regions. ADULT X (breeding): crown

Ploceus cucullatus

olive-green; face yellowish olive with indistinct dull yellow supercilium; sides of neck, back and rump greyish olive, back indistinctly streaked with brownish olive; tail feathers greyish olive with dull yellow margin to inner web on all except central pair. Underparts yellow, paler on belly, faintly washed with buff on breast, undertail-coverts pale buffy yellow. Remiges brownish olive, narrowly edged with dull yellow on outer web, more broadly on inner web; upperwing-coverts brownish olive, median and greater coverts narrowly tipped buffy yellow; underwing-coverts largely yellow. Bill black to brown, fading to ivory on basal lower part of lower mandible; eyes brown in 2nd-year females (which can breed), red by second breeding season; legs and feet light brown. ADULT X (non-breeding): like breeding X above, but underparts much less yellow, sometimes largely dull greyish white; bill paler. SIZE (10 YY, 10 XX): wing, Y 85–94 (887), X 73– 84 (774); tail, Y 51–57 (534), X 41–49 (462); bill, Y 20–21 (208), X 17–19 (183); tarsus, Y 22–25 (238), X 20–23 (217). WEIGHT (SW Nigeria, all months: Adegoke 1983b): Y (n ¼ 621) 34–46 (413), X (n ¼ 708) 32–37 (338). IMMATURE: resembles non-breeding X, eyes brown in X, brown to reddish-orange in Y. NESTLING: naked except for wisps of down; skin and palate pinkish; at this stage 25–8 g.

164

PLOCEIDAE

BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius (Ghana: Macdonald 1980). Colony in Uganda destroyed by party of Black-and-white Casqued Hornbills Bycanistes subcylindricus (the site not re-occupied: Hall 1970b). Nests robbed by Pied Crow Corvus alba, Senegal Coucal Centropus senegalensis and African

Plate 9 (Opp. p. 138)

Harrier Hawk Polyboroides typus (Macdonald 1980, Verheyen 1955); also by boomslang Dispholidus typus (Elgood and Ward 1963) and egg-eating snake Dasypeltis (Pitman 1958). Key References Crook, J. H. (1963), Din, N. A. (1992a, b, c, d), Hall, J. R. (1970a, b, c), Verheyen, R. (1955).

Ploceus cucullatus (Mu ¨ ller). Village Weaver; Black-headed Weaver; Spotted-backed Weaver. Tisserin gendarme. Oriolus cucullatus P. L. S. Mu ¨ ller, 1776. Natursyst., Suppl., p. 87; Senegal. Range and Status. Endemic resident and local intraAfrican migrant. Introduced in Mauritius, Re´union and West Indies (Hispaniola, Martinique). Widespread in subsaharan Africa except arid SW and NE, especially along watercourses, lake shores and coast lines. Generally abundant, but local and seasonal in dry areas on periphery of range. Centres of abundance are clearings in tropical rain forests in central and W Africa, and moist open country. Northern limits: Mauritania (Tagant), 18
N; Mali, delta region around Tombouctou, c. 17
N; Niger, (Tahoua, Tanout), 15
N; Chad, (Ouadi Haddat), 14
320 N; Sudan (where uncommon in E), 14
N; Eritrea, c. 16
N. Absent from desert regions of E Ethiopia and from Somalia north of 5
N; abundant along Jubba and Shabeelle river systems in S Somalia, and reappears in extreme S near Kenya border. Extreme N Kenya at Moyale and Daua R. on Ethiopian border, otherwise absent from arid regions north of Mt Elgon and Kapenguria and east of Tana R. drainage (except on coast). Southern limits: Angola (except central plateau) south to Mossamedes and Huı´la; Namibia, mainly in Caprivi, extending west along Okavango R.; in Botswana restricted to Okavango region and Limpopo drainage; Transvaal except SW, absent from western Highveld and Bushveld; Free State, mainly in moist E, scattered records elsewhere; rare in lowlands of Lesotho (ButheButhe, Leribe, Maseru, Mohale’s Hoek); E Cape Prov. west to Port Elizabeth and lower Gamtoos R. Occurs on Bioko, Sa˜o Tome´ (probably recently introduced: Christy and Clarke 1998); Pemba, Zanzibar, Mafia. Density, Acacia sevannas, Swaziland, 3, 20 and 51 birds per 100 ha. Description. P. c. cucullatus (Mu ¨ ller): Mauritania to Chad, south to Bioko, N Gabon and NW Zaı¨re. ADULT Y (breeding): head black; nape and hind neck chestnut; mantle yellow with 2 lateral black bands converging posteriorly on back to form distinct ‘V’; rump yellow, lightly washed chestnut; tail olive, outer feathers edged with yellow on inner web. Breast and belly yellow, washed with chestnut which grades into black of throat; undertail-coverts yellow. Remiges dark brownish olive narrowly edged yellow on outer web and broadly on proximal portion of inner web; lesser coverts dark grey edged with olive-green; median coverts dark grey broadly tipped yellow, forming wing bar; greater and primary coverts dark brownish olive edged pale yellow; underwing-coverts bright yellow. Bill black; eyes orange-red to bright red; legs and feet light brown. ADULT Y (non-breeding): like non-breeding X but larger. When testes regress bill loses much of black colour which depends on testosterone (Collias et al. 1961). Y may have no off-season dress in equatorial regions. ADULT X (breeding): crown

Ploceus cucullatus

olive-green; face yellowish olive with indistinct dull yellow supercilium; sides of neck, back and rump greyish olive, back indistinctly streaked with brownish olive; tail feathers greyish olive with dull yellow margin to inner web on all except central pair. Underparts yellow, paler on belly, faintly washed with buff on breast, undertail-coverts pale buffy yellow. Remiges brownish olive, narrowly edged with dull yellow on outer web, more broadly on inner web; upperwing-coverts brownish olive, median and greater coverts narrowly tipped buffy yellow; underwing-coverts largely yellow. Bill black to brown, fading to ivory on basal lower part of lower mandible; eyes brown in 2nd-year females (which can breed), red by second breeding season; legs and feet light brown. ADULT X (non-breeding): like breeding X above, but underparts much less yellow, sometimes largely dull greyish white; bill paler. SIZE (10 YY, 10 XX): wing, Y 85–94 (887), X 73– 84 (774); tail, Y 51–57 (534), X 41–49 (462); bill, Y 20–21 (208), X 17–19 (183); tarsus, Y 22–25 (238), X 20–23 (217). WEIGHT (SW Nigeria, all months: Adegoke 1983b): Y (n ¼ 621) 34–46 (413), X (n ¼ 708) 32–37 (338). IMMATURE: resembles non-breeding X, eyes brown in X, brown to reddish-orange in Y. NESTLING: naked except for wisps of down; skin and palate pinkish; at this stage 25–8 g.

Ploceus cucullatus YY of P. c. cucullatus and the following 3 races have black ‘V’ on yellow back, and wings significantly longer (891±036) than in last 4 races, which are mottle-backed (wings 843±042). P. c. bohndorffi Reichenow: N Gabon, forests of N Zaı¨re and Central African Republic; Uganda; W Kenya (L. Victoria basin). Intergrades with abyssinicus in NE Zaı¨re, Uganda and adjacent areas in SW Sudan. Y normally has black of crown less extensive than in cucullatus, chestnut on nape reaches hind crown, breast with more chestnut brown. SIZE (13 YY, 10 XX): wing, Y 82–91 (880), X 80–88 (832). WEIGHT: (Uganda, April–May) Y (n ¼ 10) 42–46 (436), X (n ¼ 9) 31–41 (356). P. c. frobenii Reichenow: Zaı¨re from c. 4
S to c. 8
S, west to near Lubue R. and east to c. 25
E. Y browner on breast and flanks than bohndorffi, but much individual variation and differences only trivial. Common in Kasai district. SIZE (13 YY, 9 XX): wing, Y 86–90 (895), X 77–82 (791). P. c. abyssinicus (Gmelin): Ethiopia and SE Sudan. Yellowest of the 4 ‘V-back’ races; black of crown more restricted than in bohndorffi, nape, hindneck and underparts yellow. SIZE (10 YY, 4 XX): wing, Y 84–94 (903), X 81–84 (823). P. c. collaris Vieillot: Gabon (lower Ogooue´ R.) south to Angola (Lobito Bay), east to c. 18
E in SW Zaı¨re. Whole head black, black throat bordered by band of deep chestnut on breast. Y with back feathers black tipped with yellow producing a scaly, spotted or mottled pattern. Interbreeds with cucullatus, bohndorffi and frobenii where ranges meet. SIZE (11 YY, 10 XX): wing, Y 78–89 (840), X 71–78 (751). P. c. nigriceps Layard (includes ‘paroptus’): Sa˜o Tome´, S Angola, S and SE Zaı¨re to E Kenya (below 2500 m), NE Kenya locally and S Somalia, south to N Botswana, S Zimbabwe, and Mozambique; Mafia I. Like collaris but lighter and yellower, Y with less black on head and little or no chestnut bordering foreneck. SIZE (10 YY, 12 XX): wing, Y 81–86 (835), X 75–84 (774). WEIGHT: (Kenya) Y ˆ i, (n ¼ 8, breeding) 34–42 (379), X (n ¼ 10) 25–38 (292); (Malaw lower Shire Valley, Nchalo) Y (n ¼ 229, in whole or partial breeding dress) 33–46 (377), X (n ¼ 134) 26–37 (314). P. c. graueri Hartert: extreme E Zaı¨re from S end of L. Kivu (below 2000 m) and NW shores of L. Tanganyika to Rwanda, Burundi, and NW Tanzania south to Kasulu (4
340 S, 30
060 E). Y very like Y nigriceps but has richer yellow underparts, more light wash of brown on breast, extending over flanks and belly. Intergrades with bohndorffi and abyssinicus northeast of L. Kivu near Zaı¨re border in Kigezi, SW Uganda. SIZE (13 YY, 10 XX): wing, Y 78–86 (837), X 75–80 (773). WEIGHT: (E Zaı¨re) Y (n ¼ 10) 36–43 (382), X (n ¼ 4) 30–33 (312). P. c. spilonotus Vigors (includes ‘dilutescens’): S Mozambique (south of c. 22
S), SE Botswana (Gaborone); South Africa and Lesotho. Introduced in Mauritius and Re´union. Y with yellow crown; intergrades with nigriceps widely in S Mozambique (Save

Valley and from Vilanculos north to Beira) and more widely in Zimbabwe and NE Transvaal (A: 4 hybrid types – (1) and (4) ‘dilutescens’  nigriceps, (2) and (3) ‘dilutescens’  ‘paroptus’: Clancey 1988). SIZE (9 YY, 8 XX): wing, Y 84–91 (867), X 78–82 (798). WEIGHT: (E Transvaal, Sept, Nov), Y (n ¼ 33) 33–47 (405), X (n ¼ 31) 29–40 (342). TAXONOMIC NOTE: Sibley and Monroe (1990) recognize only 4 races: P. c. cucullatus (V-backed group), and 3 mottle-backed groups, P. c. collaris with chestnut breast band, P. c. nigriceps with black head, and P. c. spilonotus with yellow crown. Underparts of breeding YY are more chestnut in humid climates, yellower in dry climates.

Field Characters. Length 135–17 cm. A common and familiar bird, larger than most weavers, with red eye (both sexes) and heavy black bill. Plumage varies considerably throughout its wide range. Breeding Y has head mainly black (except crown yellow in spilonotus); nape, hind neck and breast below black throat chestnut in W Africa, yellow elsewhere. Back either yellow with black ‘V’ formed by scapulars (W Africa to Ethiopia) or spotted. Y Speke’s Weaver P. spekei has back streaked rather than spotted, pale yellowish eye, and very different song. In South Africa, spilonotus told from Vitelline Masked and Lesser Masked Weavers P. velatus and P. intermedius by yellow forehead, and from Lesser Masked by red, not white, eye. Breeding X yellow below, brightest on throat, whiter on belly (varies with race: yellow more extensive in W, breast and belly white in S); most races have a yellow superciliary stripe. Non-breeding birds: Y like X but larger; both sexes duller, sometimes dull whitish below with little yellow. Similar-sized X Taveta Golden Weaver P. castaneiceps has smaller bill, heavily striped back and brown eye; X Speke’s Weaver lacks yellow supercilium and has pale eye; X Vieillot’s Black Weaver P. nigerrimus is dark olive above with heavily streaked back, greyish olive below. Voice. Tape-recorded (2, 4, 9, 22, 85, 86, 88, 99, 104, B, C, F, COLL, LEM, NOR, PAY); sonagrams of P. c. cucullatus in Collias (1963, 2000). About 26 different vocalizations. Commonest are buzzy or wheezing notes of Y song often preceded by brief, variable notes. Song directed at X often ends with a twitter. In territorial boundary disputes between YY, brief threat notes often precede buzzy notes. Alarm cries harsh, reaching high frequencies; threat calls also harsh but emphasizing low frequencies; calls of attraction are soft, brief, repetitive notes, or longer calls with clear harmonics. Mild alarm, sharp ‘chirp’; high intensity alarm in breeding season, short but loud, harsh chatter with very wide band of frequencies; this inhibits noisy cheeping of hungry nestlings. Distress cries of captured bird are loud, strident squawks. Low-intensity defensive threat a low-pitched, deep ‘chuck’; high-intensity threat when 1 Y chases another a very harsh snarl or growl. Flock contact call a soft, low ‘tsuk’; in flight a sharp higher ‘chick’. Y carrying nest materials gives rapid series of brief, almost humming, soft ‘tuk-tuktuk’ notes. Individually distinctive and repetitive nestattraction call of Y to unmated XX is 2-note, somewhat chirping ‘look-see’. When unmated X enters his territory, Y may at first chase her out with sharp ‘cut-cut’ notes; when unmated X first enters his nest Y gives either high intensity ‘wake’ or ‘wake-up’ of sexual excitement (a clear harmonic

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call with marked upswing in pitch), or a long soft purr. While visiting X inspects his nest, guarding Y utters a scold-chatter (usually followed by attempts to copulate if X perches in his territory). Soft whirring notes accompany copulation. Incubating X gives loud protest-squeal if Y attempts to enter nest. Y may give special complex call when X leaves nest and chuckling chatter when she returns. Nestling begs for food with loud high-pitched chirps which drop in pitch at the end; with age, chirps change to strident squawks. Just before feeding fledgling that has left nest, either parent utters very soft attraction call, a gentle harmonic ‘choo’ with marked drop in pitch. Juvenile song: see Development and Care of Young, below. General Habits. Occupies almost any open habitat: clearings in forest, moist savanna, coastal areas, river valleys, cultivation and suburban gardens; absent from arid zones, dense tropical rain forest and miombo woodland. For colony sites prefers large isolated trees. In moist country almost every village has a nesting colony, but in dry savanna is local, restricted to immediate vicinity of water; not necessarily near villages. Near equator breeds mainly below ˆ i up to c. 1000 m, in Natal up to c. 1500 m. 2000 m, in Malaw Highly gregarious. Forages in small, compact flocks well away from nest sites; hops on ground, flock moving along in one direction, with rear birds flying up toward front. Also forages among tree leaves and blossoms. Breeding Y forages infrequently, spending most of the time either in his small territory in colony tree or gathering nest materials. When not breeding, birds form huge, nomadic flocks that congregate near good feeding areas; they roost at night in groves of leafy trees, in reed-mace Typha latifolia, or reed beds or in tall grasses, e.g. elephant grass Pennisetum purpureum. Where food is less abundant, some flocks change roosts each day (Adegoke 1983a). Requires water for daily drinking and bathing; drinks at almost any time; does not dust bathe. When stressed by heat becomes hyperthermic, reduces general activity, and cools down by evaporative water loss from skin and by panting. Does not allopreen. Flight rather direct at c. 10 m per s, wings flapping almost continuously. Metabolic rate in flight about 8 times standard metabolic rate of 05 kcal per h (Da Camara-Smeets 1981a). Largely sedentary but in very dry areas arrives seasonally when there is adequate rainfall. Food. Insects, especially caterpillars, grasshoppers, crickets, small beetles, and winged termites taken in flight; grass seeds; fruits, e.g. Ehretia rigida and epicarp of oil-palm nut Elaeis guineensis, and in dry season stamens and green ovaries of flowers, and nectar of Aloe, Schotea brachypetala, Ziziphus mucronata. In breeding season prefers insects, but X also eats many green seeds or other green parts of plants. Captive birds ate av. 6 g of sorghum seeds per day; each bird needed 45 g to maintain basal metabolic rate. Aviary Y in breeding condition averaged 100 mealworms per day (196 kcal) for 3 days and ate almost no grain; standard metabolic rate approximately 12 kcal per day. Small stones and bits of snail shell eaten for grit. Nestlings fed almost entirely on insects.

Eats cultivated cereals (sorghum, rice, maize, wheat, millet) when available. Crop depredation a problem over much of its range, although preference for large wild grains, e.g. Rottboellia exaltata and Sorghum arundicearum, may reduce damage. In SW Nigeria, over half of diet (dry weight) was from cultivated plants. In Somalia, 63% of stomachs of 1484 birds contained cultivated grain, 36% wild seeds, and 79% insects. Near Ndjamena, Chad, in breeding season, 99 crops contained seeds (75% cultivated and 25% wild), also a few insects. One ad. ate 250 g of cultivated seed in 30 days, but 50 g of insects were taken during this period to feed one chick (Da Camara-Smeets 1981a). Captive birds preferred large wild seeds to cultivated grain and to small wild seeds (Da CamaraSmeets and Manikowski 1981). In central Uganda, 63% of 201 crops or stomachs had Rottboellia seeds and 76% had maize grains, but damage to maize heads was generally slight and wild grass seeds were the preferred food. In direct field observations over 1 year, insects averaged 21% of total food items and plant material 79%, but during the 2 rainy seasons birds took more insects than plant items (Kasoma 1987). Not a pest in southern Africa. Breeding Habits. Polygamous; colonial nester. Colony size varies from a few nests to hundreds in 1 tree. Uses a wide variety of trees, including eucalypts. In arid country, colonies are generally in acacia trees, in moist country often in palms, in rain forest clearings in almost any large tree, especially if it stands well alone, e.g. silk-cotton Ceiba (Eriodendron) tree or oil palm Elaeis. Nest placed at or near end of drooping branch over water when colony is on

Ploceus cucullatus

shore of stream or lake. In southern Africa, large colonies also nest in reeds Phragmites communis. Breeding coincides with rainy season, but YY may begin building nests well before rains begin and weeks before XX arrive. Each Y defends small territory in tree; his nests are up to c. 1 m apart, or may touch one another. Y sleeps in one of his nests. When defending territory, Y erects mantle feathers, sings and pecks at neighbouring YY. Y in breeding condition frequently vibrates slightly drooped wings, exposing bright yellow rump. Aggressive displays emphasize the black and courtship the yellow parts of plumage. When unmated XX visit colony, YY engage in spectacular communal displays. Each Y hangs upside down beneath entrance of his nest (B), chirping nest-invitation calls and flapping his wings to reveal bright yellow linings. When X chooses and enters a nest, Y looks up into entrance and sings to her. X chooses mate based on number of his nest invitation displays, and freshness and strength of nest; she favours previous mate and territory, mate more important than territory. X inspects inside of nest with her bill for up to 20 min, then either rejects it and departs, or accepts it, lines it with soft grass tops, and copulates with Y. She perches in territory with upraised bill, spreads and quivers wings slightly, and leans forward as Y mounts with wings upraised; she lifts her tail to one side as he brings his tail down, and vents meet. In next few days Y builds other nests and attempts to attract additional mates. If more than 1 X comes to Y’s nest at the same time, he may choose one and drive the others away. One Y can have up to 5 mates, but most have only 1, or none. When a nest is repeatedly rejected and fades from green to dull brown, Y tears it down and builds fresh one in its place. Brown colour stimulates nest destruction (Jacobs et al. 1978). Nest completed in 9–15 (av. 11) h (12 nests built by 6 YY, S Nigeria: Din 1992a). Persistently unsuccessful Y may shift territory or even quit the colony tree. If breeding season is long, X may have more than 1 brood; she sometimes changes mates between broods. Most of metabolic work of Y is for nest-building, of X for feeding nestlings. Doublebrooded; X may lay second clutch even before she stops feeding 1st brood. After independence, young Y sometimes builds onto nest being constructed by mature Y (and may be tolerated, probably because of its ‘X’ plumage). X often first breeds in its 2nd year (when eyes still brown); Y does not normally breed in first season after it hatched. NEST: ovoid structure 140–170 long, 110–130 high, with domed roof; bottom entrance (diam. c. 50) separated by

inner ridge from egg chamber. Y weaves outer shell in 5 stages, starting with ring (C). Using long thin strips torn from leaves of tall green grasses, e.g. elephant grass Pennisetum purpureum, or palm fronds, he weaves intricate patterns (D), pushing and pulling each strip through the ring (E) or nest wall. Perching on threshold, he pushes out with bill in all directions as far as he can reach, giving form and size to nest. Just under roof he thatches rain-proof ceiling of short, rectangular pieces of grass or of dicotyledonous leaves (hatched in F, a vertical section through nest). X lines nest with strips of grass or palm leaf, feathery grass tops, and sometimes feathers (stippled in F); Y usually adds 40–80 mm entrance tube (F) while X is incubating, and strengthens external nest attachments. After brood leaves nest, Y tears it down and often builds another in its place. In Senegal, Y cucullatus does not help feed nestlings but builds twice as many nests and averages more mates than in E Zaı¨re, where Y graueri helps X feed young (for biology of nest-building behaviour see Collias and Collias 1984). Often nests in same colony tree as other weaver species (Lahti et al. 2002; for studies of courtship and nesting behaviour in colonies with P. nigerrimus see that species, Crook 1963 and Din 1992a). Known to hybridize with P. nigerrimus.

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EGGS: 1–5, laid in morning on successive days. Just before laying, X perches on threshold (inner ridge of egg chamber) facing entrance, visible abdominal contractions begin 6–16 s before egg emerges, final contraction strongest, tail elevated, rear end body depressed, and egg is deposited into chamber (Victoria 1969). In aviary birds from Senegal, XX in first breeding season lay 1–3 eggs, av. 17 (38 clutches), and ad. XX 1–4, av. 23 (265 clutches). 139 clutches, South Africa, 2–5, av. 26. Ovoid; glossy, colour highly variable: white, turquoise, emeraldturquoise, emerald, occasionally greenish emerald; unspotted or finely or coarsely spotted with dark brown, reddish brown or lilac-grey. Colour, spotting, size and shape of eggs are characteristic and consistent for each X throughout her lifetime. At Ile-Ife, Nigeria, in 9 colonies over 2 years, 5600 were of 7 colour types: light blue with small specks (35%), plain light blue (29%), white with small specks (13%), plain white (11%), light blue with large specks (7%), white with large specks (3%) and dense

blue-blotched (2%) (Din 1992b). SIZE: (n ¼ 986 from 34 XX from Senegal in aviary: Collias 1984) 19–26  13–17 (226  155); also Nigeria (n ¼ 29), 19–26  14–19 (229  152); central Africa (bohndorffi), 217–257  153– 164; southern Africa (n ¼ 180), 20–25  13–17 (231  149). WEIGHT: (spilonotus, n ¼ 3), 15 g. LAYING DATES: Mauritania, July–Oct; Senegal, May–Nov (Aug–Sept in N); Gambia, Aug–Oct; Mali, June–Sept; Guinea-Bissau, Apr–Sept; Guinea, July–Oct; Sierra Leone, July–Oct; Liberia (nest-building all months, main season Aug–Apr; Ghana, Mar–Apr, July–Aug, Oct–Nov (nestbuilding all year); Burkina Faso, Aug–Sept; Togo, Jan– Oct; Nigeria (Ile-Ife), Apr–Aug; Niger (active colonies Apr–Sept); SW Chad (Ndjamena), Aug–Oct; Sudan Mar, Aug–Dec; Ethiopia, Jan, Mar–Oct; Somalia, May (juvs Jan); S Cameroon and Gabon, all year; Bioko, Apr–Sept; Central African Republic (breeding condition June). Zaı¨re, equatorial areas, all year; SW, Dec–May; S-central, Oct–May; SE, Sept–Feb; E, Nov–June; NE, Aug–Nov. E Africa: Uganda (equatorial areas), all year; Kenya, mostly Feb–June; Tanzania coast, Mar–Apr, Sept–Dec; Region A, Apr, Aug; Region B, Feb–Dec, peaking Feb–June and Oct; Region C, Feb–Mar, May, Oct, Dec; Region D, Feb–Nov, peaking Mar–May and Oct–Nov; Region E, July; Angola, ˆ i, Oct–Nov, Jan–Mar; Zambia, Jan–Mar, Sept–Dec; Malaw Feb–Apr, Nov–Apr (Nchalo); Zimbabwe, Aug–Apr; S Mozambique, Sept–Feb; Botswana, Mar–Apr (Shakwe), Jan (Kasane); Okavango Dec–Apr; South Africa, Aug–Feb (Transvaal, Aug–Feb; KwaZulu-Natal, Sept–Jan, E Cape, Oct–Jan). In southern Africa, breeding activity reported in all months (Harrison et al. 1997). Normally, breeds in rainy season when green nest materials and insect food most abundant, but may respond to increase in day-length independently of rainfall. In outdoor aviaries in California, a region of winter rains, over a 10-year period, Village Weavers given fresh nest materials and insect food did not breed in winter but bred during long days of dry season in spring and summer. YY not allowed to breed may remain in breeding plumage (Collias and Collias 1970b). In Nigeria, 4 main peaks of hatching during rains (Mar–Aug) and 1–2 in dry season Nov–Dec.

Ploceus cucullatus INCUBATION: by X only; begins (and copulation ends) with first egg. 4 XX (Senegal) remained in nest for bouts of 1–38 min (mean 61), and out of nest for 1–17 min (mean 40), during 10 h of observation (Collias and Collias 1969); timing of different XX varies greatly. Less attentive in warm weather, more attentive when strong wind tosses nest about. Good nest insulation allows X more time to forage. In aviary, 24-h mean egg temperature was 359
C (353
C by day and 365
C at night: White and Kinney 1974). Periodically, X stands and pokes bill down to turn eggs or rearrange nest material, and before settling down faces another direction. When clutch lost, she may lay again within 6 days. Period: (n ¼ 11 eggs, in aviary) 12–13 (122) days. DEVELOPMENT AND CARE OF YOUNG: young hatch sequentially on different days; X removes egg shells from nest. At weight of 9–17 g, feather sheaths appear in main tracts, eyes still closed; at 21–31 g, some body and wing feathers breaking out of sheaths, eyes start to open; at 26–35 g, head partly feathered, primaries with more than half their length out of sheath; at 29–37 g, body covered with feathers, tail about half its final length, last vestiges of down on head (P. c. spilonotus, Collias and Collias 1971a). Growth in body weight slows down greatly when chicks are about to leave nest, at c. 17–21 days. Fledged young resemble ad. X but are more greyish-olive above, eyes dark grey-brown, yellowish white mark along base of gape, tail short. Young may crawl in and out of nest before leaving. Nestlings fed by X or both parents (mainly X) on insects, with seeds added late in nestling period. In Senegal, Y fed nestlings only once in c. 10 h if at all, X 41 times per h; in E Zaı¨re, Y fed 23 times per h, X 47 per h; in South Africa, Y fed 14 per h, X 42 per h (Collias and Collias 1967, 1971a– c). Y cucullatus does not help feed nestlings at L. Chad or in NE Gabon, but Y abyssinicus does help in Ethiopia. In a large colony many different broods may leave nests at about the same time; they are then fed by X, sometimes also by Y, who remains in territory while X wanders off with fledged young. XX in aviary fed young out of nest for 2–3 weeks before young learned to feed themselves. Small nestling ejects faeces in rubbery sac which parent removes; older nestlings defaecate toward nest entrance. Unlike young X, young Y sings, from 1 month after hatching, and builds crude practice nests of short, less appropriate materials. It needs practice but is not taught by other YY. Young YY form small ‘play colonies’ in low trees and bushes, but XX do not visit these colonies. Juvenile Y song is soft, continuous, formless warbling which gradually becomes segmented into discrete notes. The characteristic wheeze appears later. Young YY imitate complex introductory notes of mature song given by other young YY, particularly from dominant ones. By contrast, learning is apparently not important in development of Y’s individually distinctive nest invitation calls (Jacobs 1979). BREEDING SUCCESS/SURVIVAL: in Guinea, in 130 nests, av. 186 eggs per nest, producing av. 128 nestlings per nest. In Chad, contents of 20 nests counted repeatedly had 214 eggs and 19 nestlings. No direct evidence for proportion of fledglings that survive to become breeders. In E Zaı¨re (Kivu), 9 or 10 nests were woven for every one from which a brood fledged: of 342 nests woven during 9-month

breeding season, broods fledged from only 32; XX rejected 208 nests, eggs were lost in 28, and nestlings hatched but did not fledge in 23; 51 other nests failed during period of violent storms (Collias and Collias 1967). YY vary greatly in breeding success: the more XX present, the more nests YY build. In NW Senegal, nesting colonies with fewer than 10 YY attracted significantly fewer XX and failed to increase. Larger colonies usually had about 7 XX for every 3 YY. Colonies may be deserted during prolonged spells of dry weather. In Nigeria (Ile-Ife), breeding commenced early in large colonies (>500 nests) in broadleafed trees; in small colonies (14 years later. Captive Y lived 24 years; Y >14 years old builds fewer nests and has fewer mates; X lays fewer eggs when 9–18 years old. Nestlings taken by rock python Python sebae, boomslang Dispholidus typus and mamba Dendroaspis sp. (snakes a major predator: G); Chacma Baboons Papio ursinus take eggs and nestlings. African Harrier Hawk Polyboroides typus hangs from nest and takes nestlings out of entrance; Gabar Goshawk Micronisus gabar tears hole in nest roof to extract nestlings; once Bat Hawk Machaerhamphus alcinus tore down nest, and African Goshawk Accipiter tachiro caught ad. Y. People often take nestlings for food, and destroy large numbers of young birds and ads to protect crops. Parasitized by Diederik Cuckoo Chrysococcyx caprius, which may remove egg of host before laying. X Village Weaver often removes any egg that does not match her own. (For evolution of weaver reaction to parasitic cowbird Molothrus eggs in Hispaniola see, Cruz and Wiley 1989 and Robert and Sorci 1999).

169

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PLOCEIDAE

Ectoparasites include bird lice Myrsidea textoris, Mallophaga and gamasid mites. Blood parasites include nematodes Filaroidea and protozoans Haemoproteus, Leucocytozoon and Plasmodium.

Key References Chapin, J. P. (1954), Collias, E. C. (1993), Collias, N. E. (2000), Collias, N. E. and Collias, E. C. (1959, 1970a, 1971a, 1984).

Plate 9

Ploceus grandis (Gray). Giant Weaver. Tisserin ge ´ant.

(Opp. p. 138)

Hyphantornis grandis Gray, 1844. Genera of Birds, ii, p. [1] of Ploceinae, nom. nov. pro Ploceus collaris Fraser (nec Vieillot), Proc. Zool. Soc., 1842, p. 42; Sa˜o Tome´. Range and Status. Endemic resident, restricted to Sa˜o Tome´. Common. Description. ADULT Y: forehead, crown, lores and sides of head black, passing into chestnut-brown of nape with no sharp division; mantle greenish yellow with thin dark central streaks; rump dull yellow; tail dark olive-green; chin and throat black, passing into chestnut-brown band across breast; flanks yellow washed with chestnut; belly, thighs and undertail-coverts yellow; wings dark olive-green, tertials and coverts with yellow edges, yellow tips of upperwing-coverts forming a wing-bar. Bill black; eyes yellow; legs brown. No non-breeding plumage. ADULT X: forehead, crown and nape olive-grey with dark central lines; heavier streaking on olive-grey mantle; rump plain grey; tail dark olive-green; wings dark brown with narrow yellow edges to remiges, broader pale tips to coverts, underwing cream; chin and throat cream; breast with broad buffy band; flanks buffy grey; belly, and undertail-coverts white; thighs grey. Upper mandible dark brown, lower mandible horn; eyes brown; legs brown. SIZE (7 YY, 6 XX): wing, Y 108–119 (112), X 101–111 (106); tail, Y 68–78 (721), X 61–73 (673); bill, Y 305–332 (315), X 288–313 (301); tarsus, Y 280–301 (289), X 266–303 (280). WEIGHT: unsexed (n ¼ 4) 62–65. IMMATURE: like X but upperparts brown rather than grey; broad buffy edges to wing-coverts. Underparts also generally buffy rather than grey, buffy on sides of head. NESTLING: not described. TAXONOMIC NOTE: often regarded as most closely related to Village Weaver P. cucullatus (e.g. Hall and Moreau 1970, Christy and Clarke 1998), but on the basis of plumage characters and body proportions, de Naurois and Wolters (1975) suggested that it is closest to Yellow-backed Weaver P. melanocephalus.

Ploceus grandis

Voice. Tape-recorded (B, JO PJ, TYE). Chorus of screeches and swizzles at colony. Song more raucous and less complex than that of Village Weaver, ‘pche pche pche pche pche pche gzzzuinuiuinuinuinuin’; birds pursuing each other in flight give repeated ‘prrric prrrric prrrric’ (Christy and Clarke 1998).

dotted with small patches of woodland, large isolated clumps of tall trees, forest on slopes which are too steep to log; rare in savanna, commonest in degraded habitats, especially in thick clumps of Alchornea cordifolia which is abundant along roadsides and in plantations, and in grass clumps in plantations. Originally possibly occupied forests on the coast and on the plains of the N and E, which would have been the first to be felled for plantations (Christy and Clarke 1998). Occurs singly, in pairs or small family groups. Does not form large noisy flocks like Village Weaver, nor does it join mixed-species flocks of insectivores, but typically seen alone. Forages at all levels in vegetation. In primary forest undergrowth acts like an insectivore; X was observed exploring a Uapaca, climbing its roots, going through lower clumps of foliage, and clinging to moss-covered bark; in cocoa plantations around 500 m hunts insects by probing acrobatically into rotting holes, piercing the outer layer and extracting larvae.

General Habits. Inhabits natural forest, especially undergrowth, up to 1500 m, plantations (coffee, oil palm, coconut, bamboo, cocoa shaded by Erythrina trees), crops

Food. Fruit, including fruits of Alchornea cordifolia, pawpaws and especially oil-palm Elaeis; on ground feeds on pulp of coconuts crushed by tractors; grass seeds;

Field Characters. Length 22 cm. World’s largest weaver, with huge bill. Y looks like giant version of Village Weaver P. cucullatus, with black head surrounded by chestnut, but eye pale, back plain; local race of Village Weaver lacks chestnut and has red eye, streaked back. X plumage dull, with pale eye, no supercilium, breast buff, not yellow. In flight appears short and broad-winged (Christy and Clarke 1998).

170

PLOCEIDAE

Ectoparasites include bird lice Myrsidea textoris, Mallophaga and gamasid mites. Blood parasites include nematodes Filaroidea and protozoans Haemoproteus, Leucocytozoon and Plasmodium.

Key References Chapin, J. P. (1954), Collias, E. C. (1993), Collias, N. E. (2000), Collias, N. E. and Collias, E. C. (1959, 1970a, 1971a, 1984).

Plate 9

Ploceus grandis (Gray). Giant Weaver. Tisserin ge ´ant.

(Opp. p. 138)

Hyphantornis grandis Gray, 1844. Genera of Birds, ii, p. [1] of Ploceinae, nom. nov. pro Ploceus collaris Fraser (nec Vieillot), Proc. Zool. Soc., 1842, p. 42; Sa˜o Tome´. Range and Status. Endemic resident, restricted to Sa˜o Tome´. Common. Description. ADULT Y: forehead, crown, lores and sides of head black, passing into chestnut-brown of nape with no sharp division; mantle greenish yellow with thin dark central streaks; rump dull yellow; tail dark olive-green; chin and throat black, passing into chestnut-brown band across breast; flanks yellow washed with chestnut; belly, thighs and undertail-coverts yellow; wings dark olive-green, tertials and coverts with yellow edges, yellow tips of upperwing-coverts forming a wing-bar. Bill black; eyes yellow; legs brown. No non-breeding plumage. ADULT X: forehead, crown and nape olive-grey with dark central lines; heavier streaking on olive-grey mantle; rump plain grey; tail dark olive-green; wings dark brown with narrow yellow edges to remiges, broader pale tips to coverts, underwing cream; chin and throat cream; breast with broad buffy band; flanks buffy grey; belly, and undertail-coverts white; thighs grey. Upper mandible dark brown, lower mandible horn; eyes brown; legs brown. SIZE (7 YY, 6 XX): wing, Y 108–119 (112), X 101–111 (106); tail, Y 68–78 (721), X 61–73 (673); bill, Y 305–332 (315), X 288–313 (301); tarsus, Y 280–301 (289), X 266–303 (280). WEIGHT: unsexed (n ¼ 4) 62–65. IMMATURE: like X but upperparts brown rather than grey; broad buffy edges to wing-coverts. Underparts also generally buffy rather than grey, buffy on sides of head. NESTLING: not described. TAXONOMIC NOTE: often regarded as most closely related to Village Weaver P. cucullatus (e.g. Hall and Moreau 1970, Christy and Clarke 1998), but on the basis of plumage characters and body proportions, de Naurois and Wolters (1975) suggested that it is closest to Yellow-backed Weaver P. melanocephalus.

Ploceus grandis

Voice. Tape-recorded (B, JO PJ, TYE). Chorus of screeches and swizzles at colony. Song more raucous and less complex than that of Village Weaver, ‘pche pche pche pche pche pche gzzzuinuiuinuinuinuin’; birds pursuing each other in flight give repeated ‘prrric prrrric prrrric’ (Christy and Clarke 1998).

dotted with small patches of woodland, large isolated clumps of tall trees, forest on slopes which are too steep to log; rare in savanna, commonest in degraded habitats, especially in thick clumps of Alchornea cordifolia which is abundant along roadsides and in plantations, and in grass clumps in plantations. Originally possibly occupied forests on the coast and on the plains of the N and E, which would have been the first to be felled for plantations (Christy and Clarke 1998). Occurs singly, in pairs or small family groups. Does not form large noisy flocks like Village Weaver, nor does it join mixed-species flocks of insectivores, but typically seen alone. Forages at all levels in vegetation. In primary forest undergrowth acts like an insectivore; X was observed exploring a Uapaca, climbing its roots, going through lower clumps of foliage, and clinging to moss-covered bark; in cocoa plantations around 500 m hunts insects by probing acrobatically into rotting holes, piercing the outer layer and extracting larvae.

General Habits. Inhabits natural forest, especially undergrowth, up to 1500 m, plantations (coffee, oil palm, coconut, bamboo, cocoa shaded by Erythrina trees), crops

Food. Fruit, including fruits of Alchornea cordifolia, pawpaws and especially oil-palm Elaeis; on ground feeds on pulp of coconuts crushed by tractors; grass seeds;

Field Characters. Length 22 cm. World’s largest weaver, with huge bill. Y looks like giant version of Village Weaver P. cucullatus, with black head surrounded by chestnut, but eye pale, back plain; local race of Village Weaver lacks chestnut and has red eye, streaked back. X plumage dull, with pale eye, no supercilium, breast buff, not yellow. In flight appears short and broad-winged (Christy and Clarke 1998).

Ploceus weynsi

insects, including beetles and larvae, primarily gleaned from branches and creepers; also snails.

pursue each other frequently, even though, away from nest site, they are shy. In display, Y perches under palm leaf near nest, fluffs out plumage, beats wings spasmodically and rather feebly, giving typical weaver call (Christy and Clarke 1998). NEST: large ball with no entrance tunnel (A) in cocoa plantations built solely of strips of palm leaves; supported by branches and well concealed in foliage; placed in crown of palm tree, in heart of coconut palm, or in leafy branches of tall trees. EGGS: 1–2, plain blue. LAYING DATES: Jan (B. Fisher, pers. comm.); perhaps Aug–Mar (copulation Dec, juvs Mar–Apr, June–Aug, nestbuilding July, Aug, Dec–Jan). No other information.

Breeding Habits. Monogamous. Usually a solitary nester, but in favourable sites like plantations there may be one in each tree. YY in these loose colonies are aggressive and

Key References Christy, P. and Clarke, W. V. (1998), de Naurois, R. and Wolters, H. E. (1975).

171

Ploceus weynsi (Dubois). Weyns’s Weaver. Tisserin de Weyns.

Plate 10

Melanopteryx weynsi Dubois, 1900. Orn. Monatsb., 8, p. 69; Bumba, Upper Congo.

(Opp. p. 139)

Forms a superspecies with P. golandi. Range and Status. Endemic resident. Zaı¨re, Mbandaka to Bumba and Buta. Uganda, Budongo, Ankole, Kigezi, Masaka, Sese I., Sango Bay, Mpumu, Entebbe, Gaba, Kibanga Port, Kifu, Mabira, Busoga, Bumasifwa; common in Malabigambo Forest. Tanzania, Bukoba. Local and perhaps seasonal in occurrence. Description. ADULT Y: forehead, crown, nape and mantle black; rump olive-green with some black barring or flecking; tail olive-green; wings dark olive-green with narrow yellow edges on remiges, broad margins to coverts and tertials, with yellow tips to upperwing-coverts forming distinct wing-bar; underwing yellow; chin, throat, cheeks, ear-coverts and breast black, chestnut-brown feathers at border with belly; flanks chestnutbrown; belly chestnut-brown, with variable area of yellow in centre; thighs yellow flecked with olive-green; undertail-coverts yellow. Bill black; eyes pale yellow with chestnut inner ring; legs brown. No seasonal change in plumage. ADULT X: forehead, crown, nape and sides of head plain olive-green, with no supercilium; mantle olive-green with faint darker central streaks; rump olive-green, unstreaked but faint dark tips to feathers; tail olive-green; wings dark olive-green with narrow yellow edges to remiges, broader tips to upperwing-coverts forming wing bar; chin, throat and breast pale yellow in midline passing into white on belly; flanks dull olive-green; thighs olive-green on outer surface, inner side white tinged with yellow; undertail-coverts white. Bill dark brown, lower mandible paler than upper; eyes pale yellow; legs brown. SIZE (12 YY, 12 XX): wing, Y 76–84 (806), X 70–78 (740); tail, Y 45–51 (473), X 41–49 (439); bill, Y 191–216 (202), X 186–200 (193); tarsus, Y 182–215 (205), X 181–204 (191). WEIGHT: Y (Uganda, n ¼ 31) 239–358 (309), (Zaı¨re, n ¼ 5) 226–301 (264), X (Uganda, n ¼ 28) 228–335 (273), (Zaı¨re, n ¼ 5) 204–246 (221); juv. Y (Zaı¨re, n ¼ 4) 257– 350 (297), juv. X (Zaı¨re) 241; also, (Uganda) 3 YY 36–37, 4 XX 30–33 (Friedmann and Williams 1969). IMMATURE: like X; bill brown, eyes brown.

Ploceus weynsi

?

NESTLING:

?

unrecorded.

Field Characters. Length 15 cm. Black head, back and breast and pale eye of Y suggest Vieillot’s Black Weaver P. nigerrimus, but rest of underparts chestnut and yellow, and wings have yellow wing-bar and feather edgings. X has

Ploceus weynsi

insects, including beetles and larvae, primarily gleaned from branches and creepers; also snails.

pursue each other frequently, even though, away from nest site, they are shy. In display, Y perches under palm leaf near nest, fluffs out plumage, beats wings spasmodically and rather feebly, giving typical weaver call (Christy and Clarke 1998). NEST: large ball with no entrance tunnel (A) in cocoa plantations built solely of strips of palm leaves; supported by branches and well concealed in foliage; placed in crown of palm tree, in heart of coconut palm, or in leafy branches of tall trees. EGGS: 1–2, plain blue. LAYING DATES: Jan (B. Fisher, pers. comm.); perhaps Aug–Mar (copulation Dec, juvs Mar–Apr, June–Aug, nestbuilding July, Aug, Dec–Jan). No other information.

Breeding Habits. Monogamous. Usually a solitary nester, but in favourable sites like plantations there may be one in each tree. YY in these loose colonies are aggressive and

Key References Christy, P. and Clarke, W. V. (1998), de Naurois, R. and Wolters, H. E. (1975).

171

Ploceus weynsi (Dubois). Weyns’s Weaver. Tisserin de Weyns.

Plate 10

Melanopteryx weynsi Dubois, 1900. Orn. Monatsb., 8, p. 69; Bumba, Upper Congo.

(Opp. p. 139)

Forms a superspecies with P. golandi. Range and Status. Endemic resident. Zaı¨re, Mbandaka to Bumba and Buta. Uganda, Budongo, Ankole, Kigezi, Masaka, Sese I., Sango Bay, Mpumu, Entebbe, Gaba, Kibanga Port, Kifu, Mabira, Busoga, Bumasifwa; common in Malabigambo Forest. Tanzania, Bukoba. Local and perhaps seasonal in occurrence. Description. ADULT Y: forehead, crown, nape and mantle black; rump olive-green with some black barring or flecking; tail olive-green; wings dark olive-green with narrow yellow edges on remiges, broad margins to coverts and tertials, with yellow tips to upperwing-coverts forming distinct wing-bar; underwing yellow; chin, throat, cheeks, ear-coverts and breast black, chestnut-brown feathers at border with belly; flanks chestnutbrown; belly chestnut-brown, with variable area of yellow in centre; thighs yellow flecked with olive-green; undertail-coverts yellow. Bill black; eyes pale yellow with chestnut inner ring; legs brown. No seasonal change in plumage. ADULT X: forehead, crown, nape and sides of head plain olive-green, with no supercilium; mantle olive-green with faint darker central streaks; rump olive-green, unstreaked but faint dark tips to feathers; tail olive-green; wings dark olive-green with narrow yellow edges to remiges, broader tips to upperwing-coverts forming wing bar; chin, throat and breast pale yellow in midline passing into white on belly; flanks dull olive-green; thighs olive-green on outer surface, inner side white tinged with yellow; undertail-coverts white. Bill dark brown, lower mandible paler than upper; eyes pale yellow; legs brown. SIZE (12 YY, 12 XX): wing, Y 76–84 (806), X 70–78 (740); tail, Y 45–51 (473), X 41–49 (439); bill, Y 191–216 (202), X 186–200 (193); tarsus, Y 182–215 (205), X 181–204 (191). WEIGHT: Y (Uganda, n ¼ 31) 239–358 (309), (Zaı¨re, n ¼ 5) 226–301 (264), X (Uganda, n ¼ 28) 228–335 (273), (Zaı¨re, n ¼ 5) 204–246 (221); juv. Y (Zaı¨re, n ¼ 4) 257– 350 (297), juv. X (Zaı¨re) 241; also, (Uganda) 3 YY 36–37, 4 XX 30–33 (Friedmann and Williams 1969). IMMATURE: like X; bill brown, eyes brown.

Ploceus weynsi

?

NESTLING:

?

unrecorded.

Field Characters. Length 15 cm. Black head, back and breast and pale eye of Y suggest Vieillot’s Black Weaver P. nigerrimus, but rest of underparts chestnut and yellow, and wings have yellow wing-bar and feather edgings. X has

172

PLOCEIDAE

plain olive-green upperparts and face, with pale eye but no supercilium; separated from very similar X Vieillot’s by blackish bill, green tail and yellow underwings.

Food. Fruit, including wild figs. Stomachs of 3 YY contained fruit of Beilschmiedia discolor, Alchornea cordifolia (Demey et al. 2000).

Voice. Not tape-recorded. Contact call a high-pitched ‘chip’ reminiscent of Southern Red Bishop Euplectes orix. General Habits. Lives high up in tall forest trees in seasonally dry swamp forest, also in lush lake shore vegetation. In Kampala area, Uganda, apparently absent from lake shore June–Sept. Roosts and feeds in noisy flocks of 10–20, and up to 200 birds in Sept (Carswell 1986); about 100 birds roosted on forested islands in Congo R. with other weavers. Restless, with fast flight. Moult of ads in Zaı¨re Sept–Nov, juvs June–Oct. Resident, but seems to be absent from N shores of L. Victoria in June–early Sept (Carswell 1986); wanders erratically (Stevenson and Fanshawe 2002).

Breeding Habits. Presumed to be colonial as always encountered in flocks. Neither nest nor eggs have been described. LAYING DATES: Zaı ¨re, (gonads enlarged Apr–June); Uganda, (perhaps June–July: van Someren 1916; young birds, Nov).

Key References Carswell, M. (1986), Demey, R. et al. (2000), Friedmann, H. and Williams, J. G. (1969), Stevenson, T. and Fanshawe, J. (2002).

Plate 11

Ploceus golandi (Clarke). Clarke’s Weaver. Tisserin de Clarke.

(Opp. p. 154)

Heterhyphantes golandi Clarke, 1913. Bull. Br. Orn. Club, 31, p. 32; Mombasa, Kenya. Forms a superspecies with P. weynsi. Range and Status. Endemic resident, known only from Arabuko-Sokoke Forest Dakatcha area (Malindi), Sabaki R., and from Galana Ranch to Marafa Forest and Hadu, SE Kenya. Flocks of >200 including juvs, Marafa, July, and 200–400, Sokoke, July–Aug. Total range 535 km2, population estimated at 1000–2000 pairs in 1985; vulnerable to forest clearance.

Ploceus golandi

Description. ADULT Y: forehead, crown, nape and mantle black; rump bright olive-green with some feathers yellow-fringed; tail dark olive-green; chin, throat and breast black, ending in a sharp line at the belly; flanks white with yellow flecks, belly white overlaid with yellow in centre; thighs black; undertail-coverts pale whitish yellow; wings and wing-coverts blackish brown with yellow edges, broad on tertials and coverts, producing 2 prominent yellow wing-bars; underwing grey. Bill black; eyes dark brown; legs light brown. ADULT X: forehead, crown and nape bright olive-green; mantle olive-green with black central streaks; rump olive-green, unstreaked; tail dark olive-green; narrow yellow supercilium, lores, cheeks and ear-coverts dull yellow; chin, throat and breast bright yellow except for olivegreen patches on breast level with bend of wings; belly and flanks whitish with yellow streaks, thighs grey with yellow streaks, undertail-coverts white; wings blackish brown with yellow edges. Bill horn-grey; eyes brown; legs pale brown, feet pinkish. SIZE (4 YY, 3 XX): wing, Y 72–75 (738), X 69–72 (710); tail, Y 43–47 (448), X 43–46 (443); bill, Y 175–200 (190), X 172–186 (178); tarsus, Y 204–209 (206), X 185–191 (189). WEIGHT: Y (n ¼ 17) 225–270 (246); X (n ¼ 16) 208–246 (225). IMMATURE: dull greenish above, hardly streaked; underparts washed with dull yellow; bill brown. Sub-adult Y olive and yellowish, head with black patches and trace of superciliary stripe. NESTLING: not described.

extremely restricted range, by pale yellow and white belly, olive rump and tail, yellow wing feather edgings; XX cannot be confused because X Dark-backed is like Y, while X Clarke’s a typical X weaver with streaked upperparts, yellow underparts, yellow wing edgings.

Field Characters. Length 135 cm. Y has black head, breast and back, dark eye, slender bill; told from Darkbacked Weaver P. bicolor, the only other forest weaver in its

Voice. Tape-recorded (B, CART, KEI, McVIC). Foraging flocks keep up continual choruses of chattering, chirping and sizzling sounds which rise and fall in volume, audible

172

PLOCEIDAE

plain olive-green upperparts and face, with pale eye but no supercilium; separated from very similar X Vieillot’s by blackish bill, green tail and yellow underwings.

Food. Fruit, including wild figs. Stomachs of 3 YY contained fruit of Beilschmiedia discolor, Alchornea cordifolia (Demey et al. 2000).

Voice. Not tape-recorded. Contact call a high-pitched ‘chip’ reminiscent of Southern Red Bishop Euplectes orix. General Habits. Lives high up in tall forest trees in seasonally dry swamp forest, also in lush lake shore vegetation. In Kampala area, Uganda, apparently absent from lake shore June–Sept. Roosts and feeds in noisy flocks of 10–20, and up to 200 birds in Sept (Carswell 1986); about 100 birds roosted on forested islands in Congo R. with other weavers. Restless, with fast flight. Moult of ads in Zaı¨re Sept–Nov, juvs June–Oct. Resident, but seems to be absent from N shores of L. Victoria in June–early Sept (Carswell 1986); wanders erratically (Stevenson and Fanshawe 2002).

Breeding Habits. Presumed to be colonial as always encountered in flocks. Neither nest nor eggs have been described. LAYING DATES: Zaı ¨re, (gonads enlarged Apr–June); Uganda, (perhaps June–July: van Someren 1916; young birds, Nov).

Key References Carswell, M. (1986), Demey, R. et al. (2000), Friedmann, H. and Williams, J. G. (1969), Stevenson, T. and Fanshawe, J. (2002).

Plate 11

Ploceus golandi (Clarke). Clarke’s Weaver. Tisserin de Clarke.

(Opp. p. 154)

Heterhyphantes golandi Clarke, 1913. Bull. Br. Orn. Club, 31, p. 32; Mombasa, Kenya. Forms a superspecies with P. weynsi. Range and Status. Endemic resident, known only from Arabuko-Sokoke Forest Dakatcha area (Malindi), Sabaki R., and from Galana Ranch to Marafa Forest and Hadu, SE Kenya. Flocks of >200 including juvs, Marafa, July, and 200–400, Sokoke, July–Aug. Total range 535 km2, population estimated at 1000–2000 pairs in 1985; vulnerable to forest clearance.

Ploceus golandi

Description. ADULT Y: forehead, crown, nape and mantle black; rump bright olive-green with some feathers yellow-fringed; tail dark olive-green; chin, throat and breast black, ending in a sharp line at the belly; flanks white with yellow flecks, belly white overlaid with yellow in centre; thighs black; undertail-coverts pale whitish yellow; wings and wing-coverts blackish brown with yellow edges, broad on tertials and coverts, producing 2 prominent yellow wing-bars; underwing grey. Bill black; eyes dark brown; legs light brown. ADULT X: forehead, crown and nape bright olive-green; mantle olive-green with black central streaks; rump olive-green, unstreaked; tail dark olive-green; narrow yellow supercilium, lores, cheeks and ear-coverts dull yellow; chin, throat and breast bright yellow except for olivegreen patches on breast level with bend of wings; belly and flanks whitish with yellow streaks, thighs grey with yellow streaks, undertail-coverts white; wings blackish brown with yellow edges. Bill horn-grey; eyes brown; legs pale brown, feet pinkish. SIZE (4 YY, 3 XX): wing, Y 72–75 (738), X 69–72 (710); tail, Y 43–47 (448), X 43–46 (443); bill, Y 175–200 (190), X 172–186 (178); tarsus, Y 204–209 (206), X 185–191 (189). WEIGHT: Y (n ¼ 17) 225–270 (246); X (n ¼ 16) 208–246 (225). IMMATURE: dull greenish above, hardly streaked; underparts washed with dull yellow; bill brown. Sub-adult Y olive and yellowish, head with black patches and trace of superciliary stripe. NESTLING: not described.

extremely restricted range, by pale yellow and white belly, olive rump and tail, yellow wing feather edgings; XX cannot be confused because X Dark-backed is like Y, while X Clarke’s a typical X weaver with streaked upperparts, yellow underparts, yellow wing edgings.

Field Characters. Length 135 cm. Y has black head, breast and back, dark eye, slender bill; told from Darkbacked Weaver P. bicolor, the only other forest weaver in its

Voice. Tape-recorded (B, CART, KEI, McVIC). Foraging flocks keep up continual choruses of chattering, chirping and sizzling sounds which rise and fall in volume, audible

Ploceus dicrocephalus for several hundred metres; rendered ‘ch-ch-sss-sss-sss’ or ‘zsss-zs-zs-zs’ (Zimmerman et al. 1996). General Habits. In all habitats within Sokoke Forest, but commonest in Brachystegia woodland, feeding in canopy. Occurs in noisy flocks of 5–30, sometimes up to 100 birds. In mixed-species flocks with Scimitarbill Phoeniculus cyanomelas, Black-headed Oriole Oriolus larvatus, Black Cuckoo-Shrike Campephaga flava, and Dark-backed Weaver Ploceus bicolor (Britton and Zimmerman 1979). Often associates with Retz’s Helmet-Shrike Prionops retzi and Chestnut-fronted Helmet-Shrike P. scopifrons; other species apparently hawk insects flushed by the weavers, which are shy and restless. Single-sex flocks at times. Flight described as almost quelea-like (Short and Horne 1985). At times conspicuous and gregarious, yet flocks appear erratically. Resident (Collar et al. 1994).

173

Food. Fruits of Brachystegia spiciformis, Loranthus; insects, including cockchafers and other beetles (Coleoptera), larvae (Lepidoptera) and scale insects (Hemiptera).

Breeding Habits. Unrecorded; nest and eggs undescribed. Possibly nests high in trees, at low density, perhaps from Feb–Mar when rains begin, as juveniles noted in Apr, July and Aug, and juv. being fed Mar (Marafa: J. B. Fisher, pers. comm.); however, Y in breeding condition in Sept (Williams 1957).

Key References Clancey, P. A. and Williams, J. G. (1959), Stevenson, T. and Fanshawe, J. (2002), Taylor, P. B. (1984), Williams, J. G. (1957).

Ploceus dicrocephalus (Salvadori). Juba Weaver. Tisserin de Salvadori.

Plate 9

Hyphantornis dicrocephala Salvadori, 1896. Ann. Mus. Civ. Genova, 16 (2), p. 45; Somalia.

(Opp. p. 138)

Range and Status. Endemic resident. Ethiopia, local in S Rift Valley as at Genale R., Dawa-Wachile. Kenya, locally common in Daua Valley immediately west of Mandara. Somalia, uncommon on Jubba R; and mid-section of Shabeelle R. Description. ADULT Y (breeding): forehead, crown and nape blackish brown; sharp dividing line to mantle, which is greenish yellow and unstreaked; rump yellow, contrasting with mantle; tail olive-green with yellow margins; wings olive-brown with narrow yellow edges on remiges, broad yellow margins on tertials and coverts, yellow tips with chestnut spots on upperwingcoverts, bright yellow inner webs to remiges; lores, cheeks, earcoverts, chin and throat blackish brown, with central bib blending into rich chestnut-brown on breast; flanks chestnutbrown; belly yellow in centre, with some chestnut wash; thighs yellow with chestnut wash; undertail-coverts yellow. Bill black; eyes red-brown to red; legs brown. ADULT Y (non-breeding): no information; presumed to resemble X. ADULT X: forehead, crown and nape dull olive-brown; mantle olive-brown with indistinct dark centres to feathers; rump lighter brown, but no sharp contrast; tail olive-green with yellow margins; wings dull olive with narrow yellow outer margins, broad yellow inner webs; tertials and wing-coverts with yellowish buff margins; supercilium buff; cheeks and ear-coverts light brown; chin and throat whitish; breast white with buffy wash; belly off-white with buffy wash on flanks, thighs and undertail-coverts. Upper mandible dark brown, lower mandible light brown; eyes brown; legs brown. SIZE (8 YY, 5 XX): wing, Y 69–75 (730), X 66–68 (664); tail, Y 47–53 (493), X 44–47 (456); bill, Y 168–188 (179), X 158–166 (162); tarsus, Y 192–215 (203), X 181–191 (188). WEIGHT: 2 XX (Kenya) 18, 19. IMMATURE: like X but upperparts much browner, underparts with strong buffy wash. NESTLING: not described.

Field Characters. Length 13 cm. Y golden yellow with entire head to nape and breast dark chestnut to blackish; plain yellow-green back lacks streaking of Village Weaver

Ploceus dicrocephalus

P. cucullatus. X lacks yellow on underparts, has buff breast and flanks, brown eye; best told from similar X Vitelline Masked Weaver P. vitellinus by 2-tone bill (dark above, light below). Voice. Not described. General Habits. Inhabits lush riverine bush in Kenya, short grass savanna, Acacia-Commiphora thornbush and semi-desert savanna, mainly below 1500 m, in Ethiopia.

Ploceus dicrocephalus for several hundred metres; rendered ‘ch-ch-sss-sss-sss’ or ‘zsss-zs-zs-zs’ (Zimmerman et al. 1996). General Habits. In all habitats within Sokoke Forest, but commonest in Brachystegia woodland, feeding in canopy. Occurs in noisy flocks of 5–30, sometimes up to 100 birds. In mixed-species flocks with Scimitarbill Phoeniculus cyanomelas, Black-headed Oriole Oriolus larvatus, Black Cuckoo-Shrike Campephaga flava, and Dark-backed Weaver Ploceus bicolor (Britton and Zimmerman 1979). Often associates with Retz’s Helmet-Shrike Prionops retzi and Chestnut-fronted Helmet-Shrike P. scopifrons; other species apparently hawk insects flushed by the weavers, which are shy and restless. Single-sex flocks at times. Flight described as almost quelea-like (Short and Horne 1985). At times conspicuous and gregarious, yet flocks appear erratically. Resident (Collar et al. 1994).

173

Food. Fruits of Brachystegia spiciformis, Loranthus; insects, including cockchafers and other beetles (Coleoptera), larvae (Lepidoptera) and scale insects (Hemiptera).

Breeding Habits. Unrecorded; nest and eggs undescribed. Possibly nests high in trees, at low density, perhaps from Feb–Mar when rains begin, as juveniles noted in Apr, July and Aug, and juv. being fed Mar (Marafa: J. B. Fisher, pers. comm.); however, Y in breeding condition in Sept (Williams 1957).

Key References Clancey, P. A. and Williams, J. G. (1959), Stevenson, T. and Fanshawe, J. (2002), Taylor, P. B. (1984), Williams, J. G. (1957).

Ploceus dicrocephalus (Salvadori). Juba Weaver. Tisserin de Salvadori.

Plate 9

Hyphantornis dicrocephala Salvadori, 1896. Ann. Mus. Civ. Genova, 16 (2), p. 45; Somalia.

(Opp. p. 138)

Range and Status. Endemic resident. Ethiopia, local in S Rift Valley as at Genale R., Dawa-Wachile. Kenya, locally common in Daua Valley immediately west of Mandara. Somalia, uncommon on Jubba R; and mid-section of Shabeelle R. Description. ADULT Y (breeding): forehead, crown and nape blackish brown; sharp dividing line to mantle, which is greenish yellow and unstreaked; rump yellow, contrasting with mantle; tail olive-green with yellow margins; wings olive-brown with narrow yellow edges on remiges, broad yellow margins on tertials and coverts, yellow tips with chestnut spots on upperwingcoverts, bright yellow inner webs to remiges; lores, cheeks, earcoverts, chin and throat blackish brown, with central bib blending into rich chestnut-brown on breast; flanks chestnutbrown; belly yellow in centre, with some chestnut wash; thighs yellow with chestnut wash; undertail-coverts yellow. Bill black; eyes red-brown to red; legs brown. ADULT Y (non-breeding): no information; presumed to resemble X. ADULT X: forehead, crown and nape dull olive-brown; mantle olive-brown with indistinct dark centres to feathers; rump lighter brown, but no sharp contrast; tail olive-green with yellow margins; wings dull olive with narrow yellow outer margins, broad yellow inner webs; tertials and wing-coverts with yellowish buff margins; supercilium buff; cheeks and ear-coverts light brown; chin and throat whitish; breast white with buffy wash; belly off-white with buffy wash on flanks, thighs and undertail-coverts. Upper mandible dark brown, lower mandible light brown; eyes brown; legs brown. SIZE (8 YY, 5 XX): wing, Y 69–75 (730), X 66–68 (664); tail, Y 47–53 (493), X 44–47 (456); bill, Y 168–188 (179), X 158–166 (162); tarsus, Y 192–215 (203), X 181–191 (188). WEIGHT: 2 XX (Kenya) 18, 19. IMMATURE: like X but upperparts much browner, underparts with strong buffy wash. NESTLING: not described.

Field Characters. Length 13 cm. Y golden yellow with entire head to nape and breast dark chestnut to blackish; plain yellow-green back lacks streaking of Village Weaver

Ploceus dicrocephalus

P. cucullatus. X lacks yellow on underparts, has buff breast and flanks, brown eye; best told from similar X Vitelline Masked Weaver P. vitellinus by 2-tone bill (dark above, light below). Voice. Not described. General Habits. Inhabits lush riverine bush in Kenya, short grass savanna, Acacia-Commiphora thornbush and semi-desert savanna, mainly below 1500 m, in Ethiopia.

174

PLOCEIDAE

Spends much time in vegetation emergent from river, so possibly overlooked (Somalia: Ash and Miskell 1998). Food. Seeds, probably also insects. Breeding Habits. Colonial, nesting in small groups in trees. NEST: oval, with virtually no tunnel, woven of grass, lined with fine grass, suspended from a single reed at each side. SIZE: 90 high by 115 long.

EGGS: 2, pale greenish blue or greyish green, spotted and freckled with brown and lilac. SIZE: (n ¼ 1, Ethiopia) 205  145. LAYING DATES: Ethiopia, Mar. Nothing further known

Key Reference Benson, C. W. (1947).

Plate 9

Ploceus melanocephalus (Linnaeus). Yellow-backed Weaver. Tisserin a` te ˆte noire.

(Opp. p. 138)

Loxia melanocephala Linnaeus, 1758. Syst. Nat., ed. 10, p. 175; Guinea. Range and Status. Endemic resident. Mauritania, Senegal R. area, coast near Nouakchott in rains. Senegal, Senegal R. and delta, L. de Guier, in S at Toubakouta and near permanent water. Gambia, localized on coast, common along riverbanks. Guinea-Bissau, local near coast. Guinea, Senegal R. and Niger R. watershed. Mali, Boucle de Baoule´, Ansongo, common between 12
and 15
N. Burkina Faso, southern and central region, north to Ouagadougou. Ghana, 1 Mole, 1968. Togo, Sio R., Tokpli, Oti R. in north. Benin, on rivers as far inland as Be´te´rou. Niger, Niamey north along Niger R. to Tillaberi. Nigeria, Sokoto to L. Chad, along Niger R. and Benue R. and tributaries, south to Onitsha, also old records from Niger delta. Cameroon, from Benue Plain north, not south of Buffle Noir. Chad, L. Chad south along SE and S border. N Central African Republic, Mboko, Landjia, Ge´ringou, Kapou; S CAR, Lobaye region. Congo, along Congo R. and Ubangi R., Zaı¨re, along Congo R. from Kinshasa to Kisangani, Uele R., Maniema, in E from L. Albert to L. Kivu. Sudan, Kassala, S Equatoria. Eritrea, W lowlands. Rwanda, Burundi, Uganda, widespread except Karamoja. Kenya, locally common on L. Victoria, L. Kanyaboli and Nzoia R. upstream to Mumias. Tanzania, current records only from Mwanza area. Zambia, around L. Mweru, Chisenga and Kilwa islands, Luapula R. Common over much of range. Description. P. m. melanocephalus S Mauritania, N Senegal, Mali, Niger. ADULT Y (breeding): forehead, crown, lores and cheeks black; narrow yellow collar on nape; mantle greenish yellow, unstreaked; rump yellow; tail olive-green; chin and throat black, black extending as bib down to centre of breast; breast yellow except for black bib and some chestnut-brown at margins of bib (sometimes absent); flanks, belly, thighs and undertail-coverts yellow; wings brown with yellow rims to primary remiges, broad yellow edges to wing-coverts, underwing grey. Bill black; eyes brown; legs brown. ADULT Y (non-breeding): forehead, crown and nape dull olive-green; mantle brown with broad, dark central streaks on feathers; rump plain brown with greenish wash; tail olive-green; small yellowish supercilium, cheeks and ear-coverts greenish yellow; chain and throat dull white with yellowish tinge; breast washed with buffy grey; flanks and thighs buffy; belly and undertail-coverts dull white; wing brown with paler edges to remiges and coverts; underwing cream. Upper mandible black, lower horn-coloured; eyes brown; legs brown. ADULT X: like non-breeding Y except that yellow edges of wing feathers and

Ploceus melanocephalus

yellow throat are duller. May hybridize with P. castanops (Louette 1987). SIZE (12 YY, 10 XX): wing, Y 73–79 (758), X 61–68 (653); tail, Y 46–53 (501), X 38–47 (427); bill, Y 170–197 (187), X 157–188 (167); tarsus, Y 199–226 (215), X 180–197 (189). WEIGHT: (Senegal, unsexed) 19–22. IMMATURE: like X, except that underparts pale buffy, broad buffy margins to wing-coverts, less streaking on mantle. Captive-bred Y did not acquire breeding plumage in first year (Schu ¨ rer 1978). NESTLING: crown and nape olive-green, mantle brown with darker streaks, rump plain brown; chin, throat and upper breast dull yellow; flanks, thighs, belly and undertail-coverts saffron buff (Serle 1957). P. m. capitalis (Latham): Guinea-Bissau, Guinea, Togo, Nigeria to Cameroon, Chad, N Central African Republic. Breeding Y with much more chestnut below, yellower on upperparts; X more buffy. WEIGHT: (Niger) Y (n ¼ 17) 201–250 (225); X (n ¼ 11) 170–211 (191). P. m. dimidiatus (Antinori and Salvadori) (includes ‘fischeri’): Eritrea, Sudan, E Zaı¨re, Uganda, Kenya, Tanzania, Zambia.

174

PLOCEIDAE

Spends much time in vegetation emergent from river, so possibly overlooked (Somalia: Ash and Miskell 1998). Food. Seeds, probably also insects. Breeding Habits. Colonial, nesting in small groups in trees. NEST: oval, with virtually no tunnel, woven of grass, lined with fine grass, suspended from a single reed at each side. SIZE: 90 high by 115 long.

EGGS: 2, pale greenish blue or greyish green, spotted and freckled with brown and lilac. SIZE: (n ¼ 1, Ethiopia) 205  145. LAYING DATES: Ethiopia, Mar. Nothing further known

Key Reference Benson, C. W. (1947).

Plate 9

Ploceus melanocephalus (Linnaeus). Yellow-backed Weaver. Tisserin a` te ˆte noire.

(Opp. p. 138)

Loxia melanocephala Linnaeus, 1758. Syst. Nat., ed. 10, p. 175; Guinea. Range and Status. Endemic resident. Mauritania, Senegal R. area, coast near Nouakchott in rains. Senegal, Senegal R. and delta, L. de Guier, in S at Toubakouta and near permanent water. Gambia, localized on coast, common along riverbanks. Guinea-Bissau, local near coast. Guinea, Senegal R. and Niger R. watershed. Mali, Boucle de Baoule´, Ansongo, common between 12
and 15
N. Burkina Faso, southern and central region, north to Ouagadougou. Ghana, 1 Mole, 1968. Togo, Sio R., Tokpli, Oti R. in north. Benin, on rivers as far inland as Be´te´rou. Niger, Niamey north along Niger R. to Tillaberi. Nigeria, Sokoto to L. Chad, along Niger R. and Benue R. and tributaries, south to Onitsha, also old records from Niger delta. Cameroon, from Benue Plain north, not south of Buffle Noir. Chad, L. Chad south along SE and S border. N Central African Republic, Mboko, Landjia, Ge´ringou, Kapou; S CAR, Lobaye region. Congo, along Congo R. and Ubangi R., Zaı¨re, along Congo R. from Kinshasa to Kisangani, Uele R., Maniema, in E from L. Albert to L. Kivu. Sudan, Kassala, S Equatoria. Eritrea, W lowlands. Rwanda, Burundi, Uganda, widespread except Karamoja. Kenya, locally common on L. Victoria, L. Kanyaboli and Nzoia R. upstream to Mumias. Tanzania, current records only from Mwanza area. Zambia, around L. Mweru, Chisenga and Kilwa islands, Luapula R. Common over much of range. Description. P. m. melanocephalus S Mauritania, N Senegal, Mali, Niger. ADULT Y (breeding): forehead, crown, lores and cheeks black; narrow yellow collar on nape; mantle greenish yellow, unstreaked; rump yellow; tail olive-green; chin and throat black, black extending as bib down to centre of breast; breast yellow except for black bib and some chestnut-brown at margins of bib (sometimes absent); flanks, belly, thighs and undertail-coverts yellow; wings brown with yellow rims to primary remiges, broad yellow edges to wing-coverts, underwing grey. Bill black; eyes brown; legs brown. ADULT Y (non-breeding): forehead, crown and nape dull olive-green; mantle brown with broad, dark central streaks on feathers; rump plain brown with greenish wash; tail olive-green; small yellowish supercilium, cheeks and ear-coverts greenish yellow; chain and throat dull white with yellowish tinge; breast washed with buffy grey; flanks and thighs buffy; belly and undertail-coverts dull white; wing brown with paler edges to remiges and coverts; underwing cream. Upper mandible black, lower horn-coloured; eyes brown; legs brown. ADULT X: like non-breeding Y except that yellow edges of wing feathers and

Ploceus melanocephalus

yellow throat are duller. May hybridize with P. castanops (Louette 1987). SIZE (12 YY, 10 XX): wing, Y 73–79 (758), X 61–68 (653); tail, Y 46–53 (501), X 38–47 (427); bill, Y 170–197 (187), X 157–188 (167); tarsus, Y 199–226 (215), X 180–197 (189). WEIGHT: (Senegal, unsexed) 19–22. IMMATURE: like X, except that underparts pale buffy, broad buffy margins to wing-coverts, less streaking on mantle. Captive-bred Y did not acquire breeding plumage in first year (Schu ¨ rer 1978). NESTLING: crown and nape olive-green, mantle brown with darker streaks, rump plain brown; chin, throat and upper breast dull yellow; flanks, thighs, belly and undertail-coverts saffron buff (Serle 1957). P. m. capitalis (Latham): Guinea-Bissau, Guinea, Togo, Nigeria to Cameroon, Chad, N Central African Republic. Breeding Y with much more chestnut below, yellower on upperparts; X more buffy. WEIGHT: (Niger) Y (n ¼ 17) 201–250 (225); X (n ¼ 11) 170–211 (191). P. m. dimidiatus (Antinori and Salvadori) (includes ‘fischeri’): Eritrea, Sudan, E Zaı¨re, Uganda, Kenya, Tanzania, Zambia.

Ploceus jacksoni Breeding Y has rich chestnut underparts, yellow restricted to the belly. WEIGHT: Y (Kenya, n ¼ 57) 210–363 (287), (Uganda, n ¼ 35) 200–305 (267), X (Kenya, n ¼ 24) 199–275 (228), (Uganda, n ¼ 37) 182–270 (213). P. m. duboisi Hartlaub: Congo, S Central African Republic and N Zaı¨re on middle Congo and Ubangi R. Breeding Y with yellow underparts, lacking any brown wash; bill longer, more slender than in other populations. WEIGHT: (Zaı¨re) Y (n ¼ 32) 200–294 (225), X (n ¼ 10) 15–19 (17).

Field Characters. Length 14–16 cm. Breeding Y has solid black head; underparts vary from pure bright yellow in W to chestnut with yellow belly in E; differs from very similar Golden-backed Weaver P. jacksoni by yellow hind collar contrasting with plain green back, slender bill with curved culmen, brown (not red) eye. Larger Village Weaver P. cucullatus has yellow back with black ‘V’, red eye, heavy bill. X much browner than X Golden-backed, no yellow on underparts except for wash on throat. In W Africa X and non-breeding Y very like Vitelline Masked Weaver P. vitellinus but have pale (not orange) eye, 2-tone bill (black above, pinkish brown below); waterside habitat also useful (Barlow et al. 1997). Voice. Tape-recorded (104, B, CHA, McVIC, MOR). Song a brief medley of thin, high-pitched notes, clear and sharp, followed by hissing call; also rendered ‘squeee-ki-kee’ (Barlow et al. 1997). Calls, harsh grating ‘jaayaa’ or ‘djick’. General Habits. Inhabits damp areas with tall grass along rivers and edges of standing water; also away from water in tree-savanna, especially with Ziziphus and Acacia. In Zambia, restricted to reedbeds and papyrus, though may forage in surrounding drier country. Generally below 1500 m (Ethiopia). Occurs in pairs or small parties, forming larger flocks in non-breeding season. Sedentary. A complete prenuptial moult (Verheyen 1953); testes start enlarging once tertials are growing, but are inactive during primary wing-moult. Food. Seeds and, particularly while feeding young, insects. Breeding Habits. Colonial or solitary nester; monogamous or sometimes polygynous, with 2–3 XX simultaneously in same territory (Mundy and Cook 1974). Breeds alongside Slender-billed Weaver P. pelzelni and Village Weaver P. cucullatus, also Southern Red Bishop Euplectes orix and Lesser Masked Weaver P. intermedius. In S Zaı¨re, nests in

175

ambatch Aeschynomene elaphroxylon are c. 10 m apart and apparently solitary. Highly territorial; dominates smaller species, but displaced by Y Fan-tailed Widowbird Euplectes axillaris. Neighbouring YY threaten each other with nape feathers erected, wings and tail spread, and wing-flicking, may fall into water during fights. Y uses short song in territorial encounters, in courtship away from the nest, and when X enters nest; there is a distinctive vocalization during nest advertisement display (Crook 1969). When X appears, Y hangs below nest entrance, with open fluttering wings. Displays generally similar to those of P. cucullatus. NEST: round or onion-shaped, tightly woven of grasses, and lined with fine grass tops; no tunnel around entrance which faces vertically downwards. Placed near tops of elephant grass stalks, in papyrus or reeds in water, also in bushes and small trees and maize in cultivated fields, generally >2 m (range 1–5 m) from ground or water. SIZE: 120–140 long, 95–105 high, entrance diam. 35. Leaves stripped around nest, making it conspicuous. Nest may be slung between upright stems, supported at 1 side only, or suspended from twigs or leaves. Outer shell of strips of grass blades built by Y, then inner shell of grass heads added by Y and X; a few feathers in lining. Distinct ceiling layer, with strips of grass leaf. EGGS: 2–3. Variable: dull white, blue-green, pinkish grey or brown, plain or heavily spotted or freckled with olive-brown. SIZE: (n ¼ 10, Nigeria) 175–202  132–142 (189  138), (n ¼ 2, Zaı¨re) 199  147. LAYING DATES: Mauritania, May–Oct; Senegal, Apr– Nov; Gambia, June–Sept; Mali, July–Oct; Burkina Faso, July–Aug at Ouagadougou, elsewhere Aug–Oct; Togo, Aug (breeding plumage Sept, nest-building Oct); Nigeria, May, July–Oct; Niger, (colony active June–Nov); Zaı¨re, June– Aug in NE, Kimbombo, Nov–Apr, Itombwe and E, Jan– May, Aug–Sept in central region; Sudan, Aug–Sept; Ethiopia, (breeding July–Sept); Uganda, all months except Aug, Nov, with peaks Feb, Apr, Sept; Zambia, Feb–Mar. INCUBATION: by X only. DEVELOPMENT AND CARE OF YOUNG: both Y and X feed nestlings and fledglings. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius. Key References Collias, N. E. and Collias, E. C. (1964), Guichard, K. M. (1947).

Ploceus jacksoni Shelley. Golden-backed Weaver. Tisserin a ` dos d’or.

Plate 9

Ploceus jacksoni Shelley, 1888. Ibis, 1888, p. 293; Kilimanjaro, Tanzania.

(Opp. p. 138)

Range and Status. Endemic resident. Sudan, rare: only in Dongatona and Imatong Mts. Uganda, Kidepo Valley, Agaru and Kabalega Falls south to L. Victoria, west to Masaka. Burundi, NE shores of L. Tanganyika. Kenya, S Turkana region to Kerio Valley, L. Baringo, Solai, Nyanza, Kilgoris, Amboseli and Namanga, L. Jipe and Konza. Tanzania, local on coast north of Dar es Salaam, Mwanza,

Bibondo east and south to Kilimanjaro, Arusha, Marogoro, Ruaha, Njombe. Locally common. Description. ADULT Y: forehead, crown and nape black, ending in sharp line; mantle chrome yellow, with variable greenish wash or streaking at posterior end; rump chrome yellow; tail olivegreen with narrow yellow margins; chin, lores, cheeks, ear-coverts

Ploceus jacksoni Breeding Y has rich chestnut underparts, yellow restricted to the belly. WEIGHT: Y (Kenya, n ¼ 57) 210–363 (287), (Uganda, n ¼ 35) 200–305 (267), X (Kenya, n ¼ 24) 199–275 (228), (Uganda, n ¼ 37) 182–270 (213). P. m. duboisi Hartlaub: Congo, S Central African Republic and N Zaı¨re on middle Congo and Ubangi R. Breeding Y with yellow underparts, lacking any brown wash; bill longer, more slender than in other populations. WEIGHT: (Zaı¨re) Y (n ¼ 32) 200–294 (225), X (n ¼ 10) 15–19 (17).

Field Characters. Length 14–16 cm. Breeding Y has solid black head; underparts vary from pure bright yellow in W to chestnut with yellow belly in E; differs from very similar Golden-backed Weaver P. jacksoni by yellow hind collar contrasting with plain green back, slender bill with curved culmen, brown (not red) eye. Larger Village Weaver P. cucullatus has yellow back with black ‘V’, red eye, heavy bill. X much browner than X Golden-backed, no yellow on underparts except for wash on throat. In W Africa X and non-breeding Y very like Vitelline Masked Weaver P. vitellinus but have pale (not orange) eye, 2-tone bill (black above, pinkish brown below); waterside habitat also useful (Barlow et al. 1997). Voice. Tape-recorded (104, B, CHA, McVIC, MOR). Song a brief medley of thin, high-pitched notes, clear and sharp, followed by hissing call; also rendered ‘squeee-ki-kee’ (Barlow et al. 1997). Calls, harsh grating ‘jaayaa’ or ‘djick’. General Habits. Inhabits damp areas with tall grass along rivers and edges of standing water; also away from water in tree-savanna, especially with Ziziphus and Acacia. In Zambia, restricted to reedbeds and papyrus, though may forage in surrounding drier country. Generally below 1500 m (Ethiopia). Occurs in pairs or small parties, forming larger flocks in non-breeding season. Sedentary. A complete prenuptial moult (Verheyen 1953); testes start enlarging once tertials are growing, but are inactive during primary wing-moult. Food. Seeds and, particularly while feeding young, insects. Breeding Habits. Colonial or solitary nester; monogamous or sometimes polygynous, with 2–3 XX simultaneously in same territory (Mundy and Cook 1974). Breeds alongside Slender-billed Weaver P. pelzelni and Village Weaver P. cucullatus, also Southern Red Bishop Euplectes orix and Lesser Masked Weaver P. intermedius. In S Zaı¨re, nests in

175

ambatch Aeschynomene elaphroxylon are c. 10 m apart and apparently solitary. Highly territorial; dominates smaller species, but displaced by Y Fan-tailed Widowbird Euplectes axillaris. Neighbouring YY threaten each other with nape feathers erected, wings and tail spread, and wing-flicking, may fall into water during fights. Y uses short song in territorial encounters, in courtship away from the nest, and when X enters nest; there is a distinctive vocalization during nest advertisement display (Crook 1969). When X appears, Y hangs below nest entrance, with open fluttering wings. Displays generally similar to those of P. cucullatus. NEST: round or onion-shaped, tightly woven of grasses, and lined with fine grass tops; no tunnel around entrance which faces vertically downwards. Placed near tops of elephant grass stalks, in papyrus or reeds in water, also in bushes and small trees and maize in cultivated fields, generally >2 m (range 1–5 m) from ground or water. SIZE: 120–140 long, 95–105 high, entrance diam. 35. Leaves stripped around nest, making it conspicuous. Nest may be slung between upright stems, supported at 1 side only, or suspended from twigs or leaves. Outer shell of strips of grass blades built by Y, then inner shell of grass heads added by Y and X; a few feathers in lining. Distinct ceiling layer, with strips of grass leaf. EGGS: 2–3. Variable: dull white, blue-green, pinkish grey or brown, plain or heavily spotted or freckled with olive-brown. SIZE: (n ¼ 10, Nigeria) 175–202  132–142 (189  138), (n ¼ 2, Zaı¨re) 199  147. LAYING DATES: Mauritania, May–Oct; Senegal, Apr– Nov; Gambia, June–Sept; Mali, July–Oct; Burkina Faso, July–Aug at Ouagadougou, elsewhere Aug–Oct; Togo, Aug (breeding plumage Sept, nest-building Oct); Nigeria, May, July–Oct; Niger, (colony active June–Nov); Zaı¨re, June– Aug in NE, Kimbombo, Nov–Apr, Itombwe and E, Jan– May, Aug–Sept in central region; Sudan, Aug–Sept; Ethiopia, (breeding July–Sept); Uganda, all months except Aug, Nov, with peaks Feb, Apr, Sept; Zambia, Feb–Mar. INCUBATION: by X only. DEVELOPMENT AND CARE OF YOUNG: both Y and X feed nestlings and fledglings. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius. Key References Collias, N. E. and Collias, E. C. (1964), Guichard, K. M. (1947).

Ploceus jacksoni Shelley. Golden-backed Weaver. Tisserin a ` dos d’or.

Plate 9

Ploceus jacksoni Shelley, 1888. Ibis, 1888, p. 293; Kilimanjaro, Tanzania.

(Opp. p. 138)

Range and Status. Endemic resident. Sudan, rare: only in Dongatona and Imatong Mts. Uganda, Kidepo Valley, Agaru and Kabalega Falls south to L. Victoria, west to Masaka. Burundi, NE shores of L. Tanganyika. Kenya, S Turkana region to Kerio Valley, L. Baringo, Solai, Nyanza, Kilgoris, Amboseli and Namanga, L. Jipe and Konza. Tanzania, local on coast north of Dar es Salaam, Mwanza,

Bibondo east and south to Kilimanjaro, Arusha, Marogoro, Ruaha, Njombe. Locally common. Description. ADULT Y: forehead, crown and nape black, ending in sharp line; mantle chrome yellow, with variable greenish wash or streaking at posterior end; rump chrome yellow; tail olivegreen with narrow yellow margins; chin, lores, cheeks, ear-coverts

176

PLOCEIDAE

Ploceus jacksoni

and throat black, extending into bib on breast; breast, apart from central black bib, chestnut-brown; flanks yellow under wings, rest chestnut-brown; belly chestnut-brown, in centre yellow with brown wash; thighs and undertail-coverts chrome yellow; wings dull brown, narrow yellow margins on remiges, bright yellow edges of coverts, and yellow tips to upperwing-coverts; underwing yellow. Bill black; eyes crimson; legs brown. ADULT Y (nonbreeding): forehead, crown and nape dull olive-brown; mantle olive-brown with broad, dark central streaks; rump plain greenish brown; tail olive-green with paler margins; supercilium clear yellow, cheeks and ear-coverts olive-brown; chin and throat yellowish white; upper breast with buff band, lower breast, belly and undertail-coverts white with yellow wash; thighs and flanks buffy; wings brown with yellowish or buffy edges to coverts; underwing yellow. Bill dark brown upper mandible, pale brown lower mandible; eyes red; legs brown. ADULT X: in breeding dress, like non-breeding Y, but eyes dark brown. Non-breeding X apparently lacks yellow wash on underparts, upperparts duller and greyer rather than green. Bill upper mandible dark greyish, lower mandible light grey-brown; eyes dark brown; legs brown. May hybridize with P. castanops (Louette 1987). SIZE (12 YY, 12 XX): wing, Y 75–78 (757), X 66–72 (708); tail, Y 47–51 (490), X 45–50 (474); bill, Y 166–188 (175), X 160–179 (168); tarsus, Y 187–214 (203), X 178–208 (193). WEIGHT: Y (Kenya, n ¼ 38) 223–300 (267), (Uganda, n ¼ 22) 260–310 (274); X (Kenya, n ¼ 22) 1889–223 (205), (Uganda, n ¼ 18) 190–273 (215). IMMATURE: like non-breeding X, but more buffy on underparts, bill paler brown. NESTLING: not described.

Field Characters. Length 14–15 cm. Breeding Y with black head and chestnut and yellow underparts very like Yellow-backed Weaver P. melanocephalus but black of head reaches further down onto back of neck and meets solid yellow back (no contrasting yellow hind-collar); eye red, culmen straight. X and non-breeding Y are olive above and extensively yellow below, eye reddish.

Voice. Tape-recorded (B, GREG, McVIC, STJ). Chattering chorus at colony includes a variety of notes, harsh ‘chee’, ‘chaa’, chit’ and ‘chuk’, thin, down-slurred ‘tzeew’, grating ‘churrrrr’, dry ‘bzzzzz’, clicking ‘tikatikatikatika . . . ’ and a few swizzles. Short song used in nest advertisement display, long song during territorial encounters and when X enters nest (Crook 1969).

Ploceus rubiginosus General Habits. In swamp, ambatch, reeds, papyrus, and riverine habitats, also acacia scrub and woodland, up to 1800 m. In Kenya, irregular irruptive movements in response to heavy rainfall. Locally sympatric with P. melanocephalus in L. Victoria basin. Food. Seeds, and presumably also insects. Breeding Habits. Colonial, often nesting alongside other weavers such as Village Weaver P. cucullatus, Orange Weaver P. aurantius, P. melanocephalus, Slender-billed Weaver P. pelzelni; polygynous. NEST: oval, compact, woven of strips torn from grass or palm leaves (A, unfinished; B, finished), lined with grass

177

tops and some feathers. Built over water in papyrus or reeds; also in acacia trees, ambatch bushes and in maize fields. SIZE: 125 long, 105–110 high, entrance diam. 35–40 (Collias and Collias 1964). EGGS: 2–3, blue, with purple-brown or dark red spots, which may be very fine and dense, almost obscuring ground colour. Very like eggs of P. melanocephalus (van Someren 1916). SIZE: 199–212  135–150 (Chapin 1954). LAYING DATES: Uganda, all months except July, Sept– Oct, but clear peak in Apr–May; Kenya, Mar–June; Tanzania, Sept, Nov–June, (building Sept, Whybrow 1948). BREEDING SUCCESS/SURVIVAL: Y ringed in Kenya, recovered 55 years later, 28 km from ringing site (Backhurst 1977).

Ploceus rubiginosus Ru ¨ ppell. Chestnut Weaver. Tisserin roux.

Plate 11

Ploceus rubiginosus Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, pp. 93, 100; Abyssinia.

(Opp. p. 154)

Forms a superspecies with P. badius. Range and Status. Endemic, local migrant or at least nomadic seasonally. Sudan, Ero, Lokorowa valley (possibly non-breeding visitors only). Ethiopia, southern and SE region, NE and Rift Valley. Eritrea, in W region. Uganda, Karamoja. Kenya, widespread except for L. Victoria basin, coast, and highlands. Tanzania, interior plateau, Mkomazi and L. Natron, south and west to Dodoma, Ruaha Nat. Park, Singida, E Talwa, Shinyanga. Somalia, regular in NW, west of 44
300 E, in S along Shabeelle R. but very rare on Jubba River. Angola, on coast from Catumbela south, SW Namibe and Cunene to S Huı´la. Namibia, from Windhoek northwards on central plateau. Botswana, at Gomare, Nokaneng and near Maun, apparently regular visitor to Okavango Delta. Common; locally abundant. Description. P. r. rubiginosus Ru ¨ ppell: E Africa. ADULT Y (breeding): forehead, crown, lores, cheeks and ear-coverts black; sharp line on nape at border with chestnut-brown which extends to mantle and rump, with some mantle feathers near wings having dark central streaks; tail dark brown; chin and throat black, forming curved bib; breast, flanks, belly and undertailcoverts chestnut-brown; wings dark brown, with paler edges to wing-coverts. Bill black; legs brown; eyes cinnamon-brown to orange-red. ADULT Y (non-breeding): forehead, crown and nape grey-brown with faint darker central streaks; mantle brown (sometimes washed with chestnut) with very dark central streaks; rump light brown, unstreaked; tail brown; sides of head light brown to buff, sharp line at level of lower mandible; chin and throat white; breast with a buffy collar; flanks and thighs with buffy wash, varying individually, sometimes pale chestnut; belly and undertail-coverts white; wings brown, as in breeding Y. Bill brown, upper mandible darker. ADULT X: like non-breeding Y, but always light brown or buffy rather than chestnut. SIZE (12 YY, 12 XX): wing, Y 78–86 (825), X 71–79 (753); tail, Y 52–61 (561), X 47–55 (502); bill, Y 197–210 (204), X 177–202 (191); tarsus, Y 198–224 (213), X 186–210 (200). WEIGHT: (Kenya) Y (n ¼ 15) 292–370 (324); X (n ¼ 8) 253–312 (280). IMMATURE: like X, but streaking on breast. Bill pale brown; eyes brown. NESTLING: flesh-coloured, almost naked (Archibald 1987).

Ploceus rubiginosus

P. r. trothae Reichenow: Angola, Namibia, Botswana. Chestnut plumage more orange in tone; rump light brown, not chestnut brown. SIZE (42 YY, 29 XX): wing, Y 80–87 (837), X 74–79 (775); tail, Y 47–55 (524), X 45–49 (474); bill, Y 195–225 (209), X 180–210 (196); tarsus, Y 210–245 (229), X 200–225 (214). WEIGHT: Y (n ¼ 39) 28–37 (321); X (n ¼ 55) 25–30 (278) (Komen 1990). TAXONOMIC NOTE: Clancey (1989) proposed 2 subspecies in western population; refuted by Komen (1990).

Field Characters. Length 14–15 cm. Breeding Y with black head and chestnut body much larger than Chestnut Sparrow Passer eminibey; distinguished by red eye, heavy

Ploceus rubiginosus General Habits. In swamp, ambatch, reeds, papyrus, and riverine habitats, also acacia scrub and woodland, up to 1800 m. In Kenya, irregular irruptive movements in response to heavy rainfall. Locally sympatric with P. melanocephalus in L. Victoria basin. Food. Seeds, and presumably also insects. Breeding Habits. Colonial, often nesting alongside other weavers such as Village Weaver P. cucullatus, Orange Weaver P. aurantius, P. melanocephalus, Slender-billed Weaver P. pelzelni; polygynous. NEST: oval, compact, woven of strips torn from grass or palm leaves (A, unfinished; B, finished), lined with grass

177

tops and some feathers. Built over water in papyrus or reeds; also in acacia trees, ambatch bushes and in maize fields. SIZE: 125 long, 105–110 high, entrance diam. 35–40 (Collias and Collias 1964). EGGS: 2–3, blue, with purple-brown or dark red spots, which may be very fine and dense, almost obscuring ground colour. Very like eggs of P. melanocephalus (van Someren 1916). SIZE: 199–212  135–150 (Chapin 1954). LAYING DATES: Uganda, all months except July, Sept– Oct, but clear peak in Apr–May; Kenya, Mar–June; Tanzania, Sept, Nov–June, (building Sept, Whybrow 1948). BREEDING SUCCESS/SURVIVAL: Y ringed in Kenya, recovered 55 years later, 28 km from ringing site (Backhurst 1977).

Ploceus rubiginosus Ru ¨ ppell. Chestnut Weaver. Tisserin roux.

Plate 11

Ploceus rubiginosus Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, pp. 93, 100; Abyssinia.

(Opp. p. 154)

Forms a superspecies with P. badius. Range and Status. Endemic, local migrant or at least nomadic seasonally. Sudan, Ero, Lokorowa valley (possibly non-breeding visitors only). Ethiopia, southern and SE region, NE and Rift Valley. Eritrea, in W region. Uganda, Karamoja. Kenya, widespread except for L. Victoria basin, coast, and highlands. Tanzania, interior plateau, Mkomazi and L. Natron, south and west to Dodoma, Ruaha Nat. Park, Singida, E Talwa, Shinyanga. Somalia, regular in NW, west of 44
300 E, in S along Shabeelle R. but very rare on Jubba River. Angola, on coast from Catumbela south, SW Namibe and Cunene to S Huı´la. Namibia, from Windhoek northwards on central plateau. Botswana, at Gomare, Nokaneng and near Maun, apparently regular visitor to Okavango Delta. Common; locally abundant. Description. P. r. rubiginosus Ru ¨ ppell: E Africa. ADULT Y (breeding): forehead, crown, lores, cheeks and ear-coverts black; sharp line on nape at border with chestnut-brown which extends to mantle and rump, with some mantle feathers near wings having dark central streaks; tail dark brown; chin and throat black, forming curved bib; breast, flanks, belly and undertailcoverts chestnut-brown; wings dark brown, with paler edges to wing-coverts. Bill black; legs brown; eyes cinnamon-brown to orange-red. ADULT Y (non-breeding): forehead, crown and nape grey-brown with faint darker central streaks; mantle brown (sometimes washed with chestnut) with very dark central streaks; rump light brown, unstreaked; tail brown; sides of head light brown to buff, sharp line at level of lower mandible; chin and throat white; breast with a buffy collar; flanks and thighs with buffy wash, varying individually, sometimes pale chestnut; belly and undertail-coverts white; wings brown, as in breeding Y. Bill brown, upper mandible darker. ADULT X: like non-breeding Y, but always light brown or buffy rather than chestnut. SIZE (12 YY, 12 XX): wing, Y 78–86 (825), X 71–79 (753); tail, Y 52–61 (561), X 47–55 (502); bill, Y 197–210 (204), X 177–202 (191); tarsus, Y 198–224 (213), X 186–210 (200). WEIGHT: (Kenya) Y (n ¼ 15) 292–370 (324); X (n ¼ 8) 253–312 (280). IMMATURE: like X, but streaking on breast. Bill pale brown; eyes brown. NESTLING: flesh-coloured, almost naked (Archibald 1987).

Ploceus rubiginosus

P. r. trothae Reichenow: Angola, Namibia, Botswana. Chestnut plumage more orange in tone; rump light brown, not chestnut brown. SIZE (42 YY, 29 XX): wing, Y 80–87 (837), X 74–79 (775); tail, Y 47–55 (524), X 45–49 (474); bill, Y 195–225 (209), X 180–210 (196); tarsus, Y 210–245 (229), X 200–225 (214). WEIGHT: Y (n ¼ 39) 28–37 (321); X (n ¼ 55) 25–30 (278) (Komen 1990). TAXONOMIC NOTE: Clancey (1989) proposed 2 subspecies in western population; refuted by Komen (1990).

Field Characters. Length 14–15 cm. Breeding Y with black head and chestnut body much larger than Chestnut Sparrow Passer eminibey; distinguished by red eye, heavy

178

PLOCEIDAE

bill, pale wing edgings; similar to Cinnamon Weaver P. badius but lacks any green or yellow in plumage. X and non-breeding Y much browner than other weavers, with buff or chestnut breast-band and flanks, heavy black streaks on brown back; in southern Africa grey bill distinctive; juv. similar but breast streaked. Voice. Tape-recorded (99, GREG, KEI, McVIC). Calls include sharp ‘tyoop’, grating ‘jit’ and dry chatter; see sonagram in Maclean (1993). Birds in breeding colony give loud, far-carrying quelea-like sizzling (Zimmerman et al. 1996). General Habits. Inhabits dry thornveld, generally below 1500 m; seasonally abundant, with regular movements in some areas; in places, numbers vary greatly from year to year. Movements generally correlated with regional rainfall patterns. In Aug moves from highlands (e.g. Harar) to lowlands, following rains (Archer and Godman 1961). At Tsavo, Kenya, present Oct–Apr (Lack 1985). In Angola, most frequent in Oct–Feb, but sometimes common all year; at Iona Springs annually in Dec–Jan. Occurrence irregular in N Namibia, with only 2 records in 1952–1954 and 1964–1971 near Windhoek (Braine and Braine 1971), but at other localities birds present throughout the year (Komen and Buys 1990). Non-breeding birds flock and roost with other granivores; mixed roost in Somalia held up to 1000 Chestnut Weavers, with Red-billed Quelea Quelea quelea, Ru ¨ ppell’s Weaver P. galbula, Lesser Masked Weaver P. intermedius and Vitelline Masked Weaver P. vitellinus (Clarke 1985). Often associated with Quelea quelea at roosts, even when alternative sites available (Ward and Zahavi 1973). Heavy moult in Somalia in Aug (Clarke 1985). In Namibia, moults in June–Nov, YY apparently starting wing-moult before XX. Juvs undergo complete moult, starting later than adults. Anting recorded in captive birds. Food. Seeds, such as grasses Panicum maximum, P. novemnerve, P. stapfianum; nectar of Aloe arborescens, A. candelabrum, A. ferox, A. marlothii. In Kenyan agricultural land, wild grass seeds made up >70% of diet in Oct–Dec, then birds switched to cereal crops in Feb–Apr, with sorghum a major component; insects composed 100 trees had 40–100 nests in each. May nest in association with Village Weaver P. cucullatus, P. intermedius, Red-billed Buffalo-Weaver Bubalornis niger and Whiteheaded Buffalo-Weaver Dinemellia dinemelli (Somalia), P. intermedius (Kenya), Speke’s Weaver P. spekei, P. vitellinus and occasionally Black-necked Weaver P. nigricollis, also with Chestnut Sparrow and Red-billed Quelea. Near Nairobi, Kenya, extensive colonies form after good rains in some years but species is totally absent in others (Cunningham-van Someren 1980). However, in Namibia, nests annually at same sites, even in dry years (Komen and Buys 1990). Monogamous. Breeding

apparently well-synchronized within a colony; YY are present during nest-building phase, displaying with beating wings while hanging below nests, but quit colony once XX are incubating. YY form nomadic flocks, leaving colony occupied only by XX and young birds. NEST: retort-shaped (A) with short spout c. 60 long (Namibia) or without spout (Kenya: Collias and Collias 1964). Usually suspended by strongly-woven grass stems from tip of branch; sometimes 3–4 nests in a string below one another or clustered (B). Interior tightly woven, but exterior ragged, with protruding grass stalks; ceiling and floor lined with grass heads, stems of which may protrude. Leaves not stripped from surrounding branches, but incorporated into nest structure. Nest built by Y, using grasses Enneapogon cenchroides, Cenchrus ciliaris, Eragrostris spp. and Stipagrostris spp. (Namibia), or Eriochloa rubica, Bothriochloa radicans and Chloris virgata with some leaves of acacia tree in which nest sited (Kenya). X continues to add lining of grass seed heads during incubation, and a thick lining results; in Namibia, X invariably used Stipagrostris uniplumis inflorescences for lining, in E Africa often Chloris virgata. Nest size (n ¼ 26, Namibia) 150–210 (160) long by 120–180 (150) high. Favours scattered large trees in open veld, avoiding densely bushed areas; in Namibia, most nests are in Acacia, Albizia or Colophospermum mopane trees at 3–5 m; in Kenya also in large baobabs. EGGS: 1–6, (29, n ¼ 136, Namibia); clutches of 5–6 perhaps produced by more than 1 X. Pale turquoise blue, sometimes speckled or scrolled with dark green. SIZE: (n ¼ 20, Namibia) 215–245  145–168 (228  157). LAYING DATES: Ethiopia, Oct (possibly June–July, breeding condition Sept); Somalia, May (breeding condition Aug), Uganda, May–July; Kenya, Apr–July, Nov in

Ploceus badius

179

INCUBATION: by X only, starting apparently with first egg, so that hatching period of chicks is staggered. Period: 11–14 days. DEVELOPMENT AND CARE OF YOUNG: Young fed by X only (Y no longer present at colony). Chicks began chirping at 3 days, eyes open at 6 days, when first quill feathers appear. Nestling period: 13–16 days. BREEDING SUCCESS/SURVIVAL: hatching success at a colony in Namibia 52% (172 eggs); Cape Crow Corvus capensis ate some eggs and chicks. In Namibia, within a few days of a breeding colony being discovered by Red-headed Finch Amadina erythrocephala, c. 25–30% of XX desert, abandoning incomplete and complete clutches. The finches immediately take over deserted nests and start laying, sometimes adding to weaver clutches. At one colony, 11 nests were appropriated by Red-headed Finches; colony then contained 170 weaver eggs and 64 finch eggs; 5 of these 11 contained 1–3 weaver eggs and 3–5 finch eggs. All weavers that hatched in appropriated nests died within 3 days (Archibald 1987). In Kenya, Chestnut Sparrows, Cutthroat Finches Amadina fasciata and Silverbills Euodice cantans all used abandoned Chestnut Weaver nests for breeding (van Someren and van Someren 1945). Y ringed in Kenya, recovered 5 months later, 213 km from ringing site (Backhurst 1977).

N desert region; Tanzania, Mar–Apr; Angola, Apr; Namibia, Dec–May, primarily Jan–March.

Key References Berry, H. H. et al. (1988), Braine, S. G. and Braine, J. W. S. (1971), Komen, J. (1990), Komen, J. and Buys, P. J. (1990).

Ploceus badius (Cassin). Cinnamon Weaver. Tisserin cannelle.

Plate 11

Hyphantornis badius Cassin, 1850. Proc. Acad. Nat. Sci. Philadelphia, 5, p. 57; Fazogloa, Eastern Africa [presumed to be Fazogli, Sudan].

(Opp. p. 154)

Forms a superspecies with P. rubiginosus. Range and Status. Endemic resident, Sudan. Nile R. and its tributaries, north to 16
N, south to Uganda border. Common to locally abundant. Description. ADULT Y: forehead and crown black; nape, mantle and rump dark chestnut-brown; tail olive-green; chin, lores, cheeks and ear-coverts black; throat black extending onto breast as a bib; breast dark chestnut-brown around bib, becoming lighter towards belly; flanks chestnut-brown; belly golden yellow in centre, streaked with chestnut; thighs and undertail-coverts yellow; wings dark brown with narrow yellow outer margins and broad yellow inner webs to remiges, broad yellow (or sometimes chestnut-brown) margins to coverts; underwing yellow. Bill black; eyes brown; legs brown. YY vary in amount of yellow among the chestnut on upperparts and underparts. ADULT Y (non-breeding): forehead, crown and nape olive-green with no streaking; mantle buffy brown with broad, dark central streaks; rump plain buffy brown; tail and wings as in breeding Y, but no chestnut margins to coverts; lores black; cheeks and ear-coverts buffy brown; chin and throat white; breast, flanks and belly buffy brown except for white centre to belly; thighs and undertailcoverts white. Upper mandible black, lower mandible horn; eyes brown; legs brown. ADULT X: like non-breeding Y, but

Ploceus badius

Ploceus badius

179

INCUBATION: by X only, starting apparently with first egg, so that hatching period of chicks is staggered. Period: 11–14 days. DEVELOPMENT AND CARE OF YOUNG: Young fed by X only (Y no longer present at colony). Chicks began chirping at 3 days, eyes open at 6 days, when first quill feathers appear. Nestling period: 13–16 days. BREEDING SUCCESS/SURVIVAL: hatching success at a colony in Namibia 52% (172 eggs); Cape Crow Corvus capensis ate some eggs and chicks. In Namibia, within a few days of a breeding colony being discovered by Red-headed Finch Amadina erythrocephala, c. 25–30% of XX desert, abandoning incomplete and complete clutches. The finches immediately take over deserted nests and start laying, sometimes adding to weaver clutches. At one colony, 11 nests were appropriated by Red-headed Finches; colony then contained 170 weaver eggs and 64 finch eggs; 5 of these 11 contained 1–3 weaver eggs and 3–5 finch eggs. All weavers that hatched in appropriated nests died within 3 days (Archibald 1987). In Kenya, Chestnut Sparrows, Cutthroat Finches Amadina fasciata and Silverbills Euodice cantans all used abandoned Chestnut Weaver nests for breeding (van Someren and van Someren 1945). Y ringed in Kenya, recovered 5 months later, 213 km from ringing site (Backhurst 1977).

N desert region; Tanzania, Mar–Apr; Angola, Apr; Namibia, Dec–May, primarily Jan–March.

Key References Berry, H. H. et al. (1988), Braine, S. G. and Braine, J. W. S. (1971), Komen, J. (1990), Komen, J. and Buys, P. J. (1990).

Ploceus badius (Cassin). Cinnamon Weaver. Tisserin cannelle.

Plate 11

Hyphantornis badius Cassin, 1850. Proc. Acad. Nat. Sci. Philadelphia, 5, p. 57; Fazogloa, Eastern Africa [presumed to be Fazogli, Sudan].

(Opp. p. 154)

Forms a superspecies with P. rubiginosus. Range and Status. Endemic resident, Sudan. Nile R. and its tributaries, north to 16
N, south to Uganda border. Common to locally abundant. Description. ADULT Y: forehead and crown black; nape, mantle and rump dark chestnut-brown; tail olive-green; chin, lores, cheeks and ear-coverts black; throat black extending onto breast as a bib; breast dark chestnut-brown around bib, becoming lighter towards belly; flanks chestnut-brown; belly golden yellow in centre, streaked with chestnut; thighs and undertail-coverts yellow; wings dark brown with narrow yellow outer margins and broad yellow inner webs to remiges, broad yellow (or sometimes chestnut-brown) margins to coverts; underwing yellow. Bill black; eyes brown; legs brown. YY vary in amount of yellow among the chestnut on upperparts and underparts. ADULT Y (non-breeding): forehead, crown and nape olive-green with no streaking; mantle buffy brown with broad, dark central streaks; rump plain buffy brown; tail and wings as in breeding Y, but no chestnut margins to coverts; lores black; cheeks and ear-coverts buffy brown; chin and throat white; breast, flanks and belly buffy brown except for white centre to belly; thighs and undertailcoverts white. Upper mandible black, lower mandible horn; eyes brown; legs brown. ADULT X: like non-breeding Y, but

Ploceus badius

180

PLOCEIDAE

underwing cream, not yellow. SIZE (12 YY, 5 XX): wing, Y 70–77 (731), X 63–69 (660); tail, Y 44–50 (468), X 39–43 (408); bill, Y 161–196 (179), X 167–180 (174); tarsus, Y 190–208 (202), X 167–185 (177). IMMATURE: resembles X, but crown and mantle paler; rump greener; chin and throat washed with yellow; underparts otherwise dull white. Bill light horn brown. NESTLING: undescribed. TAXONOMIC NOTE: some birds (‘axillaris’) have black of crown extending onto nape, and are darker chestnut-brown on underparts. There appears to be considerable individual variation in these characters, and we treat the species as monotypic, pending further study.

Field Characters. Length 14 cm. Breeding Y with black head and chestnut body resembles Y of allopatric Chestnut Weaver P. rubiginosus but has yellow belly, undertailcoverts and wing edgings, olive-green (not brown) tail; X and non-breeding Y streaky brown, told by olive-green head with prominent yellow supercilium, yellow in wings and tail, whiter underparts with buffy wash on breast and flanks.

Food. Seeds; no details recorded.

Breeding Habits. Colonial, possibly monogamous. Y displays on grass stalks and at nests. NEST: roughly globular, closely woven from grass blades and strips torn from sorghum leaves, with deep cup, dome and porch entrance at one side near top, between supporting stems. Nest located in dense stands of sorghum grass in flooded areas. Slung between 2 or 3 grass stems, 16–21 m above ground. First stage in construction is a bridge between 2 grass stems. Nests 5000 birds (Parker 1999); Swaziland c. 3000 (Parker 1994). Description. A. a. melanota (Heuglin): Ethiopia, S Sudan, Uganda, W Kenya, NE Zaı¨re, NW Tanzania. ADULT Y: forehead white; crown to hindneck, sides of neck and upper mantle dark rusty brown, merging with sooty black lower mantle to scapulars and uppertail-coverts. Tail sooty black. Lores black; cheeks and ear-coverts dark rusty brown, streaked blackish. Chin to upper breast dark brown with diffuse blackish streaks; lower breast, flanks and belly sooty grey with narrow blackish streaks; undertail-coverts blackish, tips fringed white. When fresh, feathers of mantle and scapulars finely fringed with buff, those of breast, flanks and belly fringed pale grey-buff. Upperwing blackish; bases of primaries white on both webs, forming patch on closed wing extending 10–12 mm beyond primary coverts; white bases of secondaries and tertials more restricted, hidden

beneath greater coverts; when fresh, secondaries finely edged and tertials edged and tipped buff, greater and median coverts finely fringed buff. Underwing-coverts and axillaries sooty black. Bill black, very stout; eyes brown or reddish brown; legs dark slate. ADULT X: forehead to lores dark brown, crown and rest of upperparts tawny brown, with diffuse darker brown mottling. Tail dark brown. Narrow, poorly-marked grey-buff superciliary stripe; cheeks and ear-coverts buffy brown with darker brown streaks. Underparts, including lower side of neck, buffy white, chin to upper breast and flanks with bold dark brown streaks, lower breast and belly with narrower streaks, undertail-coverts with blackish centres. Upperwing dark brown; edges of primaries and broader edges of secondaries and tertials tawny; broad edges and tips of greater coverts and broad tips of median and lesser coverts tawny; alula finely edged tawny-buff; feathers at bend of wing broadly edged buffy white. Underwing-coverts and axillaries buffy white. Bill yellowish with black culmen, cutting edges and tip to lower mandible; eyes and legs as ad. Y. SIZE (10 YY, 10 XX): wing, Y 95–100 (967), X 85–96 (885); tail, Y 62–68 (653), X 55–65 (607); bill, Y 22–25 (237), X 20–24 (218); tarsus, Y 22–24 (232), X 20–23 (215). WEIGHT: W Kenya, Y (n ¼ 7) 33– 455 (385), X (n ¼ 3) 28–44 (363). IMMATURE: juvenile like ad. X, but bill yellow with horn-brown culmen; upperparts and upperwing-coverts more broadly fringed tawny-buff; buffier below. See also imm. A. a. albifrons (below). NESTLING: newly hatched chick naked with some traces of white down. Fledged chicks resemble X, with all yellow bill, very short tail. YY heavier than XX (Laycock 1984b). A. a. capitalba (Bonaparte): Senegal to SW Nigeria. Y has head to upper breast and upper mantle light rufous-brown; back greyer (less black) and underparts much paler and greyer than in melanota. X somewhat paler above than in melanota, more finely streaked below. Smaller than melanota: wing, Y (n ¼ 7) 82–88, X (n ¼ 3) 78–82 (797); tail, Y (n ¼ 7) 54–57, X (n ¼ 3) 48–50 (493); bill, Y (n ¼ 7) 20–22, X (n ¼ 3) 20–20 (20); tarsus, Y (n ¼ 7) 20– 22. WEIGHT: (Liberia) Y (n ¼ 11) (362), X (n ¼ 3) 286–312 (300). A. a. saturata Sharpe: SE Nigeria, Cameroon, N Gabon, N Congo, SW Central African Republic, NW Zaı¨re. Y has rich rufous-brown head, underparts lighter and greyer than in melanota; generally less pale than capitalba, and slightly larger: wing, Y (n ¼ 6) 89–96, X (n ¼ 7) 82–85; tail, Y (n ¼ 6) 58–67, X (n ¼ 7) 55–57; bill, Y (n ¼ 6) 20–22, X (n ¼ 7) 20–21; tarsus, Y (n ¼ 6) 19–22, X (n ¼ 7) 19–22. A. a. tandae Bannerman: NW Angola, extreme W Zaı¨re. Y differs from other races in having a broad white band on forehead; head, neck, upper mantle and upper chest chestnutbrown; breast to undertail-coverts grey with narrow black streaks and buff tips to feathers. X paler than nominate race. SIZE: wing, Y 90–92, X 80; tail X 56. WEIGHT: X (n ¼ 1) 42. A. a. montana van Someren: central and SW Kenya, central and W Tanzania, Rwanda, Burundi, E Zaı¨re, Zambia (except Barotseˆ i (except extreme S), extreme NW Zimbabwe. Y land), Malaw differs from melanota in having head to breast blackish, almost concolorous with back. Wing, Y (n ¼ 9) 92–96 (936), X 84–90. WEIGHT: Y 37–437, X 36. A. a. unicolor (Fischer and Reichenow): S Somalia, coastal ˆ i. Similar to Kenya, NE and E Tanzania, extreme S Malaw montana, but head of Y tinged brown. Smaller: wing, Y (n ¼ 10) 85–91 (888), bill, Y (n ¼ 10) 21–23 (219). WEIGHT: (Nchalo, ˆ i) Y (n ¼ 177) 394–580 (477), X (n ¼ 95) 299–434 (373), Malaw imm. Y (n ¼ 223) 401–552 (458), imm. X (n ¼ 149) 305–419 (351) (Hanmer 2002b). A. a. kasaica Schouteden: Kasai (S Zaı¨re). Still blacker than montana; no contrast between head and back. A. a. maxima Roberts: SE Angola, Caprivi, Zambia (Barotseland), N Botswana, extreme NW Zimbabwe. Like montana but larger: wing Y 104–107, X 93–98. A. a. albifrons (Vigors): Transvaal (this race?), KwaZulu-Natal and Gamtoos R., E Cape Province. Y has head to breast dark grey-brown, back sooty brown; generally slightly browner than

199

200

PLOCEIDAE

unicolor. X paler brown, less tawny, above than unicolor and melanota. Larger than unicolor: SIZE (48 YY, 98 XX: Brown et al. 2001): wing, Y 92–104 (966), X 79–95 (879); tail, Y 67–80 (714), X 51–71 (641); bill, Y 205–240 (225), X 170–225 (196); tarsus, Y (n ¼ ?) 24–26, X (n ¼ ?) 21–23. WEIGHT: Y (n ¼ 48) 346– 571 (513), X (n ¼ 98) 341–530 (406); imm. YY >48, imm. XX 1200 birds ringed in Malaw year, 18 after 1 year, 8 after 2, 2 after 3, 2 after 5 and 2 after 8 years (Hanmer 2002b). Key References Brown, M. et al. (2001), Hanmer, D. B. (2002b), Laycock, H. T. (1979, 1982, 1984a, b).

201

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PLOCEIDAE

Genus Quelea Reichenbach

Small, short-tailed weavers of moist to arid grassland. Short conical bills, which may change colour in breeding season. Sexually dimorphic, with breeding YY developing black or reddish plumage about the head. Nest with side entrance; eggs plain or freckled with brown. Highly gregarious and colonial, nomadic or migratory in most regions; major granivorous bird pests. Non-breeding Y and X closely resemble members of genus Euplectes, and Quelea is probably closely related to them. Endemic, 3 species.

Plate 12

Quelea cardinalis (Hartlaub). Cardinal Quelea. Travailleur cardinal.

(Opp. p. 155)

Hyphantica cardinalis Hartlaub, 1880. J. Orn., 28, p. 325; Lado, Sudan. Range and Status. Endemic; nomadic or migratory. Sudan, local and uncommon in Imatong Mts. Ethiopia, occasional in south. Zaı¨re, Mahagi, Faradje, Kivu. Uganda, throughout non-forested areas. Kenya from S Turkana, Indunumara and Mt Marsabit to Kisumu, Londiani, Ruiru, Nairobi and Ngong, S Nyanza to Narok, Olorgesailie and Amboseli and Tsavo Nat. Park. Tanzania, in West Lake and Kibondo, to Mpanda, Namanyere, Rukwa, Ruaha Nat. Park, Iringa area, plains north of Ulugurus and ˆ i, 2 records, Lilongwe. Zambia, Ngerengere R. Malaw uncommon breeding visitor to Luangwa Valley region, south to lower Zambezi. No old records south of the Zambezi (Vernon 1987), but recent reports from Kazungula, Zimbabwe, and Caprivi region of Namibia (C. Spottiswoode, in Harrison et al. 1997). Locally abundant. Description. ADULT Y (breeding): forehead, crown and part of nape red; some birds lack red on nape and have sharp line between red crown and streaked nape. Lower nape and mantle light brown, feathers with broad dark central streaks; some birds have scattered red-tipped feathers on lower nape and mantle. Rump light brown, less heavily streaked than mantle; tail dark brown; lores dark blackish, chin, throat and cheeks red; breast partly red, but much individual variation in extent of red on underparts; lower breast, belly and undertail-coverts creamy white; flanks tawny with some streaking, thighs tawny; wings dark brown, primaries with narrow yellow outer margins, coverts with pale tawny margins; underwing cream. Bill black; eyes brown; legs brown. ADULT Y (non-breeding): forehead, crown, nape and mantle light brown with dark central streaks; rump less heavily streaked. Tail and wings as in breeding Y. Lores dark brown, cheeks and ear-coverts tawny brown; clear yellow supercilium. Chin and throat pale yellow, extending onto breast. May retain some red feathers on head and throat. Lower breast, belly, undertail-coverts creamy white. Flanks tawny with some streaking, sometimes streaking on sides of breast opposite wings; thighs tawny. Bill dark brown; eyes brown; legs brown. ADULT X: like non-breeding Y. SIZE (12 YY, 11 XX): wing, Y 58–61 (588), X 54–58 (565); tail, Y 30–34 (322), X 27–32 (301); bill, Y 133– 142 (137), X 125–140 (132); tarsus, Y 160–170 (163), X 143– 162 (152). WEIGHT: (Kenya) Y (n ¼ 13) 113–134 (125); X (n ¼ 19) 114–150 (129). IMMATURE: upperparts dark brown with broad buffy margins; prominent buffy yellow supercilium; cheeks buff; underparts white, unstreaked, with broad buff band across breast, buff wash on flanks and thighs. Bill brown; eyes brown; legs brown.

Quelea cardinalis

NESTLING: not described. TAXONOMIC NOTE: ‘pallida’

and ‘rhodesiae’ have been described, but Y colour varies considerably (cf. Chapin 1954, Granvik 1934) and, in a mobile species with poorly-understood movements, clear separation of the populations is unlikely.

Field Characters. Length 10–11 cm. Small, with short stubby bill. Breeding Y separated from similar Red-headed Quelea Q. erythrops by red of head extending onto breast but not onto nape, which is streaked like back (sometimes with reddish wash). X has yellowish face and superciliary stripes; told from X Red-headed Quelea by smaller, darker bill, yellowish throat, but probably not separable in the field. Non-breeding Y like X but with some red on head. Voice. Tape-recorded (99, 104, B, GREG, HOR, McVIC, PAY). 4-part song starts with sharp ‘tsap’ notes which

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PLOCEIDAE

Genus Quelea Reichenbach

Small, short-tailed weavers of moist to arid grassland. Short conical bills, which may change colour in breeding season. Sexually dimorphic, with breeding YY developing black or reddish plumage about the head. Nest with side entrance; eggs plain or freckled with brown. Highly gregarious and colonial, nomadic or migratory in most regions; major granivorous bird pests. Non-breeding Y and X closely resemble members of genus Euplectes, and Quelea is probably closely related to them. Endemic, 3 species.

Plate 12

Quelea cardinalis (Hartlaub). Cardinal Quelea. Travailleur cardinal.

(Opp. p. 155)

Hyphantica cardinalis Hartlaub, 1880. J. Orn., 28, p. 325; Lado, Sudan. Range and Status. Endemic; nomadic or migratory. Sudan, local and uncommon in Imatong Mts. Ethiopia, occasional in south. Zaı¨re, Mahagi, Faradje, Kivu. Uganda, throughout non-forested areas. Kenya from S Turkana, Indunumara and Mt Marsabit to Kisumu, Londiani, Ruiru, Nairobi and Ngong, S Nyanza to Narok, Olorgesailie and Amboseli and Tsavo Nat. Park. Tanzania, in West Lake and Kibondo, to Mpanda, Namanyere, Rukwa, Ruaha Nat. Park, Iringa area, plains north of Ulugurus and ˆ i, 2 records, Lilongwe. Zambia, Ngerengere R. Malaw uncommon breeding visitor to Luangwa Valley region, south to lower Zambezi. No old records south of the Zambezi (Vernon 1987), but recent reports from Kazungula, Zimbabwe, and Caprivi region of Namibia (C. Spottiswoode, in Harrison et al. 1997). Locally abundant. Description. ADULT Y (breeding): forehead, crown and part of nape red; some birds lack red on nape and have sharp line between red crown and streaked nape. Lower nape and mantle light brown, feathers with broad dark central streaks; some birds have scattered red-tipped feathers on lower nape and mantle. Rump light brown, less heavily streaked than mantle; tail dark brown; lores dark blackish, chin, throat and cheeks red; breast partly red, but much individual variation in extent of red on underparts; lower breast, belly and undertail-coverts creamy white; flanks tawny with some streaking, thighs tawny; wings dark brown, primaries with narrow yellow outer margins, coverts with pale tawny margins; underwing cream. Bill black; eyes brown; legs brown. ADULT Y (non-breeding): forehead, crown, nape and mantle light brown with dark central streaks; rump less heavily streaked. Tail and wings as in breeding Y. Lores dark brown, cheeks and ear-coverts tawny brown; clear yellow supercilium. Chin and throat pale yellow, extending onto breast. May retain some red feathers on head and throat. Lower breast, belly, undertail-coverts creamy white. Flanks tawny with some streaking, sometimes streaking on sides of breast opposite wings; thighs tawny. Bill dark brown; eyes brown; legs brown. ADULT X: like non-breeding Y. SIZE (12 YY, 11 XX): wing, Y 58–61 (588), X 54–58 (565); tail, Y 30–34 (322), X 27–32 (301); bill, Y 133– 142 (137), X 125–140 (132); tarsus, Y 160–170 (163), X 143– 162 (152). WEIGHT: (Kenya) Y (n ¼ 13) 113–134 (125); X (n ¼ 19) 114–150 (129). IMMATURE: upperparts dark brown with broad buffy margins; prominent buffy yellow supercilium; cheeks buff; underparts white, unstreaked, with broad buff band across breast, buff wash on flanks and thighs. Bill brown; eyes brown; legs brown.

Quelea cardinalis

NESTLING: not described. TAXONOMIC NOTE: ‘pallida’

and ‘rhodesiae’ have been described, but Y colour varies considerably (cf. Chapin 1954, Granvik 1934) and, in a mobile species with poorly-understood movements, clear separation of the populations is unlikely.

Field Characters. Length 10–11 cm. Small, with short stubby bill. Breeding Y separated from similar Red-headed Quelea Q. erythrops by red of head extending onto breast but not onto nape, which is streaked like back (sometimes with reddish wash). X has yellowish face and superciliary stripes; told from X Red-headed Quelea by smaller, darker bill, yellowish throat, but probably not separable in the field. Non-breeding Y like X but with some red on head. Voice. Tape-recorded (99, 104, B, GREG, HOR, McVIC, PAY). 4-part song starts with sharp ‘tsap’ notes which

Quelea erythrops accelerate into slow and fast buzz, and after pause a long, down-slurred nasal whistle: ‘tsap, tsap, tsap tsaptsaptsaptsap zee-zee-zee-zee-dzidzidzidzidzi . . . nyeeeaaaan’. For other renditions see Zimmerman et al. (1996). General Habits. Inhabits rank grass or tall wooded grassland, generally in dry areas rather than marshes (ecologically intermediate between Red-billed Quelea Q. quelea which inhabits relatively arid country, and Redheaded Quelea which occurs in high rainfall areas). Often appears after rains, especially in normally arid or semi-arid country; up to 2000 m (Imatong Mts, Sudan), but generally below 900 m (Zambia); in Kenya, mainly at 800–2000 m, though occasionally down to 400 m and up to 3000 m. Highly gregarious, in small groups, or large flocks of 600–900 birds; may join flocks of Red-billed Quelea. Initial large flocks break up into smaller groups when nesting begins. Small flocks, including YY in breeding plumage, may appear briefly and disappear again without nesting. Breeding visitor to Zambia, Feb–Apr.; at Faradje, Zaı¨re, present Aug–Nov; Uganda, in Kampala region, present Dec–May. Food. Seeds, particularly Panicum, Setaria. Nectar of Leonotis nepetifolia (Short and Horne 1978); only grass seeds in stomachs (Chapin 1954). Breeding Habits. Polygynous; in dense colonies or small groups of nests; sometimes nest is solitary; colony of several hundred nests may form in area where birds previously unknown. Y displays from elevated perch, perches upright, fluffs out body feathers, partly opens and quivers wings, fans tail jerkily, while sizzling with throat feathers erected and vibrated. When X lands in territory, Y approaches giving short song; performs upright wingbeating display (Crook 1969).

205

NEST: domed structure, tightly woven of grass such as Panicum maximum, 150 high, 100 wide; with large side entrance, without porch, diam. 30, bottom 80 from bottom of nest. Lined with finer grass. Slung between grass stems or stems of Bidens, rarely attached to branches of a shrub; 30–70 cm, or up to 15 m above ground. Sited in tall grass in abandoned cultivation, alongside Yellow and Blackwinged Bishops Euplectes capensis and E. hordeaceus. May construct initial ring, starting with very robust bridge between 2 adjacent grass stalks extended to form a shelf, and then abandons it without completing the nest. Appears that X accepts nest site at an early stage, Y then completes nest; while X incubating, Y may construct another nest and roost in it. Nest completed within a day, with Y working at it throughout. EGGS: 2–4, whitish ground colour with greyish mauve spots and darker scrawls; blue or greenish, even pinkish, finely spotted or mottled with reddish brown. SIZE: (Uganda, n ¼ 7) 170–185  123–130 (173  125); ˆ i, (Tanzania) 165–180  115–130 (Chapin 1954); (Malaw n ¼ 2) 170  120–125. LAYING DATES: Zaı ¨re, Aug–Sept, sometimes to Nov in NE; Uganda, Mar, May–June, Aug (breeding plumage Dec–May, Kampala); Kenya, Mar–July; Tanzania, Feb– ˆ i, Jan. May; Zambia, Feb; Malaw INCUBATION: by X only; period 12–14 days. DEVELOPMENT AND CARE OF YOUNG: young fed by X only, on regurgitated seeds and a few caterpillars. Nestling period: 16–18 days. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysoccocyx caprius and evidently by Paradise Whydah Vidua paradisea (van Someren 1956).

Key References Mansfield, D. M. (1963), van Someren, V. G. L. (1956), Vernon, C. J. (1987).

Quelea erythrops (Hartlaub). Red-headed Quelea. Travailleur `a te ˆte rouge.

Plate 12

Ploceus erythrops Hartlaub, 1848. Rev. Zool. 1848, pl. 109; Sa˜o Tome´.

(Opp. p. 155)

Range and Status. Endemic resident, nomad and migrant. Common to abundant in most parts of range. Senegal, restricted to S. Gambia, mainly near river. Guinea-Bissau. Guinea, Fore´cariah, Kindia, Macenta. Sierra Leone, Bo, Rokupr. Liberia, locally common resident and migrant, from coast to northern mountains. Ivory Coast, W coast at Tabou, Azagny, San Pedro; from Yapo to N, commonest in Korhogo-Ferkessedougou area. Ghana, migrant to coast; forest clearings at Korforidua and Tafo, Accra and Keta Plains; in N in Mole Nat. Park. Togo, resident and mostly intra-African migrant to savanna as at Kara, Mango, Dapaon, Bourgou, Namoundjoga and coastal strip, Lome´, Tokpli, Togokome´. Benin, in N, centre (Be´te´rou) and S (Lama Forest). Nigeria, resident with sporadic movements, mainly south of major rivers, on coast at Lagos, Sapele, Warri and Calabar, north of rivers at Zaria, Jos Plateau, Pandam Wildlife Park. Mali, locally abundant between 12


and 13
N; resident north to 14
300 N; large incursions into sahel zone in wet season (Lamarche 1993). Burkina Faso, in SW near Bobo-Dioulassou, Banfora. Niger, in SW at Gaya, also at Tazza. Chad, in SW. Sudan, uncommon in extreme S of Bahr el Ghazal, Kajo Kaji. Ethiopia, locally abundant in W highlands and in SW. Sa˜o Tome´, resident, mainly in N. Formerly on Prı´ncipe I., but now apparently extinct there. No confirmed records from Bioko. Cameroon, as mapped (M. Languy, pers. comm., 2002). Central African Republic, Lobaye Pre´f., Bamingui-Bangoran Nat. Park, Manovo-Gounda-Saint Floris Nat. Park, Birao, Bangui region. Mbini. Gabon, near coast (Libreville, N’Komi, Tchibanga), also in clearings in forest region at M’Passa. Congo, clearings along river. Zaı¨re, clearings along rivers (Boma, Lukolela, NouvelleAnvers), savannas at Medje, Faradje, Kasenji, also in S from Kasai, Marungu areas. Rwanda. Burundi. Uganda, in W,

Quelea erythrops accelerate into slow and fast buzz, and after pause a long, down-slurred nasal whistle: ‘tsap, tsap, tsap tsaptsaptsaptsap zee-zee-zee-zee-dzidzidzidzidzi . . . nyeeeaaaan’. For other renditions see Zimmerman et al. (1996). General Habits. Inhabits rank grass or tall wooded grassland, generally in dry areas rather than marshes (ecologically intermediate between Red-billed Quelea Q. quelea which inhabits relatively arid country, and Redheaded Quelea which occurs in high rainfall areas). Often appears after rains, especially in normally arid or semi-arid country; up to 2000 m (Imatong Mts, Sudan), but generally below 900 m (Zambia); in Kenya, mainly at 800–2000 m, though occasionally down to 400 m and up to 3000 m. Highly gregarious, in small groups, or large flocks of 600–900 birds; may join flocks of Red-billed Quelea. Initial large flocks break up into smaller groups when nesting begins. Small flocks, including YY in breeding plumage, may appear briefly and disappear again without nesting. Breeding visitor to Zambia, Feb–Apr.; at Faradje, Zaı¨re, present Aug–Nov; Uganda, in Kampala region, present Dec–May. Food. Seeds, particularly Panicum, Setaria. Nectar of Leonotis nepetifolia (Short and Horne 1978); only grass seeds in stomachs (Chapin 1954). Breeding Habits. Polygynous; in dense colonies or small groups of nests; sometimes nest is solitary; colony of several hundred nests may form in area where birds previously unknown. Y displays from elevated perch, perches upright, fluffs out body feathers, partly opens and quivers wings, fans tail jerkily, while sizzling with throat feathers erected and vibrated. When X lands in territory, Y approaches giving short song; performs upright wingbeating display (Crook 1969).

205

NEST: domed structure, tightly woven of grass such as Panicum maximum, 150 high, 100 wide; with large side entrance, without porch, diam. 30, bottom 80 from bottom of nest. Lined with finer grass. Slung between grass stems or stems of Bidens, rarely attached to branches of a shrub; 30–70 cm, or up to 15 m above ground. Sited in tall grass in abandoned cultivation, alongside Yellow and Blackwinged Bishops Euplectes capensis and E. hordeaceus. May construct initial ring, starting with very robust bridge between 2 adjacent grass stalks extended to form a shelf, and then abandons it without completing the nest. Appears that X accepts nest site at an early stage, Y then completes nest; while X incubating, Y may construct another nest and roost in it. Nest completed within a day, with Y working at it throughout. EGGS: 2–4, whitish ground colour with greyish mauve spots and darker scrawls; blue or greenish, even pinkish, finely spotted or mottled with reddish brown. SIZE: (Uganda, n ¼ 7) 170–185  123–130 (173  125); ˆ i, (Tanzania) 165–180  115–130 (Chapin 1954); (Malaw n ¼ 2) 170  120–125. LAYING DATES: Zaı ¨re, Aug–Sept, sometimes to Nov in NE; Uganda, Mar, May–June, Aug (breeding plumage Dec–May, Kampala); Kenya, Mar–July; Tanzania, Feb– ˆ i, Jan. May; Zambia, Feb; Malaw INCUBATION: by X only; period 12–14 days. DEVELOPMENT AND CARE OF YOUNG: young fed by X only, on regurgitated seeds and a few caterpillars. Nestling period: 16–18 days. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysoccocyx caprius and evidently by Paradise Whydah Vidua paradisea (van Someren 1956).

Key References Mansfield, D. M. (1963), van Someren, V. G. L. (1956), Vernon, C. J. (1987).

Quelea erythrops (Hartlaub). Red-headed Quelea. Travailleur `a te ˆte rouge.

Plate 12

Ploceus erythrops Hartlaub, 1848. Rev. Zool. 1848, pl. 109; Sa˜o Tome´.

(Opp. p. 155)

Range and Status. Endemic resident, nomad and migrant. Common to abundant in most parts of range. Senegal, restricted to S. Gambia, mainly near river. Guinea-Bissau. Guinea, Fore´cariah, Kindia, Macenta. Sierra Leone, Bo, Rokupr. Liberia, locally common resident and migrant, from coast to northern mountains. Ivory Coast, W coast at Tabou, Azagny, San Pedro; from Yapo to N, commonest in Korhogo-Ferkessedougou area. Ghana, migrant to coast; forest clearings at Korforidua and Tafo, Accra and Keta Plains; in N in Mole Nat. Park. Togo, resident and mostly intra-African migrant to savanna as at Kara, Mango, Dapaon, Bourgou, Namoundjoga and coastal strip, Lome´, Tokpli, Togokome´. Benin, in N, centre (Be´te´rou) and S (Lama Forest). Nigeria, resident with sporadic movements, mainly south of major rivers, on coast at Lagos, Sapele, Warri and Calabar, north of rivers at Zaria, Jos Plateau, Pandam Wildlife Park. Mali, locally abundant between 12


and 13
N; resident north to 14
300 N; large incursions into sahel zone in wet season (Lamarche 1993). Burkina Faso, in SW near Bobo-Dioulassou, Banfora. Niger, in SW at Gaya, also at Tazza. Chad, in SW. Sudan, uncommon in extreme S of Bahr el Ghazal, Kajo Kaji. Ethiopia, locally abundant in W highlands and in SW. Sa˜o Tome´, resident, mainly in N. Formerly on Prı´ncipe I., but now apparently extinct there. No confirmed records from Bioko. Cameroon, as mapped (M. Languy, pers. comm., 2002). Central African Republic, Lobaye Pre´f., Bamingui-Bangoran Nat. Park, Manovo-Gounda-Saint Floris Nat. Park, Birao, Bangui region. Mbini. Gabon, near coast (Libreville, N’Komi, Tchibanga), also in clearings in forest region at M’Passa. Congo, clearings along river. Zaı¨re, clearings along rivers (Boma, Lukolela, NouvelleAnvers), savannas at Medje, Faradje, Kasenji, also in S from Kasai, Marungu areas. Rwanda. Burundi. Uganda, in W,

206

PLOCEIDAE

Quelea erythrops

?

brown, lower mandible pale flesh-brown; eyes brown; legs brown. ADULT X: like non-breeding Y. SIZE (12 YY, 12 XX): wing, Y 62–67 (642), X 59–63 (609); tail, Y 32–40 (342), X 29–33 (314); bill, Y 151–159 (155), X 142–158 (150); tarsus, Y 173–200 (183), X 164–175 (171). WEIGHT: Y (Kenya, n ¼ 4) 145–191 (174), (South Africa, n ¼ 36) 175–255 (211), (Zaı¨re, n ¼ 19) 16–22 (185); sub-ad. Y (South Africa, n ¼ 66) 155–245 (183); X (Zaı¨re, n ¼ 8) 16–19 (17), (South Africa, n ¼ 63) 150–220 (177); unsexed (Ghana, n ¼ 8) 139–218 (160). IMMATURE: like X but broad buff edges to feathers, buff supercilium, buffy wash on underparts (so only centre of belly and undertail-coverts are white). NESTLING: not described. TAXONOMIC NOTE: races have been proposed, Q. e. erythrops in N and Q. e. viniceps in S (Clancey 1986). Morphological differences are slight, but timing of moult, migrations and breeding in the 2 regions do suggest effective separation with minimal interchange between the 2 populations (Craig 1987). Further study is required; presently we keep the species monotypic.

Field Characters. Length 11–12 cm. Slightly larger than Cardinal Quelea Q. cardinalis. Breeding Y differs in red of head continuing onto nape but not onto breast, black barring on throat. X extremely similar to X Cardinal Quelea, with yellowish face and supercilium, but has white throat, short dark moustachial mark, somewhat larger, paler bill; probably not safely distinguishable in the field. Non-breeding Y has some red on head. east to Kampala area. Kenya, L. Victoria basin; occasional on coast north to Mombasa. Tanzania, Bukoba, coastal lowlands, (some records uncertain because of confusion with Q. cardinalis). Angola, N Huı´la to Cuanza Norte, Malanje and Lunda Sul, south to Tombele R.; in Cabinda; isolated record in Bie´. Zambia, widespread except in ˆ i, uncommon Luangwa and Middle Zambezi valleys. Malaw in lower Shire Valley; from Liwonde north, often common in lake littoral, also L. Chilwa. Mozambique, Ngamwe; Beira, Maputo; local in marshland near coast. Namibia, local in Caprivi region. Zimbabwe, first records, at Kazungula, in 1995. Botswana, vagrants at Mpandamatenga, Chobe, R. Kwando. South Africa, summer breeding migrant to coastal lowlands of KwaZulu-Natal, sporadically as far south as Kingwilliamstown and East London. Description. ADULT Y (breeding): forehead, crown and nape red, ending in sharp line on the nape; lower nape, mantle and rump light brown with dark central streaks on feathers; tail dark brown with narrow yellowish fringes; lores dark blackish, cheeks and ear-coverts red; chin and strip in centre of throat (width of bill) very dark red with blackish barring, remainder of throat red; red sometimes extends onto breast; breast and flanks dull brown, with paler fringes to feathers; belly, thighs and undertail-coverts dull white. Wings brown, narrow greenish yellow outer margins on primaries, coverts with pale buffy margins; underwing cream. Bill black with brown base to lower mandible; eyes brown; legs brown. Once a partial albino (Zambia: Tree 1965a). Y breeding plumage in S Zaı¨re varies considerably (Verheyen 1953). ADULT Y (non-breeding) forehead, crown and nape light brown with dark central streaks, like mantle and rump. Tail and wings as in breeding Y. Lores dark brown, broad yellowish superciliary stripe, cheeks and ear-coverts light brown. Chin white with yellow tinge, throat yellowish, breast buff and unstreaked, flanks dull white with some streaking, belly and undertail-coverts plain dull white; thighs whitish with some buff. Upper mandible

Voice. Tape-recorded (99, 104, B, HOR, McVIC). Chorus from colony a shrill high-pitched chatter with some wheezes and deeper notes, ‘kwaar’ and a crow-like caw; calls include sharp ‘tyap’ or ‘tyip’. General Habits. Inhabits rank grass near water, farmland and rice fields. In Kenya, below 1400 m in higher rainfall areas, in South Africa primarily coastal, 1000. Small groups join Cardinal Queleas, Red-headed Queleas Q. quelea, Yellow-crowned Bishop Euplectes afer, Whitewinged Widowbird E. albonotatus, Red-collared Widowbird E. ardens, Southern Red Bishop E. orix and others. Sometimes sexes flock separately (Verheyen 1953). Flocks restless, some birds feeding on ground, others perching in bushes. When young birds disperse from colonies in Oct in Cameroon they can cause severe damage to rice fields (Da Camara-Smeets 1988). Anting recorded in captive birds (Simmons 1961). In E Africa, occurs from June–Nov north of equator and from Jan–Apr south of it. Kenyan birds are apparently non-breeding migrants, perhaps from both north and south. In W Africa, from May–Dec north of equator and from Jan–Mar south of it. In Gambia and Senegal mainly in July–Sept, but large numbers in Liberia in Jan–July, only small groups remaining in Aug–Dec. Nigerian birds in Oct very fat, ‘like migrants’ (Serle 1957). Apparently a breeding migrant to Mozambique (present Sept–Apr), ˆ i (present Oct–Apr). In South Africa Zambia and Malaw Oct–Apr, peaks in Nov; few winter records (occurrence shows no significant relationship to rainfall).

Quelea quelea Food. Grass seeds, including Echinochloa, Rottboellia exaltata, Scleria verrucosa, Pennisetum americana and rice; also insects. 4 stomachs contained rice, and smaller seeds. Nestlings fed on insects. Breeding Habits. Mating system not established, probably polygynous. Colonial, with several hundred nests close together; largest colonies of 3000–5000 nests. Breeding highly synchronized; YY may leave colony before young fledge. Breeding Y displays from prominent perch. A colony of c. 1000 nests in KwaZulu-Natal, South Africa, started in Dec and was deserted in Feb. Colony seldom at same site in successive years. NEST: tightly-woven dome with side entrance, built from very narrow strips of grass; entrance oblong, without porch, with small platform on lower lip. Not lined. Suspended between 2 grass stems, 05–20 m (most 18 m) up, above water in reeds Phragmites communis. Built by Y only from strips torn from leaves of reeds, collected up to ˆ i, nests built by X only 40 m away; at a colony in Malaw (Benson 1947); in Ghana, Y builds transverse bridge, X builds ring and egg chamber. SIZE (n ¼ 10): 90–100 high, 70–99 wide, entrance diam. 40–45 wide. In tall grasses such as Sorghum lanceolatum or Panicum, slung between stems 16 m above ground, or in tall legumes on inland fringe of mangrove swamps. Grass stems nipped off above active nests. Nests are sometimes usurped by Village Weaver Ploceus cucullatus and Vieillot’s Black Weaver P. nigerrimus, which add material (Nigeria: Din 1994). EGGS: 1–4. Uniform pale blue. SIZE: (n ¼ 53, Cameroon) 174–207  130–143 (189  134), (n ¼ 5, South Africa)

207

186–206  127–137 (193  133). WEIGHT: (n ¼ 27, Ghana) 183. LAYING DATES (in parentheses, YY in breeding plumage): Gambia, July (July–Sept); Guinea (Aug–Nov); Liberia, (May–July, ovarian eggs Sept–Nov); Mali, Aug– Nov; Ghana, May, June, Sept (May, nest-building June); Benin, (July–Aug); Nigeria, June, Aug–Oct (Aug–Sept); Sa˜o Tome´, (Dec–Mar); Cameroon, Sept, Nov (Aug); Gabon, May (Jan, Apr); Sudan, July–Aug; Zaı¨re, July– Aug in N, Jan–Mar in S; Uganda, (Apr); Tanzania, Apr– May; Angola, Apr (large gonads, Nov); Zambia, Jan–Feb, ˆ i, Mar (Dec–May, incomplete nests Apr (Dec–Apr); Malaw Nov); southern Africa, Dec–Jan (Dec–May). INCUBATION: by X only. Period in captivity 12–14 days. Clutches in colony hatch in tightly synchrony. Hatches nearly synchronously with adjoining Red-billed Quelea colony (Chad: Park 1975). DEVELOPMENT AND CARE OF YOUNG: young fed by X only; they are notably silent. Nestling period: 12–14 days. On leaving nest, young congregate in areas of good cover close to colony; fed by both sexes. BREEDING SUCCESS/SURVIVAL: of 122 nests (Ghana), 23 deserted mainly as result of storm damage; hatching success 76% and overall success 71% (Grimes 1977). In KwaZulu-Natal, South Africa, 2 birds recaptured at ringing site after 2 years. Captive Y mated with X Redbilled Quelea; 3 young raised (Siroki 1974). Key References Barre´, N. et al. (1976), Brickell, N. and Konigkramer, T. (1997), Cyrus, D. P. (1986), Grimes, L. G. (1977), McLean, S. and Taylor, R. H. (1986), Tree, A. J. (1965b).

Quelea quelea (Linnaeus). Red-billed Quelea. Travailleur a` bec rouge.

Plate 12

Emberiza quelea Linnaeus, 1758. Syst. Nat., 10th ed. p. 177; ‘India’, ¼ Senegal.

(Opp. p. 155)

Range and Status. Endemic resident; highly nomadic and perhaps truly migratory in some regions. May be the most abundant bird species in the world, forming huge flocks which can devastate cereal crops – Africa’s ‘feathered locust’. Mauritania, formerly abundant along Senegal R. and at temporarily flooded areas such as L. Rkiz, Aftout ech Chergui; now much reduced by massive control operations and concurrent droughts, but still extends well north into sahel as at Tmourt n’Naaj. Senegal, coastal region as at Koalak, Tambacounda; especially abundant in Senegal R. valley. Gambia, uncommon dry season visitor in Oct–May, last bred in 1960. Mali, from 17
N south to 12
N, mainly Niger valley (where breeds), Boucle de Baoule´, Mopti and Sahel zone; numbers much reduced following droughts in mid-1970s. Burkina Faso, in N half, south to Kou, Oursi, rare at Ouagadougou. Ivory Coast and Ghana, irregular dry season visitor to far N (Ferke´sse´dougou; Gambaga, Tono irrigation project). Niger, in SW near Niger R from Ayorou, Tillaberi, Niamey, Gaya and east along S border to BirninKonni, Maradi, Nguigmi and L. Chad. Benin, only in far N. Nigeria, formerly abundant all year between Maiduguri and

L. Chad where breeding is concentrated; less common west of Maiduguri to Sokoto; ranging south to Zaria and Benue R. at times; however, said to have been exterminated throughout N Nigeria in 1975 (Manikowski 1980). Cameroon, mainly north of 9
N, breeding north of 10
N; on Benue Plain and Adamawa Plateau in dry season. Sa˜o Tome´, June–July 1990 record is questionable (Jones and Tye 2001). Chad, throughout S, especially common around L. Chad. Central African Republic, Manovo-Gounda-Saint Floris Nat Park. Sudan, uncommon in Darfur; from Kassala and Khartoum south to Uganda border, mainly in Nile valley. Eritrea, uncommon. Somalia, mostly west of 46
E, south of 5
N is breeding area; in NW, a non-breeding migrant from Ethiopia. Vagrant to Djibouti. Ethiopia, local nomad, breeding throughout lowlands. E Zaı¨re, from L. Albert south to N end of L. Tanganyika. Uganda, widespread in N, south to Entebbe in dry season. Kenya, throughout at times, but marked seasonal movements; in E Africa, 3 breeding populations proposed: S Somalia–E Kenya, S Ethiopia–N Kenya and S Kenya–NE Tanzania (Allan 1983). Rwanda and Burundi, irregular vagrant. Tanzania, commonest on central plateau, largely

Quelea quelea Food. Grass seeds, including Echinochloa, Rottboellia exaltata, Scleria verrucosa, Pennisetum americana and rice; also insects. 4 stomachs contained rice, and smaller seeds. Nestlings fed on insects. Breeding Habits. Mating system not established, probably polygynous. Colonial, with several hundred nests close together; largest colonies of 3000–5000 nests. Breeding highly synchronized; YY may leave colony before young fledge. Breeding Y displays from prominent perch. A colony of c. 1000 nests in KwaZulu-Natal, South Africa, started in Dec and was deserted in Feb. Colony seldom at same site in successive years. NEST: tightly-woven dome with side entrance, built from very narrow strips of grass; entrance oblong, without porch, with small platform on lower lip. Not lined. Suspended between 2 grass stems, 05–20 m (most 18 m) up, above water in reeds Phragmites communis. Built by Y only from strips torn from leaves of reeds, collected up to ˆ i, nests built by X only 40 m away; at a colony in Malaw (Benson 1947); in Ghana, Y builds transverse bridge, X builds ring and egg chamber. SIZE (n ¼ 10): 90–100 high, 70–99 wide, entrance diam. 40–45 wide. In tall grasses such as Sorghum lanceolatum or Panicum, slung between stems 16 m above ground, or in tall legumes on inland fringe of mangrove swamps. Grass stems nipped off above active nests. Nests are sometimes usurped by Village Weaver Ploceus cucullatus and Vieillot’s Black Weaver P. nigerrimus, which add material (Nigeria: Din 1994). EGGS: 1–4. Uniform pale blue. SIZE: (n ¼ 53, Cameroon) 174–207  130–143 (189  134), (n ¼ 5, South Africa)

207

186–206  127–137 (193  133). WEIGHT: (n ¼ 27, Ghana) 183. LAYING DATES (in parentheses, YY in breeding plumage): Gambia, July (July–Sept); Guinea (Aug–Nov); Liberia, (May–July, ovarian eggs Sept–Nov); Mali, Aug– Nov; Ghana, May, June, Sept (May, nest-building June); Benin, (July–Aug); Nigeria, June, Aug–Oct (Aug–Sept); Sa˜o Tome´, (Dec–Mar); Cameroon, Sept, Nov (Aug); Gabon, May (Jan, Apr); Sudan, July–Aug; Zaı¨re, July– Aug in N, Jan–Mar in S; Uganda, (Apr); Tanzania, Apr– May; Angola, Apr (large gonads, Nov); Zambia, Jan–Feb, ˆ i, Mar (Dec–May, incomplete nests Apr (Dec–Apr); Malaw Nov); southern Africa, Dec–Jan (Dec–May). INCUBATION: by X only. Period in captivity 12–14 days. Clutches in colony hatch in tightly synchrony. Hatches nearly synchronously with adjoining Red-billed Quelea colony (Chad: Park 1975). DEVELOPMENT AND CARE OF YOUNG: young fed by X only; they are notably silent. Nestling period: 12–14 days. On leaving nest, young congregate in areas of good cover close to colony; fed by both sexes. BREEDING SUCCESS/SURVIVAL: of 122 nests (Ghana), 23 deserted mainly as result of storm damage; hatching success 76% and overall success 71% (Grimes 1977). In KwaZulu-Natal, South Africa, 2 birds recaptured at ringing site after 2 years. Captive Y mated with X Redbilled Quelea; 3 young raised (Siroki 1974). Key References Barre´, N. et al. (1976), Brickell, N. and Konigkramer, T. (1997), Cyrus, D. P. (1986), Grimes, L. G. (1977), McLean, S. and Taylor, R. H. (1986), Tree, A. J. (1965b).

Quelea quelea (Linnaeus). Red-billed Quelea. Travailleur a` bec rouge.

Plate 12

Emberiza quelea Linnaeus, 1758. Syst. Nat., 10th ed. p. 177; ‘India’, ¼ Senegal.

(Opp. p. 155)

Range and Status. Endemic resident; highly nomadic and perhaps truly migratory in some regions. May be the most abundant bird species in the world, forming huge flocks which can devastate cereal crops – Africa’s ‘feathered locust’. Mauritania, formerly abundant along Senegal R. and at temporarily flooded areas such as L. Rkiz, Aftout ech Chergui; now much reduced by massive control operations and concurrent droughts, but still extends well north into sahel as at Tmourt n’Naaj. Senegal, coastal region as at Koalak, Tambacounda; especially abundant in Senegal R. valley. Gambia, uncommon dry season visitor in Oct–May, last bred in 1960. Mali, from 17
N south to 12
N, mainly Niger valley (where breeds), Boucle de Baoule´, Mopti and Sahel zone; numbers much reduced following droughts in mid-1970s. Burkina Faso, in N half, south to Kou, Oursi, rare at Ouagadougou. Ivory Coast and Ghana, irregular dry season visitor to far N (Ferke´sse´dougou; Gambaga, Tono irrigation project). Niger, in SW near Niger R from Ayorou, Tillaberi, Niamey, Gaya and east along S border to BirninKonni, Maradi, Nguigmi and L. Chad. Benin, only in far N. Nigeria, formerly abundant all year between Maiduguri and

L. Chad where breeding is concentrated; less common west of Maiduguri to Sokoto; ranging south to Zaria and Benue R. at times; however, said to have been exterminated throughout N Nigeria in 1975 (Manikowski 1980). Cameroon, mainly north of 9
N, breeding north of 10
N; on Benue Plain and Adamawa Plateau in dry season. Sa˜o Tome´, June–July 1990 record is questionable (Jones and Tye 2001). Chad, throughout S, especially common around L. Chad. Central African Republic, Manovo-Gounda-Saint Floris Nat Park. Sudan, uncommon in Darfur; from Kassala and Khartoum south to Uganda border, mainly in Nile valley. Eritrea, uncommon. Somalia, mostly west of 46
E, south of 5
N is breeding area; in NW, a non-breeding migrant from Ethiopia. Vagrant to Djibouti. Ethiopia, local nomad, breeding throughout lowlands. E Zaı¨re, from L. Albert south to N end of L. Tanganyika. Uganda, widespread in N, south to Entebbe in dry season. Kenya, throughout at times, but marked seasonal movements; in E Africa, 3 breeding populations proposed: S Somalia–E Kenya, S Ethiopia–N Kenya and S Kenya–NE Tanzania (Allan 1983). Rwanda and Burundi, irregular vagrant. Tanzania, commonest on central plateau, largely

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In South Africa in 2000, up to 21 million birds per month reportedly killed in control operations, and annual kill may exceed 180 million in this country alone. Red-billed Quelea is the most abundant bird species in Kruger Nat. Park, with a population estimate of 225; in Kalahari Transfrontier Park it ranks 6th, with c. of 214 birds (Kemp et al. 2001). The map highlights some areas where breeding occurs and where species may be most abundant; but sporadic breeding can occur anywhere in range, and regionally its abundance fluctuates enormously and unpredictably.

Quelea quelea

?

Red: abundant, breeds. Pink: common, but does not breed regularly. Grey: less common; breeds?

absent from SE and from miombo habitat. Gabon, irregular on southern coastal belt. Congo, occasional in S along the river. W Zaı¨re, coast near Congo R. delta. Angola, from Cabinda, Lunda Sul, NE Moxico, southern Cuanza Norte, Bengo, through Huı´la south to Cunene and Cuando Cubango. SE Zaı¨re, Katanga and Marungu plateau. Zambia, sporadically throughout; breeding colonies only in Luangwa and middle Zambezi valleys and ˆ i, dry season visitor below adjacent plateau areas. Malaw 1100 m; occasional breeder in small numbers in Shire valley. Botswana, common, mainly absent from SW and central Kalahari. Zimbabwe, breeds in river valleys and drier southern lowveld; non-breeding birds can occur anywhere, but vagrant in E Highlands. Mozambique, Zambezi valley, Mt Gorongosa, Beira, Save R., Limpopo R. southwards. Namibia, primarily on central plateau, absent from coastal desert, sporadic in S. South Africa, once SW Cape (1963: Hockey et al. 1989); otherwise absent from macchia habitat and winter rainfall area; commonest in central and northern regions. Abundant, and currently the major pest of planted grain crops in Africa. Density assessed by aerial survey of Chari-Logone population in area of c. 115,000 km2 in N Cameroon and W Chad, 314 birds per km2 (i.e. 36 million birds) in Jan–Mar 1975; in Apr, when rains displaced birds to northern part of region, 1000 birds per km2 in 55,000 km2 (55 million birds). Density of 25,000 nests per ha in this population, and 30,000 nests per ha in NE Nigeria (in all, colonies covered c. 1125 ha, i.e. 31 million nests or 62 million birds before and c. 130 million birds after nesting: Manikowski 1980). FAO Quelea Project control operations are likely to have reduced L. Chad population greatly since 1970s.

Description. Q. q. lathami (Smith) (includes ‘spoliator’: Jones et al. 2002): South Africa. ADULT Y (breeding): forehead, lores, cheeks and ear-coverts black; crown and nape straw-coloured (or sometimes pink); mantle light brown, feathers with heavy central streaks; rump light brown, feathers with much finer central streaks; tail brown; wings brown with dull yellow edges to remiges, buffy edges to coverts; underwing cream; chin and throat black, with straw-coloured (or pink) surround extending to upper breast; lower breast brown, flanks light brown with narrow streaks; belly and undertail-coverts creamy white; thighs buffy white. Bill bright red; eyes brown with narrow, bare red orbital ring; legs orange. Mask (forehead, lores, cheeks, ear-coverts, chin, throat) usually black as described but regionally, variable proportion of YY may have mask: (1) creamy buff, often with pinkish margins; (2) pink to purplish, feathers with cream margin, (3) forehead buff, crown buff or straw-coloured, cheeks and throat creamy white. Moreover, upper breast sometimes pink, and pink may extend onto lower breast, belly, and even as flecks on undertail-coverts. 2 morphs recognized by Ward (1966): YY black-faced or white-faced. Black-faced YY in 7 categories according to amount of black; YY in Sudan and Ethiopia have almost no black on face, nearly all YY in Senegal, Zimbabwe and South Africa are very black-faced, and those in Lake Chad region (Nigeria, Cameroon, Chad) and from Kenya to Zambia are intermediate (Ward 1966). ADULT Y (non-breeding): forehead and crown grey-brown with faint streaks; chin and throat creamy white; breast buffy with faint streaks; otherwise like breeding Y. Bill red to pink; eyes brown with red orbital ring; legs fleshcoloured. ADULT X: like non-breeding Y, but in breeding condition bill and eye-ring dull yellow. An albino known (Zambia, Tree 1965a). SIZE (12 YY, 12 XX): wing, Y 63–69 (660), X 62–67 (642); tail, Y 36–40 (379), X 33–39 (358); bill, Y 142–158 (151), X 143–151 (146); tarsus, Y 168–186 (178), X 168–182 (176). Seasonal changes in bill-length may be related to increased wear when food is picked off hard ground in dry season (Ward 1965a). WEIGHT: Y (Kenya, n ¼ 69) 168–225 (192), (South Africa, n ¼ 1164) 150–265 (196); X (Kenya, n ¼ 30) 150–200 (179), (South Africa, n ¼ 419) 145–250 (187); unsexed (South Africa, n ¼ 163) 147–236 (197) (Skead 1974); juv. (South Africa, n ¼ 643) 140–280 (197). Lays down up to 5 g of fat before migrating, population ‘intermedia’ laying down av. 27 g (Ward and Jones 1977). IMMATURE: like X but head plain grey; cheeks whitish; broad buffy margins to rectrices and wing-coverts. Bill horn brown with paler lower mandible. NESTLING: on hatching naked with small wisps of down on crown and shoulders, bill soft and white. Eyes begin to open on day 4, first feathers emerge from skin. By day 10 covering of contour feathers virtually complete. Q. q. quelea (Linnaeus) (includes ‘intermedia’) Senegal to Chad. Breeding Y has buffy crown, nape and underparts; black facial mask forms broad band on forehead. Q. q. aethiopica (Sundevall): Sudan to Somalia, NE Zaı¨re, Uganda, Kenya and Tanzania. Breeding Y like nominate race, but black frontal band normally absent. TAXONOMIC NOTE: there is no support for separation of southern African birds as Q. q. spoliator (Clancey 1960, 1973),

Quelea quelea on either morphological or distributional grounds (Lourens 1961, Jones et al. 2002). Mask types vary in all regions. Genetic separation of the 3 races (Ward 1966) depends primarily on the proposed migratory patterns for these populations, and their consequent separation when breeding; in all populations males are extremely variable with regard to the characters listed above.

Field Characters. Length 11–12 cm. Breeding Y variable, mask usually black surrounded by varying amounts of pink, but sometimes pink or straw-coloured, and pink may extend onto underparts; told from Ploceus weavers by lack of yellow in plumage. X and non-breeding Y separated from other queleas and from X bishops and widowbirds by large pink bill (sometimes yellow in X); crown and earcoverts grey-brown; pale supercilium and crescent below eye create spectacled effect; narrow pink eye-ring visible at close range. X Straw-tailed and Pin-tailed Whydahs Vidua fischeri and V. macroura have small pink bills, X Pin-tailed a heavily striped head. Brown bill of juv. heavier than most look-alikes. Often in vast flocks which at a distance of several km look like smoke wafting over the horizon. Voice. Tape-recorded (3, 86, 88, 99, 104, B, F, PAY). Sonagrams in Crook (1969). Short 3-part song: preliminary chatter, warbled ‘tweedle toodle tweedle’ and long high whistle; or ‘chee-chee-chee-chee . . . seee-seeew-seee . . . sweeeeee’. Several YY sing in synchronized bursts. Other songs include high thin swizzle, long grating ‘jaaaaaaaaa’ and chattering rapid ‘jijijijijiji . . . ’. Contact call when perched, short ‘chirt chirt’; in aggression, harsh ‘chak’; call given on take-off or in response to another flying overhead ‘tseep tseep’. Special mobbing call in response to snake. Screams in extreme distress. Loud ‘chyzzz’ given by begging juv. and by X during courtship. General Habits. May occur sporadically in very wet and very dry areas, but most abundant in semi-arid habitats; in W Africa concentrated in Sahel zone; in E Africa, commonly in bushed grassland and cultivated areas at 500–1500 m altitude (but occurs up to 3000 m); in southern Africa, preferred breeding habitat is dry thornveld below 1000 m. Occurs commonly in farmland, particularly in cereal fields (see Food). Highly gregarious, feeding and breeding usually in hundreds, often in millions. In flight, flocks are dense and highly synchronised, resembling smoke clouds at a distance. Around water-holes and at roosts, combined weight of birds may break large tree branches. Once nesting is over, huge swarms break up into much smaller flocks of tens or hundreds, which scatter widely, but can continue to congregate nightly in densely-packed roosts. Feeds almost exclusively on grass and cereal seeds, taken equally in situ in fruiting grass heads and from ground. When foraging on ground in large flock, shows spectacular ‘roller feeding’, birds from the back constantly flying to the front, so that flock seems to roll across the savanna. Drinking in shallow water may follow a similar pattern. Appears to drink every day, and huge concentrations may appear at standing water in dry country. Small flocks (60–80 birds) sometimes mix with other granivores, when foraging and at roosts, e.g. with bishops

Euplectes spp., weavers Ploceus spp., Golden Sparrows Passer luteus (on rice fields in Senegal) and often with Red Bishops E. orix (in winter, South Africa; Skead 1965; pers. obs.). Large flocks mix with African Silverbills Euodice cantans, Cut-throat Finches Amadina fasciata and other estrildids (Mali: Paludan 1936). Forms enormous roosts in dry season, of several million birds; roosting sites often the same from year to year. (Roosts, and breeding colonies, may serve as ‘information centres’ which enhance food-finding: Ward and Zahavi 1973). Birds from one roost may forage at different sites and take different spectra of seeds. Sometimes roosts with other granivores, e.g. weavers Ploceus galbula, P. intermedius, P. rubiginosus and P. vitellinus, African Silverbills, and sparrows Passer gongonensis and P. luteus (may roost with the last 2 even when alternative sites are available). Small numbers roosted with Cuckoo-Finches Anomalospiza imberbis (Zimbabwe). Thousands roosted in reeds with hundreds of other ploceids and a million Barn Swallows Hirundo rustica (South Africa: Rudebeck 1955). At hottest time of day, congregates in secondary or daytime roosts, commonly in trees near feeding areas; sometimes with e.g. Red-collared Widowbirds E. ardens (Zambia). May bathe in wet grass. Anting occurs in wild and captive birds (Simmons 1961, Matthews 1987). Locally resident; but large-scale movements occur throughout range, distance and timing varying from year to year. Migrates regularly at start of wet season, 300 km to southwest in NE Nigeria (where migration induced partly by rain, making seeds on ground germinate, becoming unavailable as food), 600–700 km to southeast in N Cameroon and SE Chad, 1200 km to northeast in E Africa, and 550 km to southeast in southern Africa. NE Nigerian and N Cameroon populations, both in L. Chad basin, are separate, former migrating parallel to Komadugu Gana and Kedseram R. valleys, latter along Chari-Logone valley (Ward 1971, Manikowski 1980). Amount of premigratory fattening seems to correlate with distance moved, suggesting that birds normally migrate the full distance without stopping (Ward and Jones 1977); but in E Africa migrants seem to have insufficient fat and probably make several short flights rather than a single non-stop one (Manyanza 1980). Nevertheless, present in all months in E and southern Africa at some localities where food (crops, feedlots) available all year (Whittington-Jones et al. 2001). At Gaborone, Botswana, arrives to breed in Dec–Jan, most birds leaving in Mar, and non-breeding birds appear in May–June (Beesley and Irving 1976). Ringing recoveries: from Ethiopia to ˆ i, Zambia Torit, Sudan, after 2 years; Zimbabwe to Malaw and South Africa; and South Africa to Namibia, Botswana, S ˆ i, Zimbabwe and Mozambique. Of 420 Zaı¨re, Zambia, Malaw birds ringed in South Africa and recovered up to 1981, 10 had moved >1000 km, 113 >100 km and 224 up to 10 km. One moved 2545 km from South Africa to Congo (Bull. Afr. Bird Club 7, 2000, 8). Juvs complete moult within 3 months of fledging (Elliott 1990). In South Africa juv. wing-moult starts later than in ads. In E Cape, wing-moult well synchronized, in Apr– Sept; interrupted moult very rare. Individual takes c. 124 days to moult (significantly longer than in other sympatric ploceids: Craig et al. 2001). In KwaZulu-Natal, ads moult

209

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PLOCEIDAE

in Apr–June and juvs start to moult in June (Berruti 1995). In Kenya, moulting can overlap with breeding or be interrupted; wing moult, suspended early in cycle, resumes later at the same point; if arrested later in cycle, entire cycle starts again with 1st primary (Thompson 1988). In Kenya, YY appear to start post-nuptial moult earlier than XX. In Mali, pre-nuptial moult occurs in July (Paludan 1936); in Senegal post-nuptial moult from Nov to May (Morel and Bourlie`re 1955). Food. Seeds, typically about 2  1 mm in size; also insects and spiders. Damages crops of wheat, sorghum, manna, millet, oats, buckwheat and rice. Seeds of grasses Dactyloctenium aegyptium, Echinochloa colonum, E. pyramidalis, Ischaemum brachyantherum, Oryza bartlii, Panicum laevigatum, Paspalum commersonii, P. dilatatum, P. urvillei, Pennisetum ramosum, Schoenfeldia gracilis, Setaria chevallieri, S. flabellate, S. nigrirostris, S. pallidefusca, S. sphacelata, Urochloa spp. and Pennisetum spp. Appears to prefer cultivated millet to available wild seeds (Manikowski and Da Camara-Smeets 1979), but in most regions major crop damage occurs when wild food sources are inadequate (Ward 1979). In Nigeria, wild grasses made up 75% of diet (Ward 1965a). In Kenyan agricultural lands, 40%; prefers millet to sorghum, and eats only a little maize (Gichuki 1984). At cattle feeding troughs, eats finer parts of cattle feed, especially finely-crushed maize. Invertebrate food includes beetles, caterpillars, butterflies, grasshoppers, crickets, hemipterans, ants, harvester termites, dragonflies and spiders. Termite alates caught and eaten on ground, also caught on the wing when eaten either on ground or at perch (Brooke 1970). X takes snail shells, eggshell fragments and calcrete when laying, apparently to supplement calcium levels (Jones 1976). Gives nestlings fresh seeds, or seeds and insects (mainly grasshoppers and caterpillars), by regurgitation; insects compose 35–50% of nestling food volume (Morel et al. 1957). Colony of c. 12,400 nests with young per ha consumes 1845 kg per ha of seeds and 214 kg per ha of insects in a month (Morel 1968). Breeding Habits. Breeds in vast colonies, with 600 nests practically touching in a small tree 5 m tall and up to 6000 nests in a large tree; a 200 ha site may hold 10 million nests. Occasional colonies much smaller; colony of only 5 pairs can reproduce successfully (Morel et al. 1957). Monogamous, but birds are evidently itinerant breeders, nesting successively at different colonies during one season, so species may in fact be serially polygamous, YY and XX mating with different partners at different sites (some birds have 3 separate moult foci in wings, suggesting 3 breeding episodes in a season: Elliott 1990). Nesting activities closely synchronized; av. time from start of nestbuilding until first egg is 8–9 days (but may be as little as 4 days); entire colony of millions of nests can be established in little more than a week; all birds normally leave area within 41 days from first nesting. Active nests may be abandoned if food or drinking water fails in surrounding area, and entire flock then leaves together; in sahel zone, some breeding birds die of thirst or starvation before

abandoning colony. Availability of fresh grass for nest construction is important; in a colony of 12,400 nest per ha, c. 186 kg per ha of grass was used (Morel 1968). Territorial. Territory confined to immediate vicinity of nest. Y sings in territory (i.e. at or very near nest; he may also sing at roost) and in territorial encounters. YY very aggressive, often fighting in territory, sometimes leading to death. Y threatens with head lowered, tail depressed, body feathers ruffled, and wings sometimes slightly raised, accompanied by aggressive chatter. Y advertises nest as soon as first ring completed, displaying in conspicuous posture, raising and fluttering wings, raising tail, fluffing head and body feathers a little, and singing. He does not sing during nest-advertisement display, but sings when X enters nest or during courtship near nest. When nest nearly completed, Y stops building and pair-formation takes place, both sexes quivering half-spread, drooping wings, fanning tails and fluffing body feathers (Morel et al. 1957). X may give begging call like that of young bird when approaching Y. X solicits copulation with head retracted, tail slightly depressed and wings quivering; after copulation both birds may posture with wings spread widely and slightly vibrated at tips, tail spread and body crouched. Protein reserves of X determine her breeding condition and clutch size; X eats more insects at start of breeding period (Jones and Ward 1976). NEST: small oval ball of grass, with side entrance at top under a small porch (A, B). Surface has distinctive knobbly appearance, with protruding loops and grass ends. Not lined. Nest is started as a bridge of woven grass which becomes a crescent, then an upright ring; roof and side walls added, then floor, finally entrance and porch. Built usually of 600–700 grass strips 200–300 mm long, from e.g.

Quelea quelea Chloris virgata, Digitaria milanjiana, Heteropogon contortus, Panicum maximum and Urochloa mosambicensis. Only green, flexible grass is used. Building completed in 2–3 days (Morel and Morel 1980). SIZE: (Zimbabwe, n ¼ 32) av. 122 high, 96 wide, 104 from front to back, entrance diam. 39; (South Africa) 100–115 high, 90 wide, entrance 50–60; long axis of nest is vertical. Nest sited 1–6 m above ground, often c. 2 m. Birds do not strip leaves around nest, and do not demolish old nests.

Nests built mainly in thornbush with complete grass cover and 20–50% tree cover; in South Africa, mainly thorn trees Acacia erubescens, A. gerrardii, A. karoo, A. mellifera, A. nigrescens, A. nilotica, A. polycantha, A. robusta, A. sieberiana, A. tortilis, Dichrostachys cineracea, Ziziphus mucronota; also in other savanna trees such as Albizia amara, Combretum imberbe, Commiphora schimperi, Cordia sinensis, Dalbergia melanoxylon, and in reeds Phragmites communis. In Senegal, nests mainly in Acacia raddiana, A. senegal and Balanites aegyptiaca, and in flooded areas

sometimes in A. scorpioides, once in bulrushes Typha (Morel and Morel 1974). In Mali, nests in bushes near water or in temporarily flooded areas, and often uses Sesbania thickets. Near L. Chad often nests in reeds and sugarcane Saccharum officinarum. Sometimes builds in scant growth of exotic gum trees in sand dunes (Mauritania, Rift Valley) and poplars (South Africa). Providing fresh green grass to captive birds stimulates immediate nest-building. Persistent attempts to build nests of needle-shaped leaves of ornamental trees failed and birds abandoned the area, leaving tops of branches stripped where they had been building (Lane 1930); nests built of pine needles also abandoned (Oschadleus 1989). In E Cape, South Africa, aborted nests often occur in Red Bishop colonies (Skead 1965). May nest alongside Sudan Golden Sparrows in W Africa (Morel and Morel 1980). 1st egg laid while nest partly finished, and Y continues adding material during incubation. EGGS: 1–5; av. clutch size throughout range 28, except for Chari-Logone population (N Cameroon, W Chad) where 31 (n ¼ 122 clutches) or 335 (n ¼ 1200 clutches, 6 colonies, 1975; Manikowski 1980); av. clutch size in Senegal 272 (n ¼ 127 clutches), 277 (n ¼ 2064), 306 (n ¼ 1428) and 285 (n ¼ 975) in 4 successive years (Morel et al. 1957). Pale greenish or bluish, very rarely spotted with light brown. SIZE: (Senegal, n ¼ 27) 180–210  130–160 (183  143,); (South Africa, n ¼ 86) 161–210  113–148 (188  132). LAYING DATES: Mauritania, Aug–Dec; Gambia, once, Aug 1960; Senegal, Aug–Nov; Mali, July–Oct; Burkina Faso, (breeding plumage, July, Oct); Nigeria, Aug–Nov, Chad, Aug–Sept; Sudan, June, Sept; Ethiopia, Aug–Sept; Somalia, June–July, Dec; Zaı¨re, Sept near L. Albert, Jan– May in S; Angola, Feb–May, also June–July (breeding plumage from Dec); Kenya, Nov–Apr, June–July; Tanzania, Dec–Feb, May–June; Zambia, Feb–Apr; Malawi, (juvs Apr); Mozambique, Nov–Mar; Namibia, Jan–Apr; Botswana, Dec–Apr; Zimbabwe, Jan–Mar, also Nov–Dec, Apr; South Africa, Dec–Apr. INCUBATION: by X at night, Y may spend short periods on nest during day. Starts with penultimate egg. Incubating parent never stays on eggs for long; one X left and returned to eggs 21 times in 41 min. During incubation period, when mate arrives at nest, both parents greet by quivering half-spread wings, held horizontally (not drooped, and body feathers not fluffed as in pair-formation display: Morel et al. 1957). Period 10–12 days (12 days in captivity). DEVELOPMENT AND CARE OF YOUNG: fed by both Y and X, always by regurgitation, even when insects brought to nest. At a Senegal colony, Y av. 47 feeding visits per day, X 60 (Morel and Morel 1973). As soon as food is delivered to young, parent flies to top of nest tree, waits for foraging flock to fly by, and immediately joins it. Parents forage mainly within 3 km of colony, sometimes up to 25 km away. Nestling period: 10–11 days (14 days in captivity). Young emerge from nest 2–3 days before they can fly (at 12–14 days); perch in branches nearby; independent at 21 days. BREEDING SUCCESS/SURVIVAL: breeding often highly successful. In Nigeria, egg fertility rate 97%; 95% of eggs

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212

PLOCEIDAE

hatched and 87% produced flying young, with little predation evident (Ward 1965b). In Senegal av. 22 young fledged per nest, with little impact from predators (Morel and Morel 1973). In 8 samples totalling 1365 nests in Logone-Chari population, L. Chad (NE Nigeria, N Cameroon, W Chad), mainly in 1975, avs of 096–270 chicks per nest just before fledging (overall av. productivity of 185 young per nest: Manikowski 1980); c. 10% of eggs and 11% of young were taken by predators (Thiollay 1978). In Kruger Nat. Park, South Africa, predation rates at 4 colonies were c. 13%, 14%, 35% and 60% of nests (Pienaar 1969). In Mali, only 50% of eggs produced flying young (Thiollay 1975). The vast L. Chad population thought to be singled brooded; other populations may have 2 or more successive broods. Sex ratio of nestlings and fledglings is balanced (Morel and Bourlie`re 1955). Predominance of YY in dry season flocks formerly ascribed to higher X mortality (Ward 1965c). Large monthly samples in Logone-Chari population gave Y:X ratios of 36:64 in Mar and from 45:55 to 49:51 in May and Oct–Feb (Manikowski 1980). However, flocks can segregate by sex, YY and XX foraging separately (Elliott 1980), so true sex ratios remain uncertain. Twice as many YY as XX die overnight (Bruggers et al. 1981). XX may suffer higher mortality while nesting (Thiollay 1978), but likely that food shortage during non-breeding period is major population-regulating factor. Predators are legion. In Logone-Chari breeding colonies, 28 major and 24 minor bird species take clutches and broods or kill ad. queleas; in addition, 6 owl spp., 17 mammal spp., snakes, varanid and agamid lizards forage in the colonies (Thiollay 1978). Of c. 1750 instances of predation, Cattle Egrets Ardeola ibis were responsible for 30%, Little Sparrowhawks Accipiter minullus, Wahlberg’s Eagle Aquila wahlbergi and Black Kites Milvus migrans for 22%, Red-billed Hornbills Tockus erythrorhynchus and 3 spp. of Halcyon kingfishers for 15%; other major avian predators were storks and ibises (4 spp.), vultures (2 spp.), eagles (4 spp.), falcons (5 spp.) and other hawks (10 spp.). Mammals included monkeys, jackals, foxes, honey badgers Mellivora capensis, African civets Civettictis civetta, genets Genetta genetta, mongooses, hyaenas, small cats and warthogs Phacochoerus aethiopicus. Elsewhere tree squirrels, baboons, serval cats F. serval and even lions Panthera leo and leopards P. pardus prey upon quelea colonies and roosts, opportunistically if not regularly. In Somalia, predatory birds included herons and storks (9 spp.), hawks (17 spp.), owls (2 spp.), laniid and malaconotid shrikes (9 spp.) (Becker and Amir 1993). Other predator lists in Brooke et al. (1972), Thiollay (1975), Biggs (2001). In South Africa, c. 1000 Cattle Egrets destroyed colony of 3000 nests in 9 days, then moved to another colony (Van Ee 1973). Reptile predators of eggs are egg-eating snake Dasypeltis scabra and rock leguaan Varanus exanthematicus, and of nestling queleas, rock leguaan and snakes Python sebae, Philothamnus irregularis, Dispholidus typus, Thelotornis kirtlandii, Naja nigricollis, Dendroaspis polylepis (Pienaar 1969) and Rhamphiophis oxyrhynchus. Flying birds taken by many falcons and sparrowhawks including Falco subbuteo, F. chicquera, F. biarmicus, F. peregrinus, Accipiter gabar and A. minullus, and Barn Owls

Tyto alba. Birds struggling in mist-nets attacked by White Stork Ciconia ciconia; birds perched in bushes speared by Black-headed Heron Ardea melanocephala. At a roost in Somalia, 217 birds died apparently of puncture wounds from Citrus spines (Bruggers et al. 1981). Y died in cold spell in Zimbabwe in 1968. Thousands sometimes die of thirst (Morel 1968). Birds sometimes drown while drinking; corpses eaten by Marabou Storks Leptoptilos crumeniferus. Green Heron Butorides striatus caught ads whilst drinking (Stocker 1994), so did pelomedusid turtles (Namibia: Jarvis 1986). Up to 23 Nile crocodiles Crocodylus niloticus, some 3 m long, snapped at birds drinking on the wing and rushed at flocks drinking on shoreline, evidently killing many (Luangwa Valley, Zambia: Attwell 1954). Birds drinking on the wing may alight on water, and are eaten by crocodiles, Marabou Storks and African Fish Eagles Haliaeetes vocifer. At a Zimbabwe colony several km long in reeds with Ficus cupreifolia and Vernonia colorata trees, local people harvested up to 400 50-kg bags of chicks every day (Saunders 1981). Racks of nestlings are smoked and fledglings boiled and eaten in Zambia (Stewart 1959). In W Africa, traditional bird catchers, using nets on poles, harvest thousands of ads during the dry season; they are plucked, fried and sold in markets (Mullie´ 2000). National and international agencies kill millions of queleas annually in several countries, mainly by means of explosives and petrol bombing; formerly by flamethrowing and aerial spraying with e.g. fenthion, very toxic to birds and considered comparatively harmless to other wildlife (but cf. Becker and Amir 1993). Annual mortality 56% (est. from South African ringing data in 1954–1960, a period of intensive control operations; >90% of sample ringed and recovered in Aug–Jan: Jones 1980) or 60% (using Haldane’s method: Craig 1982d). In South Africa 65–134 million birds killed annually in 1954–1960 with huge fluctuations, 3–180 million annually, since then. It implies that control operations there have little impact, merely replacing naturally occurring mortality (Jones 1980). Oldest ringed bird, 7 years 10 months (D. Oschadleus, pers. comm.). Only c. 02% of birds survive to that age (Manikowski 1980); but 13% of 970 juvs survived for nearly 5 years and 26% of 228 juvs survived for 3 years (Morel et al. 1957).

Key References Crook, J. H. (1960), Bruggers, R. J. and Elliott, C. C. H. (1989), Manikowski, S. (1980), Morel, G. (1968), Morel, G. et al. (1957), Mundy, P. J. and Jarvis, M. J. F. (1989), Oschadleus, H. D. (2001), Ward, P. (1965a, b, c, 1966, 1971), Ward, P. and Jones, P. J. (1977).

Brachycope anomala

213

Genus Brachycope Reichenow

A small, very short-tailed weaver with soft plumage, long undertail-coverts, and short P10. Sexually dimorphic, with black face mask and some yellow on head and breast in Y, but both breeding and non-breeding plumage predominantly brown. Inhabits clearings in riverine forest; nest rounded with a side entrance; eggs plain dark grey. Endemic, monotypic. Thought to link Asiatic Ploceus species and bishops Euplectes (Chapin 1917, 1954; Bannerman 1949), but treated by Wolters (1979–1982) as a subgenus of Quelea. Later, Wolters (1975–1982) raised all 3 subgenera of Quelea to generic rank. Moreau (1960) included Brachycope in Euplectes, even though he thought that there was no seasonal change in Y plumage, and suggested that it was close to E. afer. White (1963) also included Brachycope in Euplectes. Hall and Moreau (1970) noted similarities in pattern and proportions to Quelea – black mask, and yellow distributed in similar way to red of Q. cardinalis – and placed Brachycope in that genus. Sibley and Monroe (1990) commented that generic affinities are uncertain, but the differences warrant a separate genus. The nest with its side entrance certainly suggests a link to Quelea and Euplectes, whereas egg colour is highly distinctive. We thus agree with Dowsett and Forbes-Watson (1993) in retaining a monotypic genus for this anomalous species.

Brachycope anomala (Reichenow). Bob-tailed Weaver. Travailleur a` queue courte.

Plate 12

Ploceus anomalus Reichenow, 1887. J. Orn. 35, p. 214; Stanley Falls, Congo.

(Opp. p. 155)

Range and Status. Endemic resident, clearings in riverine forests in Congo Basin. SE Cameroon, old record Moloundou; Kika and Ngongo (M. Languy, pers. comm., 2002). Congo; S Central African Republic; Zaı¨re, at Boma, from Bolobo upstream to Ponthierville on Congo R. and its tributaries: Sanga, Ubangi, Ruki, Itimbiri, Aruwimi-Ituri, Uele, but not Kasai; locally common.

Brachycope anomala

Description. ADULT Y (breeding): forehead and crown yellow, grading into brown on nape; mantle, back, rump, uppertailcoverts and tail dark brown with paler margins to feathers; lores, cheeks and ear-coverts black; chin and throat black, breast yellow; belly and thighs buffy brown becoming whitish around vent; wing dark brown, upperwing-coverts with pale edges; underwing and undertail-coverts buff. Bill black, eyes dark brown, legs brown. ADULT Y (non-breeding): like breeding Y, but head plain brown; chin, throat and breast buffy, bill brown. ADULT X: resembles non-breeding Y, but bill black. SIZE: (28 YY, 11 XX) wing, Y 58–63 (606), X 55–60 (578); tail, Y 28–32 (300), X 27–31 (289); bill, Y 140–154 (148), X 136–147 (140); tarsus, Y 195–223 (212), X 190–205 (200). WEIGHT: Y (n ¼ 3, Zaı¨re) 20, 26, 30 (P. Herroelen, pers. comm.). IMMATURE: like ad. X, but iris dark grey, bill pinkish brown, feet pinkish. NESTLING: unknown.

Field Characters. Length 12–13 cm. A small weaver restricted to vicinity of rivers in central Africa; soft plumage and short tail give impression of immaturity even to ads (Chapin 1954). Breeding Y distinctive, yellow with black mask and throat; X and non-breeding Y told from all queleas and Euplectes spp. by unstreaked head and uniform buff underparts.

General Habits. Inhabits grassy areas close to human habitations, common in villages and clearings along river banks but not found away from rivers; hops on the ground, perches in palms, mango trees. Forages like sparrows, in open areas. Food. Small seeds in 1 stomach.

Voice. Tape-recorded (104, C, KAE P). Call a dry, harsh ‘jit’ repeated 2–7 times; also a few liquid chirping notes.

Breeding Habits. Presumed monogamous, not colonial.

Brachycope anomala

213

Genus Brachycope Reichenow

A small, very short-tailed weaver with soft plumage, long undertail-coverts, and short P10. Sexually dimorphic, with black face mask and some yellow on head and breast in Y, but both breeding and non-breeding plumage predominantly brown. Inhabits clearings in riverine forest; nest rounded with a side entrance; eggs plain dark grey. Endemic, monotypic. Thought to link Asiatic Ploceus species and bishops Euplectes (Chapin 1917, 1954; Bannerman 1949), but treated by Wolters (1979–1982) as a subgenus of Quelea. Later, Wolters (1975–1982) raised all 3 subgenera of Quelea to generic rank. Moreau (1960) included Brachycope in Euplectes, even though he thought that there was no seasonal change in Y plumage, and suggested that it was close to E. afer. White (1963) also included Brachycope in Euplectes. Hall and Moreau (1970) noted similarities in pattern and proportions to Quelea – black mask, and yellow distributed in similar way to red of Q. cardinalis – and placed Brachycope in that genus. Sibley and Monroe (1990) commented that generic affinities are uncertain, but the differences warrant a separate genus. The nest with its side entrance certainly suggests a link to Quelea and Euplectes, whereas egg colour is highly distinctive. We thus agree with Dowsett and Forbes-Watson (1993) in retaining a monotypic genus for this anomalous species.

Brachycope anomala (Reichenow). Bob-tailed Weaver. Travailleur a` queue courte.

Plate 12

Ploceus anomalus Reichenow, 1887. J. Orn. 35, p. 214; Stanley Falls, Congo.

(Opp. p. 155)

Range and Status. Endemic resident, clearings in riverine forests in Congo Basin. SE Cameroon, old record Moloundou; Kika and Ngongo (M. Languy, pers. comm., 2002). Congo; S Central African Republic; Zaı¨re, at Boma, from Bolobo upstream to Ponthierville on Congo R. and its tributaries: Sanga, Ubangi, Ruki, Itimbiri, Aruwimi-Ituri, Uele, but not Kasai; locally common.

Brachycope anomala

Description. ADULT Y (breeding): forehead and crown yellow, grading into brown on nape; mantle, back, rump, uppertailcoverts and tail dark brown with paler margins to feathers; lores, cheeks and ear-coverts black; chin and throat black, breast yellow; belly and thighs buffy brown becoming whitish around vent; wing dark brown, upperwing-coverts with pale edges; underwing and undertail-coverts buff. Bill black, eyes dark brown, legs brown. ADULT Y (non-breeding): like breeding Y, but head plain brown; chin, throat and breast buffy, bill brown. ADULT X: resembles non-breeding Y, but bill black. SIZE: (28 YY, 11 XX) wing, Y 58–63 (606), X 55–60 (578); tail, Y 28–32 (300), X 27–31 (289); bill, Y 140–154 (148), X 136–147 (140); tarsus, Y 195–223 (212), X 190–205 (200). WEIGHT: Y (n ¼ 3, Zaı¨re) 20, 26, 30 (P. Herroelen, pers. comm.). IMMATURE: like ad. X, but iris dark grey, bill pinkish brown, feet pinkish. NESTLING: unknown.

Field Characters. Length 12–13 cm. A small weaver restricted to vicinity of rivers in central Africa; soft plumage and short tail give impression of immaturity even to ads (Chapin 1954). Breeding Y distinctive, yellow with black mask and throat; X and non-breeding Y told from all queleas and Euplectes spp. by unstreaked head and uniform buff underparts.

General Habits. Inhabits grassy areas close to human habitations, common in villages and clearings along river banks but not found away from rivers; hops on the ground, perches in palms, mango trees. Forages like sparrows, in open areas. Food. Small seeds in 1 stomach.

Voice. Tape-recorded (104, C, KAE P). Call a dry, harsh ‘jit’ repeated 2–7 times; also a few liquid chirping notes.

Breeding Habits. Presumed monogamous, not colonial.

214

PLOCEIDAE

NEST: spherical with a lateral entrance, loosely woven of grass strips, 2–5 m above ground in bushes, small trees (e.g. coffee) and oil palms. EGGS: 2–4: 4 fledglings fed by 2 ads, Kika, Cameroon (M. Languy, pers. comm.). Uniform dark grey. SIZE: (n ¼ 4) (171–185  130–136. WEIGHT: 17 (Scho ¨nwetter 1983).

LAYING DATES: Cameroon, Kika (fledglings being fed, Oct); Zaı¨re (Eala), Jan, (Lukolela), Mar, (Avakubi), Apr, June, Sept–Oct (building Oct at Lukolela).

Key References

Chapin, J. P. (1954), Schouteden, H. (1922).

Genus Euplectes Swainson

Small to medium-sized weavers; sexually and seasonally dimorphic; breeding plumage of YY solid black; or black with red, orange or yellow areas; wing feathers black in Y of some species and retained in non-breeding plumage. Some YY moult tail feathers twice a year, and have short to enormously elongated black tails in breeding plumage. XX sparrowy brown with no seasonal change in appearance, smaller than YY. Bill short and conical. Both sexes always have dull, brownish eyes. Territorial and polygynous; YY of Euplectes jacksoni display at a lek. Trend for reduced Y involvement in nest construction, in parallel with complexity of Y courtship display and ornamentation (Craig 1980), so that in E. jacksoni X alone responsible for selection of nest site and nest-building. Nest with a side entrance, without a tunnel. Only X incubates and feeds young. Occur in grassland, all predominantly seed-eaters. Delacour and Edmond-Blanc (1933) united the genera Euplectes (short-tailed species, which do not moult rectrices and have plain eggs, rarely with some fine spotting) and Coliuspasser (species which moult rectrices, usually long-tailed, eggs with heavy and varied markings); this has been followed by most subsequent authors. However, Wolters (1977–1982) split bishops and widowbirds into 3 genera and 8 sub-genera: a monotypic genus for ardens; short-tailed species except capensis in Euplectes and the rest in Coliuspasser. 2 of his sub-genera coincide with the after and orix superspecies. We use the single genus Euplectes since all species share characteristics of pigmentation, nest structure and skull morphology (A. Craig, unpub. data). Sizzling calls with the bill depressed may be common to all species during courtship. Tail-moult in the pre-nuptial moult occurs in both short-tailed capensis and other ‘Coliuspasser’ which, however, vary in tail-length and tail structure (ardens, albonotatus, axillaris, macroura, hartlaubi, jacksoni, psammocromius, progne); seasonal changes in bill colour also occur in both short- and long-tailed species (e.g. ardens), and YY of both groups may have black

Euplectes orix superspecies

Euplectes afer superspecies

1 1

1

3 1 E. franciscanus 2 E. nigroventris 3 E. orix

2

3

1 E. afer 2 E. diadematus

2

214

PLOCEIDAE

NEST: spherical with a lateral entrance, loosely woven of grass strips, 2–5 m above ground in bushes, small trees (e.g. coffee) and oil palms. EGGS: 2–4: 4 fledglings fed by 2 ads, Kika, Cameroon (M. Languy, pers. comm.). Uniform dark grey. SIZE: (n ¼ 4) (171–185  130–136. WEIGHT: 17 (Scho ¨nwetter 1983).

LAYING DATES: Cameroon, Kika (fledglings being fed, Oct); Zaı¨re (Eala), Jan, (Lukolela), Mar, (Avakubi), Apr, June, Sept–Oct (building Oct at Lukolela).

Key References

Chapin, J. P. (1954), Schouteden, H. (1922).

Genus Euplectes Swainson

Small to medium-sized weavers; sexually and seasonally dimorphic; breeding plumage of YY solid black; or black with red, orange or yellow areas; wing feathers black in Y of some species and retained in non-breeding plumage. Some YY moult tail feathers twice a year, and have short to enormously elongated black tails in breeding plumage. XX sparrowy brown with no seasonal change in appearance, smaller than YY. Bill short and conical. Both sexes always have dull, brownish eyes. Territorial and polygynous; YY of Euplectes jacksoni display at a lek. Trend for reduced Y involvement in nest construction, in parallel with complexity of Y courtship display and ornamentation (Craig 1980), so that in E. jacksoni X alone responsible for selection of nest site and nest-building. Nest with a side entrance, without a tunnel. Only X incubates and feeds young. Occur in grassland, all predominantly seed-eaters. Delacour and Edmond-Blanc (1933) united the genera Euplectes (short-tailed species, which do not moult rectrices and have plain eggs, rarely with some fine spotting) and Coliuspasser (species which moult rectrices, usually long-tailed, eggs with heavy and varied markings); this has been followed by most subsequent authors. However, Wolters (1977–1982) split bishops and widowbirds into 3 genera and 8 sub-genera: a monotypic genus for ardens; short-tailed species except capensis in Euplectes and the rest in Coliuspasser. 2 of his sub-genera coincide with the after and orix superspecies. We use the single genus Euplectes since all species share characteristics of pigmentation, nest structure and skull morphology (A. Craig, unpub. data). Sizzling calls with the bill depressed may be common to all species during courtship. Tail-moult in the pre-nuptial moult occurs in both short-tailed capensis and other ‘Coliuspasser’ which, however, vary in tail-length and tail structure (ardens, albonotatus, axillaris, macroura, hartlaubi, jacksoni, psammocromius, progne); seasonal changes in bill colour also occur in both short- and long-tailed species (e.g. ardens), and YY of both groups may have black

Euplectes orix superspecies

Euplectes afer superspecies

1 1

1

3 1 E. franciscanus 2 E. nigroventris 3 E. orix

2

3

1 E. afer 2 E. diadematus

2

Euplectes franciscanus

215

Euplectes hartlaubi superspecies

1

1

1 E. hartlaubi 2 E. psammocromius

1

2

remiges. Centre of evolution is apparently E Africa; 15 species occur in Kenya. We follow Dowsett and Forbes-Watson (1993) in recognizing E. orix and E. franciscanus as separate species, and in separating the highland E. psammocromius from E. hartlaubi. For a revision of the subspecies see Craig (1993a, b). Endemic, 17 species. 10 independent species and 3 superspecies: E. franciscanus/orix/nigroventris; E. afer/diadematus; E. hartlaubi/psammocromius.

Euplectes franciscanus (Isert). Northern Red Bishop. Euplecte franciscain.

Plate 13

Loxia franciscana Isert, 1789. Schrift. Ges. Nat. Fr. Berlin, 19, p. 332; Accra, Ghana.

(Opp. p. 202)

Forms a superspecies with E. orix and E. nigroventris. Range and Status. Endemic resident and partial migrant mainly within breeding range. Escaped birds breed near Los Angeles, California (K. Garrett, pers. comm.) and in Martinique, West Indies (Barre´ and Benito-Espinal 1985). Common to abundant throughout range. Mauritania, north to 18
N, e.g. Massif du Tagant, Tamourt n’Naaj; occasional records on coast in Nouakchott area (possibly escaped cage birds: Gee 1984). Senegal, widespread except forested areas of Casamance, from Djoudj Nat. Park to Ziguinchor, Kolda. Gambia, throughout. Guinea-Bissau. Guinea, locally common. Mali, south of 17
N, abundant in marshes of delta and lakes, along Niger R. and Bani R., at Ban Markala, in savanna of Boucle de Baoule´. Ivory Coast, not usually south of Bouake´ but recorded in Abidjan in 1982–1984 (Demey and Fishpool 1991). Ghana, resident N and E of forest, Volta region, north to Mole, Tumu, Bolgatanga; local on coast as at Cape Coast, Accra and Keta plains. Togo, abundant resident throughout. Benin, widespread throughout. Nigeria, around Ilorin and Enugu northward to Sokoto and L.

Chad; old records from Lagos, Ibadan, Abeokuta, Awgu and Afkipo. Burkina Faso, widespread and locally common, from Gaoua and Arli north to Ouahigouya. Niger, in SW near Niger R., east to Dallol Bosso, Maradi, Tanout, Zinder. Chad, south of L. Chad. Cameroon, in N and Benue plain, south to Ngounde´re´. Central African Republic, Bangui, Manovo-Gounda-Saint Floris Nat. Park. Sudan, widespread south of Dongola, primarily on Nile and its tributaries, north to 20
N. Ethiopia, locally common to abundant in lowlands. Eritrea. Somalia, in Shabeelle and Jubba valleys, adjoining coastal areas, and in extreme NW. Zaı¨re, extreme NE, Shari R., shores of L. Albert. Uganda, West Nile and Kidepo Nat. Park south to Toro, Lango and Teso. Kenya, in Rift Valley from Elgeya and Eldama Ravine to N Uaso Nyiro R., L. Baringo and L. Bogoria, 1978 sight record near Marsabit. Description. ADULT Y (breeding): forehead and crown black, nape red; mantle red to red-brown; rump and uppertail-coverts red, almost covering brown tail; lores, cheeks and ear-coverts black; chin, throat and breast red; belly and flanks black; thighs

Euplectes franciscanus

215

Euplectes hartlaubi superspecies

1

1

1 E. hartlaubi 2 E. psammocromius

1

2

remiges. Centre of evolution is apparently E Africa; 15 species occur in Kenya. We follow Dowsett and Forbes-Watson (1993) in recognizing E. orix and E. franciscanus as separate species, and in separating the highland E. psammocromius from E. hartlaubi. For a revision of the subspecies see Craig (1993a, b). Endemic, 17 species. 10 independent species and 3 superspecies: E. franciscanus/orix/nigroventris; E. afer/diadematus; E. hartlaubi/psammocromius.

Euplectes franciscanus (Isert). Northern Red Bishop. Euplecte franciscain.

Plate 13

Loxia franciscana Isert, 1789. Schrift. Ges. Nat. Fr. Berlin, 19, p. 332; Accra, Ghana.

(Opp. p. 202)

Forms a superspecies with E. orix and E. nigroventris. Range and Status. Endemic resident and partial migrant mainly within breeding range. Escaped birds breed near Los Angeles, California (K. Garrett, pers. comm.) and in Martinique, West Indies (Barre´ and Benito-Espinal 1985). Common to abundant throughout range. Mauritania, north to 18
N, e.g. Massif du Tagant, Tamourt n’Naaj; occasional records on coast in Nouakchott area (possibly escaped cage birds: Gee 1984). Senegal, widespread except forested areas of Casamance, from Djoudj Nat. Park to Ziguinchor, Kolda. Gambia, throughout. Guinea-Bissau. Guinea, locally common. Mali, south of 17
N, abundant in marshes of delta and lakes, along Niger R. and Bani R., at Ban Markala, in savanna of Boucle de Baoule´. Ivory Coast, not usually south of Bouake´ but recorded in Abidjan in 1982–1984 (Demey and Fishpool 1991). Ghana, resident N and E of forest, Volta region, north to Mole, Tumu, Bolgatanga; local on coast as at Cape Coast, Accra and Keta plains. Togo, abundant resident throughout. Benin, widespread throughout. Nigeria, around Ilorin and Enugu northward to Sokoto and L.

Chad; old records from Lagos, Ibadan, Abeokuta, Awgu and Afkipo. Burkina Faso, widespread and locally common, from Gaoua and Arli north to Ouahigouya. Niger, in SW near Niger R., east to Dallol Bosso, Maradi, Tanout, Zinder. Chad, south of L. Chad. Cameroon, in N and Benue plain, south to Ngounde´re´. Central African Republic, Bangui, Manovo-Gounda-Saint Floris Nat. Park. Sudan, widespread south of Dongola, primarily on Nile and its tributaries, north to 20
N. Ethiopia, locally common to abundant in lowlands. Eritrea. Somalia, in Shabeelle and Jubba valleys, adjoining coastal areas, and in extreme NW. Zaı¨re, extreme NE, Shari R., shores of L. Albert. Uganda, West Nile and Kidepo Nat. Park south to Toro, Lango and Teso. Kenya, in Rift Valley from Elgeya and Eldama Ravine to N Uaso Nyiro R., L. Baringo and L. Bogoria, 1978 sight record near Marsabit. Description. ADULT Y (breeding): forehead and crown black, nape red; mantle red to red-brown; rump and uppertail-coverts red, almost covering brown tail; lores, cheeks and ear-coverts black; chin, throat and breast red; belly and flanks black; thighs

216

PLOCEIDAE

Euplectes franciscanus

Migrations: see text.

brown; undertail-coverts red, almost as long as tail; wings brown with paler margins to feathers; underwing buff. Bill black; eyes dark brown; legs pale brown to flesh-coloured. ADULT Y (nonbreeding): forehead to uppertail-coverts brown, feathers with dark central streaks, tail brown with paler edges to feathers; yellowish superciliary stripe; lores, cheeks and ear-coverts brown to buff; chin and throat white; breast buffy with faint streaking in some individuals; belly and undertail-coverts white, wings brown. Bill brown; eyes brown; legs flesh-coloured. Y does not moult into breeding plumage until 2nd year (Lynes 1924–1925). ADULT X: like non-breeding Y. SIZE (94 YY, 65 XX): wing, Y 57– 66 (629), X 53–60 (562); tail, Y 29–40 (365), X 30–38 (329); bill, Y 129–155 (142), X 128–145 (138); tarsus, Y 183–213 (197), X 170–197 (183). WEIGHT: (Ghana) Y (n ¼ 30), July–Aug) 143–189 (170), (n ¼ 10, July–Sept) 15–18 (160); unsexed (n ¼ 42) 114–212 (150) (Davidson 1978); (Nigeria) Y (n ¼ 14, Apr–July) 135–178 (160), unsexed (n ¼ 131) 115–220 (160) (Fry 1971, Aiyedun 1972). IMMATURE: like X, but retains broad buff edges to feathers of the upperparts until post-juvenile moult. NESTLING: not described. TAXONOMIC NOTE: YY in breeding plumage variable; from Sudan eastwards they tend to have longer tail-coverts, less red on breast, orange-red rather than deep red plumage (‘pusillus’).

Field Characters. Length 11 cm. Black and red breeding Y extremely similar to allopatric Southern Red Bishop E. orix, but black extends further back on crown, tail coverts longer, concealing tail. Smaller than Black-winged Bishop E. hordeaceus, with small bill, brown wings and short brown tail. X and non-breeding Y indistinguishable from Southern Red Bishop, with streaked buff breast-band, short whitish supercilium; told from Black-winged Bishop by paler underparts, in flight by brown wings; less streaked below than Yellow-crowned Bishop E. afer, supercilium short and white.

Voice. Tape-recorded (104, B, BRU, GREG, PAY). Song a mixture of hard chattery notes, buzzy ‘zee-zee-zee . . . ’ and low grating swizzles. Sibilant contact calls: sharp ‘tsip’ and high thin ‘tseet’. Sonagrams of calls and songs of captive birds in Craig (1976). General Habits. Inhabits tall open or bushed grassland, at 600–1800 m in E Africa and up to 2000 m in Ethiopia; nests in seasonally flooded grassland. Closely associated with giant grasses and tall crops such as millet and sorghum, but occurs in open habitats with rank weedy vegetation; in Mali with Andropogon gayamis. Chiefly in ricefields rather than natural savanna in Ivory Coast. Moves by hopping on open ground. Forms large flocks in non-breeding season; with Black-winged Bishops (Uganda) and mixed flocks of seed-eaters included canaries, waxbills and whydahs (Nigeria). Flocks damage crops, and in Ethiopia flapping pieces of cloth are strung over fields to keep them away. Roosts in thickets or tall grass. In N-central Nigeria, wing moult begins in Oct, ends late Feb, lasting 4–5 months in population, but is up to 7 weeks out of synchrony in individual birds, some of which are resident, others migrant. YY acquire breeding plumage by partial moult in May–June, taking 6–7 weeks (Fry 1971). In Ethiopia pre-nuptial moult of YY in Aug–Sept; by Dec post-nuptial moult completed in many birds. Resident over most of range, but in far N a breeding rainy season visitor; evidence of substantial migrations within residential range (at least to west of 20
E). In Mauritania, moves north up to 17
300 N (and beyond in Tagant area) at start of rains (Lamarche 1993). In Dec–Jan all Senegal birds are in non-breeding plumage and do not occur on lower reaches of Senegal R. where known to breed (Schifter 1986). In Mali, large movements take place with arrival of rains, birds moving north to breed in sahel zone; withdrawal to S in Oct (Lamarche 1993). Present in sahel and sudan zones in NE Nigeria throughout dry season, then moves south at start of rains, and absent for a month or more; birds have fat reserves sufficient for flight of 600 km (Jones and Ward 1977). Elsewhere in Nigeria, may be only a non-breeding dry season visitor south of Niger R. and Benue R., and inferred to be on passage (as well as breeding near Zaria, 11
300 N, in early rains (May–June: Fry 1971). Appears in Ethiopia in June–July, before acquiring breeding plumage; largely nomadic in Dec after breeding, and apparently absent from many breeding areas throughout Jan–May (Reichenow 1902–1905). Food. Seeds, also insects. Few field data; common name of ‘dhurra-bird’ in Sudan reflects preference for millet; eats guinea-corn in W Africa. Only small grass seeds in stomachs (Chapin 1954). Breeding Habits. Polygynous, Y with up to 5 XX (Bowen 1926). Territorial, with boundaries well defined and respected by neighbouring YY. Forms small colonies; in Benin, 10 YY in area of 4 ha; less colonial and apparently more specialized in nesting requirements, than Southern Red Bishop E. orix (Winterbottom 1936). Territory may overlap with that of Black-winged Bishop. Breeding Y perches on prominent stem, wings make audible sound in

Euplectes orix flight. YY in non-breeding plumage also sing and display during breeding season. Captive birds reared 2 broods within 5 weeks (Steiner 1967). NEST: globular structure of coarse grass, lined with grass flower heads which project through entrance. Sited in and supported by grass or weed stems 1–2 m above ground; rarely in bamboo, bushes or trees, 3–6 m above ground; in crops, e.g. rice (Mali), sugarcane (Mauritania) and millet (Sudan). Density of nests in millet c. 10 times that in wild grasses (Huff 1979). Nest built by Y, starting with vertical ring between supporting vegetation. A captive X built her own nest, ignoring those provided by Y (Bullock 1938). EGGS: 2–4, plain blue. Laid at 1-day intervals (captive birds). SIZE: (n ¼ 48, Nigeria) 153–195  122–131 (173  128); (Darfur, Sudan) (av. 165  125); (Ethiopia) 165–175  130–132. LAYING DATES (in parentheses, YY in breeding plumage): Gambia and Senegal, Aug–Nov; Mali, May–Oct; Ivory Coast, (May–Oct); Ghana, July–Sept, (Apr–Oct; nest-building June); Togo, (nesting June–July); Benin,

217

July–Nov; Burkina Faso, June–Oct; Niger, Oct, (Aug– Oct); Nigeria, June–Oct, (from early July); Sudan, Feb– Mar, Aug-Nov; Ethiopia, May–Sept, (Dec); Somalia, in NW Apr, July, in S July, Sept (nest-building Dec, YY displaying Aug); NE Zaı¨re, May–Oct; Uganda (Oct). INCUBATION: by X only. Period: 13–14 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: young fed by X only, at first with insects. X removes faeces. Period: 14–16 days (in captivity, 23–27 days). In aviaries, young did not return to nest once fledged. Captive Y occasionally fed fledged young (Kleefisch 1971). BREEDING SUCCESS/SURVIVAL: hybridizes with E. afer, in captivity (Hopkinson 1938). Captive Y at least 8 years old, retained breeding plumage throughout year (Adlersparre 1938).

Key Reference Bowen, W. W. (1926).

Euplectes orix (Linnaeus). Southern Red Bishop. Euplecte ignicolore.

Plate 13

Loxia orix Linnaeus, 1758, Syst. Nat., ed. 10, p. 177; Angola.

(Opp. p. 202)

Forms a superspecies with E. franciscanus and E. nigroventris. Range and Status. Endemic resident. Introduced unsuccessfully to St Helena (Haydock 1954), Tahiti and Hawaii (Long 1981, Pratt et al. 1987), and S Australia (Long 1981, Blakers et al. 1984). Occasional sightings in Portugal (Costa et al. 1997) may refer to E. franciscanus. Zaı¨re, eastern region bordering lakes, south to E Katanga. Uganda, in SW, north to Equator. Kenya, L. Natron area, L. Kanyaboli, L. Magadi, wanderers north to L. Naivasha, L. Victoria basin, particularly around Ahero, Toro, Masaka, Kisumu, Siaya. Rwanda. Burundi. Tanzania, Tabora, Kigoma, West Lake east to Arusha and ˆ i at Kilimanjaro, south to Iringa and shores of L. Malaw Songea. Angola, S Huı´la, Namibe north to Benguela and W Huambo, isolated records at Dondo, Massangano. Zambia, Luapula R., L. Mweru and L. Tanganyika area, Chambeshi R., Bangweulu region, Luangwa valley, Eastern Prov., Lusaka Prov., southern Central Prov., Southern Prov., south and west parts of Western Prov. but absent from ˆ i, very other parts and from Northwestern Province. Malaw ˆ i littoral zone, also L. common in Shire valley, L. Malaw Chilwa; local at higher levels (Zomba, Dedza, Lilongwe, Mzimba and Rumphi districts). Mozambique, Mocuba, Tete and Zambezi valley, Buzi R., Beira, Save R., common in Sul do Save from Coguno southwards, in southern Mozambique, >5000 birds (Parker 1999). Namibia, Cunene and Caprivi region, central highlands south of Windhoek, Orange R. and tributaries in S. Botswana, Okavango, Linyati and Chobe R. south to L. Ngami, Mopipi and Orapa, along Shashe and Limpopo R. west to Serowe, also in SE around Lobatse, Kanye, Gaborone. Zimbabwe, widespread (probably absent from SE lowveld

Euplectes orix

before irrigation there: Brooke 1965), absent from drier NW, local in E highlands. South Africa, almost throughout except locally in North-West Prov., N Cape and Kalahari away from rivers, drier areas of Namaqualand. Swaziland, common throughout. Lesotho, in lowland valleys. Very

Euplectes orix flight. YY in non-breeding plumage also sing and display during breeding season. Captive birds reared 2 broods within 5 weeks (Steiner 1967). NEST: globular structure of coarse grass, lined with grass flower heads which project through entrance. Sited in and supported by grass or weed stems 1–2 m above ground; rarely in bamboo, bushes or trees, 3–6 m above ground; in crops, e.g. rice (Mali), sugarcane (Mauritania) and millet (Sudan). Density of nests in millet c. 10 times that in wild grasses (Huff 1979). Nest built by Y, starting with vertical ring between supporting vegetation. A captive X built her own nest, ignoring those provided by Y (Bullock 1938). EGGS: 2–4, plain blue. Laid at 1-day intervals (captive birds). SIZE: (n ¼ 48, Nigeria) 153–195  122–131 (173  128); (Darfur, Sudan) (av. 165  125); (Ethiopia) 165–175  130–132. LAYING DATES (in parentheses, YY in breeding plumage): Gambia and Senegal, Aug–Nov; Mali, May–Oct; Ivory Coast, (May–Oct); Ghana, July–Sept, (Apr–Oct; nest-building June); Togo, (nesting June–July); Benin,

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July–Nov; Burkina Faso, June–Oct; Niger, Oct, (Aug– Oct); Nigeria, June–Oct, (from early July); Sudan, Feb– Mar, Aug-Nov; Ethiopia, May–Sept, (Dec); Somalia, in NW Apr, July, in S July, Sept (nest-building Dec, YY displaying Aug); NE Zaı¨re, May–Oct; Uganda (Oct). INCUBATION: by X only. Period: 13–14 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: young fed by X only, at first with insects. X removes faeces. Period: 14–16 days (in captivity, 23–27 days). In aviaries, young did not return to nest once fledged. Captive Y occasionally fed fledged young (Kleefisch 1971). BREEDING SUCCESS/SURVIVAL: hybridizes with E. afer, in captivity (Hopkinson 1938). Captive Y at least 8 years old, retained breeding plumage throughout year (Adlersparre 1938).

Key Reference Bowen, W. W. (1926).

Euplectes orix (Linnaeus). Southern Red Bishop. Euplecte ignicolore.

Plate 13

Loxia orix Linnaeus, 1758, Syst. Nat., ed. 10, p. 177; Angola.

(Opp. p. 202)

Forms a superspecies with E. franciscanus and E. nigroventris. Range and Status. Endemic resident. Introduced unsuccessfully to St Helena (Haydock 1954), Tahiti and Hawaii (Long 1981, Pratt et al. 1987), and S Australia (Long 1981, Blakers et al. 1984). Occasional sightings in Portugal (Costa et al. 1997) may refer to E. franciscanus. Zaı¨re, eastern region bordering lakes, south to E Katanga. Uganda, in SW, north to Equator. Kenya, L. Natron area, L. Kanyaboli, L. Magadi, wanderers north to L. Naivasha, L. Victoria basin, particularly around Ahero, Toro, Masaka, Kisumu, Siaya. Rwanda. Burundi. Tanzania, Tabora, Kigoma, West Lake east to Arusha and ˆ i at Kilimanjaro, south to Iringa and shores of L. Malaw Songea. Angola, S Huı´la, Namibe north to Benguela and W Huambo, isolated records at Dondo, Massangano. Zambia, Luapula R., L. Mweru and L. Tanganyika area, Chambeshi R., Bangweulu region, Luangwa valley, Eastern Prov., Lusaka Prov., southern Central Prov., Southern Prov., south and west parts of Western Prov. but absent from ˆ i, very other parts and from Northwestern Province. Malaw ˆ i littoral zone, also L. common in Shire valley, L. Malaw Chilwa; local at higher levels (Zomba, Dedza, Lilongwe, Mzimba and Rumphi districts). Mozambique, Mocuba, Tete and Zambezi valley, Buzi R., Beira, Save R., common in Sul do Save from Coguno southwards, in southern Mozambique, >5000 birds (Parker 1999). Namibia, Cunene and Caprivi region, central highlands south of Windhoek, Orange R. and tributaries in S. Botswana, Okavango, Linyati and Chobe R. south to L. Ngami, Mopipi and Orapa, along Shashe and Limpopo R. west to Serowe, also in SE around Lobatse, Kanye, Gaborone. Zimbabwe, widespread (probably absent from SE lowveld

Euplectes orix

before irrigation there: Brooke 1965), absent from drier NW, local in E highlands. South Africa, almost throughout except locally in North-West Prov., N Cape and Kalahari away from rivers, drier areas of Namaqualand. Swaziland, common throughout. Lesotho, in lowland valleys. Very

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localized in arid zones; common to abundant. This species has benefited greatly from changes in agriculture, particularly the building of dams and irrigation canals, and expansion of seed crops. Description. ADULT Y (breeding): forehead black, crown, nape, rump and uppertail-coverts deep red to orange-red,. mantle redbrown (sometimes streaked with black: Ogilvie-Grant 1910), tail brown with paler edges; lores and cheeks black, ear-coverts red; chin and upper throat black (extent of black on chin varies individually); lower throat and breast red; belly black, thighs brown, undertail-coverts red; wings brown; underwing buff. Bill black, eyes brown, legs brown. ADULT Y (non-breeding): forehead to uppertail-coverts brown with dark central streaks; yellowish superciliary stripe; lores, cheeks and ear-coverts light brown; chin and throat whitish; breast and flanks off-white to buff with narrow central streaks on feathers; belly, vent and undertailcoverts white; wings brown. Bill brown, eyes brown, legs light brown to flesh-coloured. In Southern Africa, northern birds tend to have back feathers browner, less reddish than southern birds. ADULT X: like non-breeding Y, but underparts often less heavily streaked. SIZE (257 YY, 118 XX): wing, Y 57–81 (706), X 54–72 (648); tail, Y 35–47 (402), X 30–42 (364); bill, Y 136–175 (154), X 126–161 (145); tarsus, Y 187–255 (215), X 178–214 (195). Size varies clinally in southern Africa: wing Y (n ¼ 111, W Cape) 69–81 (748); (n ¼ 215, E Cape) 68–78 (732); (n ¼ 312, KwaZulu-Natal) 68–77 (719), (n ¼ 396, Zimbabwe) 67–75 (705). Within any population, birds can often be sexed on basis of winglength alone (Wilson 1973, Craig and Manson 1981). WEIGHT: (KwaZulu-Natal, South Africa) Y (n ¼ 704) 21–30 (255), X (n ¼ 445) 17–26 (207); (Zaı¨re) Y (n ¼ 16) 14–21 (175), 2 XX 14, ˆ i, n ¼ 7) 165–210 (193); juv. (North-west 15; sub-ad. (Malaw Province, South Africa, n ¼ 643) 162–299 (227). Heaviest in evening; in KwaZulu-Natal, Y heaviest early in breeding season and during post-nuptial moult, X heaviest during egg-laying period and during complete moult (Craig 1978). IMMATURE: like X, but broad buff margins to feathers on upperparts, buffy wash on underparts. NESTLING: on hatching, naked and pink-skinned except for small tufts of white plumes on head, centre of back. TAXONOMIC NOTE: races ‘orix’, ‘nigrifrons’, ‘sundevalli’ and ‘turgidus’ have been recognised, but we treat species as monotypic since populations are not reliably separable on measurements nor on plumage characters.

Field Characters. Length 11 cm. Black and red breeding Y smaller than Black-winged Bishop E. hordeaceus, with short brown tail and brown wings uniform with body in flight; undertail-coverts red, forehead and forecrown black. Red comes further forward on crown than similar but allopatric Northern Red Bishop E. franciscanus, tail-coverts shorter; told from Zanzibar Bishop E. nigroventris by broad red breast-band. Buffy breast-band with sharp streaks separates X and non-breeding Y from Zanzibar Bishop (indistinctly streaked), short whitish superciliary stripe from Yellow-crowned Bishop E. afer (long yellow supercilium), brown (not yellow) edges to primaries from Firefronted Bishop E. diadematus. X and non-breeding Y widowbirds have longer tails, often pointed or wedgeshaped; Red-collared Widowbird E. ardens has plain buff breast, grey underwing, pointed undertail-coverts with dark centres; Yellow-mantled Widowbird E. macroura is larger and darker below; X White-winged Widowbird E. albonotatus has cinnamon-brown shoulder patch, oliveyellow bend of wing, yellowish wash on face and breast, white underwing.

Voice. Tape-recorded (86, 88, 99, 104, B, X, LEM, PAY, WALK). Song a sharp slow sewing machine-like trill, ‘stukstukstukstukstuk . . . ’ mixed with long, low grating swizzles, high sibilant notes, nasal ‘zhaaa-ta-zhaaa’, repeated dry, rolled ‘tsurrrri, tsurrrri . . . ’, sometimes a down-slurred whistle, ‘tu-tyooo’ or ‘hee-heeooo’; sibilant form of song, ‘ss-ss-ss-ss-jaa-jaa-jaa-jaa-ss-ss’. Chorus from colony dominated by long chattering trills. Song given with fluffed plumage during ‘bumble-flights’ (see below) or from perched Y either courting or alone (Craig 1974). Calls include sparrow-like ‘chewp’ and sharp ‘chiz-chiz’; chipping notes given in contact, flight or alarm, with slight differences in tone; short rattle in response to other birds flying over may announce that territory is occupied. General Habits. Inhabits tall grassland and cultivation, typically open country near water at 600–1500 m in E Africa; up to 1700 m at Inyanga, Zimbabwe (Irwin 1981). Occurs in flocks, all year, from 200 birds. In nonbreeding season often flocks with Red-billed Queleas Quelea quelea and Red-headed Queleas Q. erythrops. A significant crop pest in wheat-lands of SW Cape, South Africa, where up to 70% of crop destroyed in some fields; mist-nests used locally in control (McVeigh 1987); damage to sorghum and other cereals noted by early travellers. Birds generally forage in 2 sessions each day, morning and late afternoon, gathering in day roosts in between. Termite alates are caught and eaten on ground, or caught on wing then eaten on ground or on perch (Brooke 1970). Forages in rotting seaweed along W Cape shorelines, alongside European Starlings Sturnus vulgaris, apparently taking kelp fly larvae and ads, and perhaps amphipods. When drinking, sips and then tips bill upwards to swallow. Uses regular roost sites in association with other ploceids such as Cape Weaver Ploceus capensis and Redcollared Widowbird, Fan-tailed Widowbird E. axillaris, Yellow Bishop E. capensis and Long-tailed Widowbird E. progne. Up to 700 birds used a single roost site in KwaZulu-Natal (pers. obs.); a significant excess of YY at roosts in Zimbabwe and KwaZulu-Natal, for 6–7 months per year (Craig and Manson 1979b). In SW Cape YY outnumber XX (in 9 monthly samples); but sex ratio of 44 YY in breeding plumage to 100 breeding XX. Juvenile sex ratios seem to be balanced. Differences may depend on behaviour of specific sex and age classes, and partial segregation of sexes in non-breeding season (Hey ¨l 1980b). Seasonal plumage change depends on a partial moult of body plumage, excluding wing- and tail-feathers; but some imm. XX may replace tail-feathers; Y usually acquires breeding plumage in 2nd year but, based on skull ossification, 13% of YY in breeding plumage in Free State were 1styear birds (Roos and Kok 1979). However, in SW Cape Y takes up to 2 years to complete skull ossification and few if any 1st-year YY come into breeding plumage; possible that X shows more rapid skull ossification (Hey ¨l 1980a); most XX breed in 1st year (Friedl 1998). A population takes 4 months to complete post-nuptial moult; timing varies regionally in relation to breeding season. Body plumage is replaced within 6 weeks, while wing-moult in individual birds takes 110–120 days. Juvs undergo complete moult

Euplectes orix within 3 months of leaving nest (Roos and Kok 1979, Craig and Manson 1979a, Craig et al. 2001). Resident, but moves locally in non-breeding season in Botswana and probably elsewhere, especially in semi-arid habitats. Only 1% of ringed birds recovered were >100 km from ringing site (Craig 1982d). Y returns to same colony and same territory for up to 5 successive breeding seasons; X return rate is much lower (pers. obs., Friedl 1998). Food. Seeds; also insects in summer (South Africa) and when feeding young. In Free State, eats 24 seed species, mostly small; greatest variety taken in summer, but largest crop masses in winter; main seeds are Zea mays, Eragrostris curvula, Setaria flabellate, Panicum maximum, Eleusine indica and Triticum (Roos and Kok 1978). Feeds in flowers of Phragmites australis and Leonotis in SW Cape, South Africa, taking either nectar or insects (Fraser and McMahon 1989); in Gauteng, feeds on flowers of Rhynchelytrum repens (Skead 1975). Insect food 05% of crop contents by dry weight, 67% composed of beetles and their pupae. In E Cape insects include scarab beetles, winged termites, and kelp flies; small crustaceans Talorchestia taken on shoreline; grit fragments the size of sugar grains in stomachs (Skead 1997). Chicks fed arthropods including dragonflies, beetles, caterpillars and spiders; eggshell fragments in chick stomachs. Breeding Habits. Polygynous, with up to 7 XX nesting in Y’s territory at once; colonial, with colonies of several hundred nests in reedbeds; nests are also in small groups and occasionally a pair nests solitarily. Territorial, defending area of 3 m2 in reedbed or 300 m2 in more open grassland. Y displaces intruders by ‘supplant chases’ with plumage not erected except around the head. At territory boundaries, neighbouring YY threaten with tail spread, neck stretched up and black bill and face mask surrounded by red ruff. Contest is silent, and seldom progresses to fighting; occasionally birds grapple, and may fall into water locked together. Once territories established, YY respect boundaries; intruders immediately recognizable by their sleeked plumage; XX ignore territories and defend only immediate vicinity of nest entrance against other birds. Aggressive interactions occur between Y Southern Red Bishop and nesting Ploceus weavers; warblers nesting very close to bishop nests in reedbeds are ignored. On arrival of X (or any bird in non-breeding plumage) Y approaches with fluffed plumage in slow ‘bumble-flight’, alights near it (A) and, if it does not fly away, proceeds with perched courtship. With body plumage fluffed, beak depressed on chest, and sizzling calls accompanying swivelling movements around reed stems (B), Y circles, progressively approaching X. X enters nest, examines and tugs at structure while Y is outside; she does not stay in territory if no unoccupied nest frame is available. Finally she solicits copulation by crouching, quivering and drooping wings. Sub-adult Y displays with sizzling calls, may build incomplete nests at satellite colony, usually occupied only for short period, late in season. Ad. Y develops testes earlier than yearlings; testes regress later; ad. X has rapid follicle development (Roos and Kok 1977). Y bill colour

closely correlated with testis development (Craig 1975). Captive birds exposed to long photoperiods remain in nuptial plumage, but follow normal cycle of wing-moult and their testes regress (Craig 1985). NEST: oval, with side entrance under a porch near the top, tightly woven of thin strips of reed or grass blades (C), lined with plant down and grass seed heads. Size 104–115 high, 70–77 wide, 70–71 deep; entrance diam. 28–39  44. Nests most often in reeds, sedges or bulrushes in shallow water; also in bamboo thickets (Zambia), weedy thickets of Indigofera (Mozambique: Vincent 1936), privet hedges (Lane 1930), weeds or crops including maize away from water. In Phragmites reeds av. 283 m tall, mean nest height above water 201 m; stem diam. 10–12 (Zimbabwe: Woodall 1984). In E Cape, breeds annually in some perennial reedbeds but only sporadically in others (Skead 1965). Nest built by Y only; starts as vertical ring between

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PLOCEIDAE

supports (D, X visiting), Y perching on cross-piece whilst building; may be completed in a single day, but usually takes 2–3 days. Initially eggs visible through weaving; X adds lining during incubation. EGGS: 1–5 (av. 30, n ¼ 670, South Africa). Plain turquoise blue, usually pale; rarely with some faint dark spots. Clutches of 7 probably produced by 2 XX (Vincent 1949). Eggs normally laid on successive days; av. clutch size is larger in years of heavier rainfall. SIZE: (Zambia, n ¼ 5) 164–180  124–135 (170  131); (Zimbabwe, n ¼ 8) 169–185  131–139 (177  135); (SW Cape, n ¼ 17) 180–202  136–148 (194  142). WEIGHT: 173; water loss from eggs 166 mg per day (Woodall and Parry 1982). LAYING DATES: Uganda, Apr–June; Kenya, Apr–June; S Zaı¨re, Jan–Apr; Angola, Jan–Apr (prenuptial moult Nov); Rwanda, Mar–May; Uganda, Apr–June; Kenya, Apr–June; Tanzania, Feb–May; Angola, Dec; Zambia, Dec–Mar; ˆ i, Dec–Apr; Mozambique, Feb–Mar; Namibia, Malaw Feb–May; Botswana, Dec–Mar; Zimbabwe, Dec–Mar, rarely Nov, Apr; SW Cape, July–Dec; Transvaal, Oct– Apr; Free State, Nov–Mar, sometimes till Apr; KwaZuluNatal, Oct–Apr; E Cape, Oct–Mar. Except for SW Cape, rarely breeds before rainy season or more than 1 month after it. Rainfall may delay start of nesting and influence number of nests built; rainfall whilst breeding can also be influential (Brooke 1966, Craig 1982a, Friedl in press). 2 peaks in egg-laying at colonies suggest that birds may be double-brooded. INCUBATION: by X only; may start before clutch is complete. In KwaZulu-Natal, X on nest for only 40% of day time whatever the clutch size, with bouts on nest of 1– 10 min and absences of 1–12 min (Slotow et al. 1995). Period: 12–13 days. Eggshells dropped 20 cm from nest, but unhatched eggs left in nest.

DEVELOPMENT AND CARE OF YOUNG: young fed by X only, with insects and seeds regurgitated from crop. Feather quills appear on day 4, eyes open on day 6, feathers open from day 8. Nestling period: 11–15 days, but young liable to leave nest after only 10 days if disturbed. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius; rates of parasitism range from 7 to 50%. Up to 4 X cuckoos can lay in large colony; Y Southern Red Bishop regularly attacks them and only small colonies have high rates of parasitism, suggesting that defence is effective (Ferguson 1994a, Lawes and Kirkman 1996). Nest height has no apparent influence on breeding success in reeds (South Africa, Zimbabwe), although flooding may be significant source of mortality in some years. In SW Cape, 66% of eggs hatched, fledging success 45% at one colony (Schmidt 1968); at another site only 50% of eggs hatched, though fledging success overall was 35% (Craig 1982b). Breeding success in 4 successive seasons at Addo Elephant Nat. Park was 13%, 38%, 14% and 32%, with predation more important than rainfall (Friedl in press). Over 2 seasons at a KwaZulu-Natal colony, 9% of eggs produced flying young; predation was main source of mortality, and in one season 36% of eggs and 40% of chicks taken by rats, water monitors Varanus niloticus and other predators; Cattle Egrets Bubulcus ibis remove chicks from some nests (Craig 1982b). At Willem Pretorius Game Res., Free State, breeding success was 336%, with rodents and egg-eating snakes Dasypeltis scabra the main predators; some chicks died after heavy rain (Kok et al. 1977). A reedbed colony in Gauteng had 25% breeding success over 2 seasons; predation was the most important mortality factor (probably rodents, slender mongoose Herpestes pulverulentus, Burchell’s Coucal Centropus burchelli: Ferguson 1994).

Euplectes nigroventris Adults preyed on by Barn Owl Tyto alba in South Africa (Dean 1973). Breeding success of 6 YY varied considerably at a colony in KwaZulu-Natal; territory size was 2–12 m2; 3–13 nests occupied by 3–8 XX which laid 9–19 eggs; most successful Y (3 chicks from 9 eggs) had 3 occupied nests in 2 m2 territory, whereas Y most active in courtship, with 8 XX laying 19 eggs, fledged no young (Craig 1982c). Duration that Y holds territory, and his nest-building performance, are main factors determining mating success, although X selection of a territory and nest site appears to be random (Friedl and Klump 1999, 2000). Old nests used by Tawny-flanked Prinia Prinia subflava and (often) Zebra Waxbill Sporaeginthus subflavus and others; in SW Cape, climbing mice Dendromus mesomelas occupy nests but leaves eggs untouched (Schmidt 1968).

223

Captive Y once mated with its daughter of previous season (Schifter 1967). In captivity Y E. orix and X E. ardens produced a hybrid (Colahan and Craig 1981). Bone structure suggests lifespan of 7 years (Brown ˆ i: Hanmer 1989), 1994); ringed birds >125 years (Malaw 118 years (South Africa: Yom-Tov et al. 1994). Annual survival rate estimated at 715% from recaptures over a 17year period (Peach et al. 2001). Calculated mortality rate for ringed birds in South Africa is 44%, and for birds recaptured in Zimbabwe 47%, with no difference between Y and X. Key References Craig, A. J. F. K. (1974, 1978, 1982a, b, c, d), Craig, A. J. F. K. and Manson, A. J. (1979a, b, c, 1981), Friedl, T. W. P. (1998 and in press), Friedl, T. W. P. and Klump, G. M. (1999, 2000), Skead, C. J. (1956b), Woodall, P. (1971).

Euplectes nigroventris Cassin. Zanzibar Bishop. Euplecte de Zanzibar.

Plate 13

Euplectes nigroventris Cassin, 1848. Proc. Acad. Nat. Sci. Philadelphia, 4, p. 66; Zanzibar.

(Opp. p. 202)

Forms a superspecies with E. orix and E. franciscanus. Range and Status. Endemic resident; coastal belt of SE Kenya, from Lamu south, inland to Tsavo, L. Jipe and Taveta; Tanzania, on coast including Zanzibar and Pemba, no further west than Eastern Arc mountains except along Pangani R., north to Nyumba Ya Mungu, inland locally west of Songea; N Mozambique, southernmost record at Mocuba. Locally common. Description. ADULT Y (breeding): forehead, crown and nape red, mantle bright rufous-brown, rump and uppertail-coverts red; tail brown with paler edging; lores, cheeks and ear-coverts black; chin, throat, breast and belly black; thighs brown; undertailcoverts red; wings brown with paler buffy edges; underwing buff. Bill black; eyes dark brown; legs brown. Variable; inland, some birds (‘rufigula’) have scattered red feathers on throat and upper breast. ADULT Y (non-breeding): from forehead to uppertailcoverts light brown with dark brown central streaks on feathers; lores, cheeks and ear-coverts light brown with faint streaks; chin, throat, breast and belly dull white, with buffy wash in centre of breast; thighs buffy; undertail-coverts white. Wings and tail as in breeding Y. Bill brown. ADULT X: like non-breeding Y. SIZE (58 YY, 24 XX): wing, Y 55–64 (588), X 52–58 (539); tail, Y 32–40 (354), X 26–36 (315); bill, Y 131–148 (139), X 127–141 (134); tarsus, Y 170–197 (181), X 159–186 (171). WEIGHT: (Kenya) 1 Y 136; 2 XX 107, 109; unsexed (n ¼ 12) 111–138 (125). IMMATURE: like ad. X, with broader and paler buff edges to feathers of upperparts. NESTLING: no information.

Field Characters. Length 10 cm. An E coast bishop. Breeding Y differs from very similar but allopatric Northern and Southern Red Bishops E. franciscanus and E. orix by entirely red crown (including forehead) and lack of red breast-band on black underparts; from sympatric Black-winged Bishop E. hordeaceus by short brown tail, brown wings, red undertail-coverts. X and non-breeding Y smaller than Black-winged Bishop, with shorter bill, paler and duller buff breast-band; they lack the yellow

Euplectes nigroventris

?

wing-edgings of Fire-fronted Bishop E. diadematus, and are less rufous brown above. Voice. Tape-recorded (B, GREG, McVIC, STJ). Song like sewing machine, thin, dry trills on 1 pitch, at varying speeds, mixed with variety of calls, including low grating chatter, nasal ‘zhhhhnn’, ‘djeeeeen’ and ‘jeeya’, long downslurred ‘peeeeeurrr’ and some short swizzles. Calls, sharp ‘tsip’, buzzy ‘dzip’ or ‘zzzeee’ and hard low ‘drrt’.

Euplectes nigroventris Adults preyed on by Barn Owl Tyto alba in South Africa (Dean 1973). Breeding success of 6 YY varied considerably at a colony in KwaZulu-Natal; territory size was 2–12 m2; 3–13 nests occupied by 3–8 XX which laid 9–19 eggs; most successful Y (3 chicks from 9 eggs) had 3 occupied nests in 2 m2 territory, whereas Y most active in courtship, with 8 XX laying 19 eggs, fledged no young (Craig 1982c). Duration that Y holds territory, and his nest-building performance, are main factors determining mating success, although X selection of a territory and nest site appears to be random (Friedl and Klump 1999, 2000). Old nests used by Tawny-flanked Prinia Prinia subflava and (often) Zebra Waxbill Sporaeginthus subflavus and others; in SW Cape, climbing mice Dendromus mesomelas occupy nests but leaves eggs untouched (Schmidt 1968).

223

Captive Y once mated with its daughter of previous season (Schifter 1967). In captivity Y E. orix and X E. ardens produced a hybrid (Colahan and Craig 1981). Bone structure suggests lifespan of 7 years (Brown ˆ i: Hanmer 1989), 1994); ringed birds >125 years (Malaw 118 years (South Africa: Yom-Tov et al. 1994). Annual survival rate estimated at 715% from recaptures over a 17year period (Peach et al. 2001). Calculated mortality rate for ringed birds in South Africa is 44%, and for birds recaptured in Zimbabwe 47%, with no difference between Y and X. Key References Craig, A. J. F. K. (1974, 1978, 1982a, b, c, d), Craig, A. J. F. K. and Manson, A. J. (1979a, b, c, 1981), Friedl, T. W. P. (1998 and in press), Friedl, T. W. P. and Klump, G. M. (1999, 2000), Skead, C. J. (1956b), Woodall, P. (1971).

Euplectes nigroventris Cassin. Zanzibar Bishop. Euplecte de Zanzibar.

Plate 13

Euplectes nigroventris Cassin, 1848. Proc. Acad. Nat. Sci. Philadelphia, 4, p. 66; Zanzibar.

(Opp. p. 202)

Forms a superspecies with E. orix and E. franciscanus. Range and Status. Endemic resident; coastal belt of SE Kenya, from Lamu south, inland to Tsavo, L. Jipe and Taveta; Tanzania, on coast including Zanzibar and Pemba, no further west than Eastern Arc mountains except along Pangani R., north to Nyumba Ya Mungu, inland locally west of Songea; N Mozambique, southernmost record at Mocuba. Locally common. Description. ADULT Y (breeding): forehead, crown and nape red, mantle bright rufous-brown, rump and uppertail-coverts red; tail brown with paler edging; lores, cheeks and ear-coverts black; chin, throat, breast and belly black; thighs brown; undertailcoverts red; wings brown with paler buffy edges; underwing buff. Bill black; eyes dark brown; legs brown. Variable; inland, some birds (‘rufigula’) have scattered red feathers on throat and upper breast. ADULT Y (non-breeding): from forehead to uppertailcoverts light brown with dark brown central streaks on feathers; lores, cheeks and ear-coverts light brown with faint streaks; chin, throat, breast and belly dull white, with buffy wash in centre of breast; thighs buffy; undertail-coverts white. Wings and tail as in breeding Y. Bill brown. ADULT X: like non-breeding Y. SIZE (58 YY, 24 XX): wing, Y 55–64 (588), X 52–58 (539); tail, Y 32–40 (354), X 26–36 (315); bill, Y 131–148 (139), X 127–141 (134); tarsus, Y 170–197 (181), X 159–186 (171). WEIGHT: (Kenya) 1 Y 136; 2 XX 107, 109; unsexed (n ¼ 12) 111–138 (125). IMMATURE: like ad. X, with broader and paler buff edges to feathers of upperparts. NESTLING: no information.

Field Characters. Length 10 cm. An E coast bishop. Breeding Y differs from very similar but allopatric Northern and Southern Red Bishops E. franciscanus and E. orix by entirely red crown (including forehead) and lack of red breast-band on black underparts; from sympatric Black-winged Bishop E. hordeaceus by short brown tail, brown wings, red undertail-coverts. X and non-breeding Y smaller than Black-winged Bishop, with shorter bill, paler and duller buff breast-band; they lack the yellow

Euplectes nigroventris

?

wing-edgings of Fire-fronted Bishop E. diadematus, and are less rufous brown above. Voice. Tape-recorded (B, GREG, McVIC, STJ). Song like sewing machine, thin, dry trills on 1 pitch, at varying speeds, mixed with variety of calls, including low grating chatter, nasal ‘zhhhhnn’, ‘djeeeeen’ and ‘jeeya’, long downslurred ‘peeeeeurrr’ and some short swizzles. Calls, sharp ‘tsip’, buzzy ‘dzip’ or ‘zzzeee’ and hard low ‘drrt’.

224

PLOCEIDAE

General Habits. Inhabits coastal grasslands and cultivation below 1000 m. Always in flocks, moves away from nesting areas after breeding; rare visitor to riverside areas in Tsavo (Lack 1985). Often in flocks with Black-winged Bishop, also with other bishops and queleas. At Mombasa, wing-moult in May–Sept. Food. Grass seeds of Echinochloa haploclada, Panicum maximum; also sorghum and rice; termites and spiders in captivity. Breeding Habits. Polygynous, up to 5 XX per Y. Territorial, territory size varies from 120 to 240 m2 in grassland and down to 10 m2 in reedbeds (Fuggles-Couchman 1943); from 75–100 m2 in pure stands of Phragmites reeds and 50– 800 m2 in Typha bulrushes to >200 m2 in grassland and mixed herbage (Moreau and Moreau 1938). In grassland, YY feed within territory. Territory may lack special vantage point, or have a favoured perch. Y postures on borders, defending territory against conspecifics (also against Black-winged Bishop, whereas Red-collared Widowbird E. ardens and White-winged Widowbird E. albonotatus are tolerated within territory). Y approaches X in short fluttering flights, with ruffled plumage; in perched courtship, turns around repeatedly, displaying both upperparts and underparts.

Plate 13 (Opp. p. 202)

NEST: thin-walled oval structure of coarse grass with side entrance, lined with seed-heads of grasses such as Panicum maximum; grass heads project over entrance to form a small porch. Attached to grass, bushes or reed stems, usually 1–15 m above ground, typically supported by 2 stems, 1 each side of entrance. Occupied nest in woody shrub measured 102  67  59, entrance 34  36 (Bowen 1931). Early in season, nests most often in reeds or bulrushes. Nest built by Y, but X adds the lining. EGGS: 1–3, av. 24 (n ¼ 19, Tanzania). Pale blue, rarely with sparse brown speckles. SIZE: (n ¼ 3, Tanzania) 157– 175  120–127. LAYING DATES: Kenya, May–Oct, peaking in Aug; Tanzania, all months; on Zanzibar, May–July, (possibly again Nov–Dec, Vaughan 1930). In one year nested in Apr, and YY were again in nuptial plumage following rain in Dec, when they built nests but XX did not lay (Tanzania: Moreau 1950). INCUBATION: in captivity by X only; period 12 days. DEVELOPMENT AND CARE OF YOUNG: young fed on insects; nestling period 13–16 days, in captivity 15–19 days. BREEDING SUCCESS/SURVIVAL: at Mombasa, eggs and young taken by House Crows Corvus splendens (Ryall 1992).

Key References Fuggles-Couchman, N. R. (1943), Moreau, R. E. and Moreau, W. M. (1938), Vaughan, J. H. (1930).

Euplectes hordeaceus (Linnaeus). Black-winged Bishop; Fire-crowned Bishop. Euplecte monseigneur. Loxia hordeacea Linnaeus, 1758. Syst. Nat., ed. 10, p. 173; Senegal. Range and Status. Endemic resident; common. Mauritania, Senegal R. delta and valley (Gorgol, Karakoro, Toya, Wadon and some localities north to 17
N). Gambia, throughout, but least numerous of the 3 bishops. Senegal, Dakar, Basse Casamance east to Niokolo Koba. Mali, quite common as mapped, mainly at 12–15
N, moving to 16
N in some years (Lamarche 1993). The Mauritania/Mali population appears to be quite separate from larger range to south. Guinea-Bissau, local on coast. Guinea, widespread in south as at Kindia, Macenta. Sierra Leone, Freetown, Koindu, Woroma, Bo, Rotifunk, Rokupr. Liberia, common inland, uncommon but widely distributed along most of coast. Ivory Coast, almost throughout: marshes and rice fields on coast, large clearings in forest zone, in savanna. Ghana, coastal zone, Accra Plains, Volta Region, N savanna; seasonal in some areas. Togo, throughout, north as far as Borgou and Namoundjoga. Benin, savannas south to Be´te´rou, occasional closer to coast. Burkina Faso, in S half, north to Ouagadougou; Arli, Boromo, Banfora, Po, Gaoua. Nigeria, throughout Guinea savanna zone north to Kano and Aliya; absent from E and SE, uncommon in S. Cameroon, Bamenda, Adamawa Plateau, Tibati Plateau, Benue plain, local in N at Koza, Maroua. Sa˜o Tome´, first recorded in 1893, and possibly introduced; common in N and NE and in city. Niger, in SW at Mari. Central African Republic, Manovo, Lobaye,

Euplectes hordeaceus

Bangui, Haute Sangha, Bamingui-Bangoran Nat. Park, Dzanga Reserves. Chad, local, as mapped. Sudan, local in

224

PLOCEIDAE

General Habits. Inhabits coastal grasslands and cultivation below 1000 m. Always in flocks, moves away from nesting areas after breeding; rare visitor to riverside areas in Tsavo (Lack 1985). Often in flocks with Black-winged Bishop, also with other bishops and queleas. At Mombasa, wing-moult in May–Sept. Food. Grass seeds of Echinochloa haploclada, Panicum maximum; also sorghum and rice; termites and spiders in captivity. Breeding Habits. Polygynous, up to 5 XX per Y. Territorial, territory size varies from 120 to 240 m2 in grassland and down to 10 m2 in reedbeds (Fuggles-Couchman 1943); from 75–100 m2 in pure stands of Phragmites reeds and 50– 800 m2 in Typha bulrushes to >200 m2 in grassland and mixed herbage (Moreau and Moreau 1938). In grassland, YY feed within territory. Territory may lack special vantage point, or have a favoured perch. Y postures on borders, defending territory against conspecifics (also against Black-winged Bishop, whereas Red-collared Widowbird E. ardens and White-winged Widowbird E. albonotatus are tolerated within territory). Y approaches X in short fluttering flights, with ruffled plumage; in perched courtship, turns around repeatedly, displaying both upperparts and underparts.

Plate 13 (Opp. p. 202)

NEST: thin-walled oval structure of coarse grass with side entrance, lined with seed-heads of grasses such as Panicum maximum; grass heads project over entrance to form a small porch. Attached to grass, bushes or reed stems, usually 1–15 m above ground, typically supported by 2 stems, 1 each side of entrance. Occupied nest in woody shrub measured 102  67  59, entrance 34  36 (Bowen 1931). Early in season, nests most often in reeds or bulrushes. Nest built by Y, but X adds the lining. EGGS: 1–3, av. 24 (n ¼ 19, Tanzania). Pale blue, rarely with sparse brown speckles. SIZE: (n ¼ 3, Tanzania) 157– 175  120–127. LAYING DATES: Kenya, May–Oct, peaking in Aug; Tanzania, all months; on Zanzibar, May–July, (possibly again Nov–Dec, Vaughan 1930). In one year nested in Apr, and YY were again in nuptial plumage following rain in Dec, when they built nests but XX did not lay (Tanzania: Moreau 1950). INCUBATION: in captivity by X only; period 12 days. DEVELOPMENT AND CARE OF YOUNG: young fed on insects; nestling period 13–16 days, in captivity 15–19 days. BREEDING SUCCESS/SURVIVAL: at Mombasa, eggs and young taken by House Crows Corvus splendens (Ryall 1992).

Key References Fuggles-Couchman, N. R. (1943), Moreau, R. E. and Moreau, W. M. (1938), Vaughan, J. H. (1930).

Euplectes hordeaceus (Linnaeus). Black-winged Bishop; Fire-crowned Bishop. Euplecte monseigneur. Loxia hordeacea Linnaeus, 1758. Syst. Nat., ed. 10, p. 173; Senegal. Range and Status. Endemic resident; common. Mauritania, Senegal R. delta and valley (Gorgol, Karakoro, Toya, Wadon and some localities north to 17
N). Gambia, throughout, but least numerous of the 3 bishops. Senegal, Dakar, Basse Casamance east to Niokolo Koba. Mali, quite common as mapped, mainly at 12–15
N, moving to 16
N in some years (Lamarche 1993). The Mauritania/Mali population appears to be quite separate from larger range to south. Guinea-Bissau, local on coast. Guinea, widespread in south as at Kindia, Macenta. Sierra Leone, Freetown, Koindu, Woroma, Bo, Rotifunk, Rokupr. Liberia, common inland, uncommon but widely distributed along most of coast. Ivory Coast, almost throughout: marshes and rice fields on coast, large clearings in forest zone, in savanna. Ghana, coastal zone, Accra Plains, Volta Region, N savanna; seasonal in some areas. Togo, throughout, north as far as Borgou and Namoundjoga. Benin, savannas south to Be´te´rou, occasional closer to coast. Burkina Faso, in S half, north to Ouagadougou; Arli, Boromo, Banfora, Po, Gaoua. Nigeria, throughout Guinea savanna zone north to Kano and Aliya; absent from E and SE, uncommon in S. Cameroon, Bamenda, Adamawa Plateau, Tibati Plateau, Benue plain, local in N at Koza, Maroua. Sa˜o Tome´, first recorded in 1893, and possibly introduced; common in N and NE and in city. Niger, in SW at Mari. Central African Republic, Manovo, Lobaye,

Euplectes hordeaceus

Bangui, Haute Sangha, Bamingui-Bangoran Nat. Park, Dzanga Reserves. Chad, local, as mapped. Sudan, local in

Euplectes hordeaceus Darfur with apparent influx in breeding season; also in S area of Nile floodplain. Ethiopia, SW, including highlands and Rift Valley. Gabon, Mouila, Tchibanga, Labamba, Mimongo, Djambala, Gamboma. Congo, widespread in S, along Congo R. Zaı¨re, along rivers and clearings in forested areas, throughout savanna below 1300 m. Uganda, south to Toro, Mubende and NE shore of L. Victoria, also Elgon and S Nyanza. Kenya, coastal lowlands inland to Tiva R. floodplain, Taru and L. Jipe, also west of Rift Valley; vagrant to Marsabit. Rwanda; Burundi, outside of forests. Tanzania, as mapped; coastal lowlands. Zanzibar and Pemba; widespread south of 5
S to Shinyanga, Biharamulo and Ngara in W. Angola, Cabinda, south to Cuanza Norte, Huı´la, east to Lunda Norte, Lunda Sul and N Moxico. Zambia, mainly east of 25
, S-central and NW sectors of North-Western Prov., locally absent in Luapula, ˆ i, widespread below Northern and Central Prov. Malaw 1200 m, especially common on lake littoral. Zimbabwe, middle Zambezi Valley, lower-lying eastern areas and Save R. area. Mozambique, Manica Platform, Manica e Sofala, Zambezi R., Mocuba. Description. ADULT Y (breeding): forehead, crown and nape orange-red to deep red (sometimes a narrow black band just above base of bill); mantle red-brown, rump and uppertail-coverts orange-red to red; tail black with narrow pale brown margins; lores, cheeks and ear-coverts black; chin and throat black; breast orange-red to red, linked to red of dorsum; belly black; thighs and undertail-coverts black with paler margins; wings black; underwing black; in Sudan, Ethiopia, Uganda and W Kenya undertail-coverts usually whitish (‘craspedopterus’). Intensity of red varies within a population. Bill black; eyes dark brown; legs dark brown. Specimen from Zambia had sooty face and breast, black wings and belly; bill black (Winterbottom 1938). ADULT Y (non-breeding): dorsal plumage brown with black central streaks; tail black with brown edges; broad yellowish superciliary stripe; lores, cheeks and ear-coverts light brown; chin and throat whitish; breast light brown with some streaking; flanks and thighs light brown; belly and undertail-coverts whitish; wings black with paler edges, and broad buff edges on wing-coverts. Bill brown. Y acquires black wing feathers before first breeding plumage. SUB-ADULT Y: dorsal plumage brown with dark brown central streaks; tail brown with paler margins; yellowish superciliary stripe; lores, cheeks and ear-coverts brown to buffy; chin and throat whitish; breast, flanks and thighs brown to buffy with variable streaking on breast; belly and undertail-coverts whitish; wings brown with buff edges; underwing dark grey. Bill brown. Y does not acquire breeding plumage until 2nd year. ADULT X: like sub-ad. Y. SIZE (137 YY, 22 sub-ad. YY, 87 XX): wing, Y 69–85 (757), sub-ad. Y 66–76 (717), X 63–74 (666); tail, Y 41–51 (460), sub-ad. Y 38–48 (426), X 35–45 (390); bill, Y 150–186 (171), sub-ad. Y 156–182 (171), X 150–173 (163); tarsus, Y 208–240 (224), sub-ad. Y 211–234 (223), X 193–222 (210). WEIGHT: Y (Zaı ¨re, n ¼ 10) 22–29 (253), (Kenya, n ¼ 12) 180–229 ˆ i, Mozambique, n ¼ 14) 23–28 (255); X (Kenya (204), (Malaw ˆ i, n ¼ 7) 18–21 (196); unsexed n ¼ 6) 172–195 (184), (Malaw (Nigeria, n ¼ 5) 155–210 (183). IMMATURE: like X, but first plumage with broad buff edges to feathers of upperparts. NESTLING: scanty white down; inside of mouth deep fleshy red, white gape.

Field Characters. Length 12–14 cm. Larger than Northern, Southern and Zanzibar Red Bishops E. franciscanus, E. orix and E. nigroventris, with heavier bill and longer tail. Red and black breeding Y has black wings, black tail

extending well beyond uppertail-coverts, entirely red crown, buffy or whitish undertail-coverts; Zanzibar Bishop has red crown but no red on breast. X and non-breeding Y have broad yellowish supercilium, rich brown breast-band; smaller and browner above than Black Bishop E. gierowii, breast lightly streaked (no spots); in flight, black wings (retained in non-breeding season) contrast with brown upperparts (wings brown in other red bishops). Birds with buffy breast closely resemble non-breeding Red-collared Widowbird E. ardens but have square (not pointed) tail, heavier bill, white throat. Voice. Tape-recorded (86, 99, 104, B, C, F, CART, MOR, PAY). Song of hard sibilant notes usually in 3 parts, with swizzle or rattle in middle, ‘chussaka-tsipa-tsipa-tsipazzzzzzh-tututututu’ or ‘chee-chee-chee-chee-cheetarrrrrrrrrr-chee-chee-chee’; central swizzle may be omitted, leaving a medley of chattering notes at different speeds. High thin whistles and buzzy notes can also be run together into a song, ‘seep-seep-seep, zzzzzray-titizzzzzray-titi-zzzzzray, sisisisisi’. Varied calls given from perch on territory, including hard ‘dzzt’, shrill sibilant trill, sharp ‘tsap’ or ‘tsip’ and brief liquid trill. In courtship gives melodious notes only audible at close range. X soliciting copulation utters soft, high-pitched call; Y on territory used distinctive twittering to greet returning X (Lack 1985). General Habits. Inhabits bushed grassland, rice fields, and tall grassland, as well as open areas in forested country; spends much time near tops of tall grasses, but forages mainly on ground in savanna, often concentrated at sites of abandoned cultivation, with secondary growth of tall grasses. E African coastal population, up to 500 m, is often sympatric with E. nigroventris; occurs inland at 100–2000 m, in wetter areas than E. orix. Usually in pairs or small groups; forms large roving flocks in non-breeding season. In W Africa, common in open Pennisetum grassland; often forms mixed flocks with E. franciscanus. On Sa˜o Tome´ in northern savannas and towns, in tall grass and damp areas; nonbreeding birds flock with E. aureus. In Sudan, forms mixed flocks with E. franciscanus and White-winged Widowbird E. albonotatus; on Zanzibar flocks with E. nigroventris and Eastern Golden Weaver Ploceus subaureus. On Zanzibar, established territories on grassy slopes in Panicum maximum; in Uganda, nests in swamps and reedbeds. Anting recorded in captive birds (Poulsen 1956). Hawks termite alates, returning to a perch to eat them. Food. Grass seeds, particularly Panicum maximum, Hyparrhenia, Pennisetum, Rottboellia; also rice and green maize. Stomach contents include unripe grass seeds and caterpillars. Breeding Habits. Polygynous, Y with up to 4 XX (and nests); highly territorial, defending area of 400 m2 (Zaı¨re) and 500–1200 m2 (Tanzania). People on Zanzibar claim that YY are attracted to red flower or red cloth (where caught on bird-lime) (Vaughan 1930). In Zimbabwe tolerates E. orix nesting within territory (R.K. Brooke, pers. comm.); nests alongside Golden-backed Bishop E. aureus on Sa˜o Tome´; in

225

226

PLOCEIDAE

Tanzania little interaction with Yellow Bishop E. capensis where territories overlap (Lack 1985). Aggressive towards Y E. ardens (Tanzania), Y displacing it from territory (Liberia). Y chases conspecific YY from territory; Y pursuing intruder into territory of neighbouring Y might be attacked by neighbouring Y and receive ‘a severe buffet’ (Lack 1935). Strange YY often visit territory while territory-holder is absent; return of resident Y is marked by fluffed-plumage descending flight with loud calls and audible wing-beats. Away from territory YY feed close together without aggression. XX appear to ignore territory boundaries. In display, Y perches prominently in territory with plumage fluffed out; he swivels about, calling constantly, flies up 4–5 m then glides back to perch. Y also performs horizontal display flights between perches with body feathers puffed out, wings flapping slowly. Body plumage fluffed in Y approach flight to X; rapid pursuits contrast with slow, almost hovering, ‘bumbleflight’. In perched courtship, Y bobs with quivering wings and gives hissing and twittering calls; but is silent during many displays. Y perches near X, leans forward and slowly flaps wings in front of breast, or extends them above the back with the tips touching and quivering. X solicits copulation with quivering wings and soft calls. NEST: resembles nest of E. orix but with larger entrance at side and with overhanging porch; placed in tall grass or coarse vegetation, 08–30 m above ground, usually supported by grass stems on either side of the large entrance (A). In Tanzania, nest often in Sorghum verticilliflorum (Moreau and Moreau 1938). In a drought, first nests are 3 weeks later than average, and more nests built in trees than in grass. In normal rainy season, 26 occupied nests were in Hyparrhenia rufa or Pennisetum purpureum, 2 in Acacia orfota, 1 on shoots of Cassia siamea, whereas in drought 17 nests were in Acacia orfota, 7 in Pennisetum, 4 in Hyparrhenia, 3 in millet and 10 in other vegetation (Whybrow 1950). Seldom in colonies, often either solitary or 2 YY close together. Y constructs nest, may work on more than one at a time. X lines nest with fine grass; defends immediate area against other bird species but tolerates conspecific XX on her territory. EGGS: 2–4; light blue-green, rarely with tiny black spots. SIZE: (n ¼ 18, Cameroon) 175–190  130–145; (Ethiopia) 175–180  142–150; (n ¼ 17, Zaire) 174–199  129– 144 (185  138); (n ¼ 9, Zambia) 167–190  122–141 (181  135). LAYING DATES: Mauritania, Karakoro, Aug–Sept; Senegal, July–Aug; Gambia, Aug–Nov; Mali, Aug–Sept; Sierra Leone, Sept–Jan; Liberia, Aug–Sept; Ghana, July– Aug (breeding plumage Apr–Oct); Togo, July–Aug; Benin, (breeding plumage, July–Nov); Nigeria, Aug–Oct; Cameroon, Sept–Dec; Sa˜o Tome´, Dec–Jan; (breeding plumage Dec–Apr, juvs Oct, Mar); Burkina Faso, Aug– Sept (breeding plumage Apr, July); Central African Republic, (breeding plumage June–Nov); Sudan, Aug– Oct; Ethiopia, Gato R., Apr–May, Blue Nile, Oct; Gabon, (breeding plumage Dec–May); Angola, Jan–May; Zaı¨re, Uele, Aug–Oct, Kasenga, Jan–Feb, southern region, Jan– May; Uganda, Apr–Aug; Kenya, mainly Apr–July, also Jan, Mar, Aug, Sept, Dec; Tanzania, Feb–June, Zanzibar,

ˆ i, Feb–May; May–Aug; Zambia, Dec–Apr; Malaw Zimbabwe, Jan–Apr; Mozambique, Feb–May. INCUBATION: by X only. Period 12–13 days, in captivity, 14 days. Y may chase X when she leaves nest. Y reported on a nest containing eggs (Jourdain and Shuel 1935). DEVELOPMENT AND CARE OF YOUNG: fed by X only. Nestling period 11–13 days, up to 16 days in unfavourable conditions; in captivity 14 days. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius. Y attacks Diederik Cuckoo only when it enters territory (not outside it: Verheyen 1953). Y, with Y E. nigroventris, mobbed White-browed Coucal Centropus superciliosus (Tanzania); 3 YY chased off a coucal (Liberia). 3 birds ringed in Zimbabwe recovered 44 km from ringing site. Captive Y survived >10 years. (Adlersparre 1938).

Key References Bannerman, D. A. (1949), Fuggles-Couchman, N. R. (1943), Lack, D. (1935), Moreau, R. E. and Moreau, W. M. (1938), Whybrow, H. (1950).

Euplectes gierowii

227

Euplectes gierowii Cabanis. Black Bishop. Euplecte de Gierow.

Plate 13

Euplectes gierowii Cabanis, 1880. J. Orn. 28, p. 106; Malange, Angola.

(Opp. p. 202)

Range and Status. Endemic resident. Widely separated, localized populations: Cameroon, recent records from Ijim, between Belo and Bamenda (Dowsett and DowsettLemaire 2000), Meiganga and Dimako on Adamawa Plateau (M. Languy, pers. comm.). Central African Republic, Bangui. Zaı¨re, Kidada (05
220 S, 14
320 E) and Kinshasa, in NE to Ubangi R. and grasslands from L. Albert to Rutshuru; S Sudan, near Yambio, Ikoto, and at 5
N on border with Ethiopia; SW Ethiopia, both in lowlands and in western highlands; Uganda, from Toro to Bunyoro, Mubende, Mengo, Busoga, Mbale, and at Kidepo Valley Nat. Park; W Kenya, Siaya, Kakamega, L. Victoria basin, along southern Uaso Nyiro R. on Tanzanian border; Tanzania, Mbulu Highlands, L. Manyara, Seronera R., Hanang, Shinyanga, recent record near Keragwe (Dec 1993, N. Baker, pers. comm.); Angola, Cahuta, Bailundo to Malange and Pungo Andongo. Uncommon to rare. Description. E. g. ansorgei: Cameroon to Ethiopia and W Kenya. ADULT Y (breeding): forehead and crown black; hindcrown and nape orange; mantle orange-yellow, rump and uppertail-coverts black; tail black, edged with buff in fresh plumage; lores, cheeks and ear-coverts black; chin and throat black, with narrow orange collar on upper breast linking to mantle; rest of breast, belly, thighs, vent and undertail-coverts black; wings black, edged with buff in fresh plumage; underwing sooty grey. Bill black; eyes dark brown; legs brown. ADULT Y (non-breeding): crown, nape and mantle feathers brown with heavy black central streaks; broad creamy superciliary stripe extends well behind eye; rump feathers brown with pale margins; chin and throat dull white; breast and flanks buff with dark spots rather than streaks; belly, vent and undertail-coverts dull white; thighs buffy. Wings and tail as in breeding plumage, underwing dark grey. Upper mandible dark brown, lower mandible yellowish brown. ADULT X: like non-breeding Y, but streaking on upperparts brown, not black; wing and tail feathers brown, not black, in fresh plumage, with broad buff margins, especially on upperwing-coverts; worn wings uniform dark brown. SIZE: (51 YY, 20 XX) wing, Y 75–95 (87), X 72–78 (753); tail, Y 55–71 (62), X 48–57 (531); bill, Y 180–208 (192), X 177–189 (184); tarsus, Y 252–288 (270), X 221–268 (249). WEIGHT: Y (Kenya, n ¼ 17) 232–362 (296); X (Kenya, n ¼ 7) 224–300 (270). IMMATURE: like ad. X. NESTLING: no information. E. g. gierowii Cabanis: Angola, Zaı¨re (Kidada, Kinshasa). Y in breeding plumage like ansorgei except that orange extends forwards to corner of bill; nape to mantle bright orange-yellow; a broad orange patch on throat and upper breast. SIZE: (1 Y) wing 83, tail 56, bill 171, tarsus 263; also wing, Y 79, X 82; tail, Y 50, bill Y 18, tarsus, Y 24 (Delacour and Edmond-Blanc 1933). E. g. friederichseni Fischer and Reichenow: N Tanzania, SW Kenya. Breeding Y differs from ansorgei in broad orange band around breast; rest of breast and belly black, but thighs, vent and undertail-coverts buff; hindcrown, nape and mantle bright orange; rump buff with dark centres to feathers. SIZE: Y (n ¼ 11) wing 75–83 (787), tail 53–58 (547), bill 151–173 (165), tarsus 240–267 (254). TAXONOMIC NOTE: a specimen from Tanzania, part of a shipment of birds for the cage bird trade, was illustrated in New Scientist, 22 Feb 1992, as a new species, but is clearly E. g. friederichseni in which red plumage replaced by yellow; collection site unknown. Cameroon birds look like E. g. ansorgei but have a

Euplectes gierowii

broader breast-band, and orange on head extends over the crown (only the forehead black); only sight records to date.

Field Characters. Length 14–16 cm. Largest and least common of the red and black bishops. Breeding Y distinguished from sympatric Black-winged Bishop E. hordeaceus by mainly black head, orange-yellow back, black rump; orange breast-band narrow or broad (varies with race). X and non-breeding Y blacker above than other bishops (especially Y), heavily streaked, with broad creamy supercilium; Y has rich buff face and breast, black spots (not streaks) on breast; X similar but face darker, underparts paler buff. Voice. Tape-recorded (C, KAE P). Song in flight display, long buzzy note followed by 7–10 short ones, ‘zeee-za-zaza-za-za-za-za . . . ’. In another aerial display, ‘rapid, thin, silvery notes followed by a clearer ‘‘tee-ee-ee-ee-eee’’ and then a sizzling ‘‘see zee see zee see zhe see zhe see-zhee’’, accelerating as volume increases’ (Zimmerman et al. 1996, where other variations described). General Habits. Little known; inhabits tall grassland in swampy areas; also (Tanzania) scrub and sugarcane. In Kenya, local in tall bushed grassland at 700–1600 m, and in areas with annual rainfall >1000; in NE Zaı¨re, up to 1800 m near Bogoro; perhaps up to 2000 m in W Ethiopia. Solitary or in pairs, but joins mixed flocks after breeding. Often sympatric with E. hordeaceus in E Africa.

228

PLOCEIDAE

Food. Grass seeds, ant and termite alates in stomachs. Breeding Habits. Polygynous, Y ‘controls three or four nests’ (Chapin 1954). Territorial, trees alongside elephant grass used as song perches by Y. NEST: rather coarsely woven of green grass with large entrance high up on side; lined with grass seed heads. In Uganda nests in groups in clumps of elephant grass, c. 2 m above ground, usually attached to one upright stalk and several leaf blades. One nest in Tanzania in fork of woody shrub in rank riverside vegetation, c. 1 m above ground (Bowen 1931).

EGGS: 2–4, usually 3. Glossy bright blue, immaculate or sometimes with fine black or red-brown spots. SIZE: 183– 230  143–163 (Chapin 1954). LAYING DATES: NE Zaı ¨re, May–Aug (Lippens and Wille 1976), (gonads enlarged Aug–Oct, Chapin 1932; YY in breeding plumage until Nov, Lippens and Wille 1976); Uganda, Apr–Oct, Dec (van Someren 1916); Kenya, May; Tanzania, June. Cameroon, breeding plumage and display in Sept (C. Bowden, pers. comm.). No further information.

Key References V. G. L. (1916).

Chapin, J. P. (1932, 1954), van Someren,

Plate 12

Euplectes afer (Gmelin). Yellow-crowned Bishop; Golden Bishop. Euplecte vorabe´.

(Opp. p. 155)

Loxia afra Gmelin, 1789. Syst. Nat. 1, pt 2, p. 857, Senegal. Forms a superspecies with E. diadematus. Range and Status. Endemic resident. Small populations established in Portugal (Costa et al. 1997). (Introduced to Hawaii (Long 1981), but not listed by Pratt et al. (1987).) Mauritania, uncommon near Senegal R.; Gambia, mainly along river, local on coast; Senegal, primarily on floodplain of rivers, Louga, Dakar, Kolda, Tambacounda; Guinea-Bissau, in coastal region; Guinea, Conakry, Kindia; Sierra Leone, Sherbro, Rokupr, Rotifunk; Liberia, 1 record at Gedetabo in 1929; Ivory Coast, in savanna region north of Bouake´, also Abidjan; Mali, from 17
N southwards; Burkina Faso, Arli, Bobo, Banfora, Ouagadougou, Oursi, in north near Gaoua, Ouahigouya; Ghana, coastal belt, Accra and Keta plains, northern savanna; Benin, N savannas as at Be´te´rou; Togo, coastal wetlands near Lome´ and northern savanna around Mango; Niger, in SW Gaya, Tillaberi, Ayorou, Niamey, and during rains west to MaradiTanout, Dallol Bosso, Zinder, Takie´ta, and Korama; Nigeria, widespread, commoner from great rivers northwards than towards coast; Cameroon, localized but widespread; Chad, from L. Chad S to Moundou; Central African Republic, Lobaye Pre´f., Bangui area, BaminguiBangoran Nat. Park, Manovo-Gounda-Saint Floris Nat. Park; Gabon, clearings at Makokou; Congo, local outside forest; Zaı¨re, NW, W and SE, Kasai, Sankuru, Upemba; Uganda, Karamajo and Teso regions; Sudan, locally common in Darfur and in south along Nile R. tributaries; Ethiopia, western S Ethiopia, western highlands, northeast E Ethiopia; Kenya, Baringo, Mwea, Thika, Nairobi, Turkwel delta, Kerio valley, L. Victoria basin, southern Uaso Nyiro swamps; Tanzania, Arusha, Masailand; Angola, from Luanda and Bengo south through Cuanza Norte, Malanje to coastal Benguela, Moxico, Huı´la, south and east to Cunene and Cuando Cubango; Zambia, lowlands between L. Mweru and L. Tanganyika, Kafue Flats, Southern Prov., W part of Western Prov.; Mozambique, Macia district, Umbeluzi, upper Limpopo R, Mafambisse, Cheringoma, Gorongosa Nat. Park, Tete (Clancey 1971), but apparently no recent records (Parker 1999); Namibia, Caprivi, Cunene R., northern seasonal pans; Botswana,

Euplectes afer

Okavango basin and SE; Zimbabwe, on central plateau, regular only in W; South Africa, E highveld areas, seldom near coast; Lesotho, in lowlands. Common, but often local and sporadic in occurrence. Description. E. a. afer (Gmelin) (includes ‘ladoensis’, ‘niassensis’). W Africa east to Sudan (except S), south to N Zaı¨re and NW Angola. ADULT Y (breeding): forehead, crown and nape golden yellow, nape feathers fringed with black; mantle feathers dark brown tipped with yellow, rump and uppertail-coverts yellow, some feathers fringed with black. Tail brown. Sides of head, chin, throat and belly black; yellow band across the breast, tinged with chestnut-brown in centre. Flanks, vent and undertail-coverts yellow, thighs whitish. Wing feathers and coverts brown with

228

PLOCEIDAE

Food. Grass seeds, ant and termite alates in stomachs. Breeding Habits. Polygynous, Y ‘controls three or four nests’ (Chapin 1954). Territorial, trees alongside elephant grass used as song perches by Y. NEST: rather coarsely woven of green grass with large entrance high up on side; lined with grass seed heads. In Uganda nests in groups in clumps of elephant grass, c. 2 m above ground, usually attached to one upright stalk and several leaf blades. One nest in Tanzania in fork of woody shrub in rank riverside vegetation, c. 1 m above ground (Bowen 1931).

EGGS: 2–4, usually 3. Glossy bright blue, immaculate or sometimes with fine black or red-brown spots. SIZE: 183– 230  143–163 (Chapin 1954). LAYING DATES: NE Zaı ¨re, May–Aug (Lippens and Wille 1976), (gonads enlarged Aug–Oct, Chapin 1932; YY in breeding plumage until Nov, Lippens and Wille 1976); Uganda, Apr–Oct, Dec (van Someren 1916); Kenya, May; Tanzania, June. Cameroon, breeding plumage and display in Sept (C. Bowden, pers. comm.). No further information.

Key References V. G. L. (1916).

Chapin, J. P. (1932, 1954), van Someren,

Plate 12

Euplectes afer (Gmelin). Yellow-crowned Bishop; Golden Bishop. Euplecte vorabe´.

(Opp. p. 155)

Loxia afra Gmelin, 1789. Syst. Nat. 1, pt 2, p. 857, Senegal. Forms a superspecies with E. diadematus. Range and Status. Endemic resident. Small populations established in Portugal (Costa et al. 1997). (Introduced to Hawaii (Long 1981), but not listed by Pratt et al. (1987).) Mauritania, uncommon near Senegal R.; Gambia, mainly along river, local on coast; Senegal, primarily on floodplain of rivers, Louga, Dakar, Kolda, Tambacounda; Guinea-Bissau, in coastal region; Guinea, Conakry, Kindia; Sierra Leone, Sherbro, Rokupr, Rotifunk; Liberia, 1 record at Gedetabo in 1929; Ivory Coast, in savanna region north of Bouake´, also Abidjan; Mali, from 17
N southwards; Burkina Faso, Arli, Bobo, Banfora, Ouagadougou, Oursi, in north near Gaoua, Ouahigouya; Ghana, coastal belt, Accra and Keta plains, northern savanna; Benin, N savannas as at Be´te´rou; Togo, coastal wetlands near Lome´ and northern savanna around Mango; Niger, in SW Gaya, Tillaberi, Ayorou, Niamey, and during rains west to MaradiTanout, Dallol Bosso, Zinder, Takie´ta, and Korama; Nigeria, widespread, commoner from great rivers northwards than towards coast; Cameroon, localized but widespread; Chad, from L. Chad S to Moundou; Central African Republic, Lobaye Pre´f., Bangui area, BaminguiBangoran Nat. Park, Manovo-Gounda-Saint Floris Nat. Park; Gabon, clearings at Makokou; Congo, local outside forest; Zaı¨re, NW, W and SE, Kasai, Sankuru, Upemba; Uganda, Karamajo and Teso regions; Sudan, locally common in Darfur and in south along Nile R. tributaries; Ethiopia, western S Ethiopia, western highlands, northeast E Ethiopia; Kenya, Baringo, Mwea, Thika, Nairobi, Turkwel delta, Kerio valley, L. Victoria basin, southern Uaso Nyiro swamps; Tanzania, Arusha, Masailand; Angola, from Luanda and Bengo south through Cuanza Norte, Malanje to coastal Benguela, Moxico, Huı´la, south and east to Cunene and Cuando Cubango; Zambia, lowlands between L. Mweru and L. Tanganyika, Kafue Flats, Southern Prov., W part of Western Prov.; Mozambique, Macia district, Umbeluzi, upper Limpopo R, Mafambisse, Cheringoma, Gorongosa Nat. Park, Tete (Clancey 1971), but apparently no recent records (Parker 1999); Namibia, Caprivi, Cunene R., northern seasonal pans; Botswana,

Euplectes afer

Okavango basin and SE; Zimbabwe, on central plateau, regular only in W; South Africa, E highveld areas, seldom near coast; Lesotho, in lowlands. Common, but often local and sporadic in occurrence. Description. E. a. afer (Gmelin) (includes ‘ladoensis’, ‘niassensis’). W Africa east to Sudan (except S), south to N Zaı¨re and NW Angola. ADULT Y (breeding): forehead, crown and nape golden yellow, nape feathers fringed with black; mantle feathers dark brown tipped with yellow, rump and uppertail-coverts yellow, some feathers fringed with black. Tail brown. Sides of head, chin, throat and belly black; yellow band across the breast, tinged with chestnut-brown in centre. Flanks, vent and undertail-coverts yellow, thighs whitish. Wing feathers and coverts brown with

Euplectes afer paler edges, underwing-coverts pale brown. Bill black; eyes brown; legs brown. ADULT Y (non-breeding): forehead, crown, nape, mantle, and uppertail-coverts brown, with dark brown central streaks on feathers. Chin, throat, breast and belly whitish, with dark shaft streaks on flanks and in band across breast. Thighs brown, undertail-coverts whitish; wings, tail and underwing coverts as in breeding Y. Upper mandible dark brown, lower mandible pale brown. ADULT X: like non-breeding Y. Breeding X said to be more yellowish white than non-breeding X, which is buffy and more streaked ventrally (Zimmerman et al. 1996), but that may represent differences between ads and subads, not between breeding and non-breeding ad. birds. SIZE (42 YY, 11 XX): wing, Y 53–64 (575), X 52–58 (556); tail, Y 31–38 (345), X 31–36 (335); bill, Y 128–150 (140), X 130–140 (135); tarsus, Y 170–193 (181); X 164–177 (172). WEIGHT: Y (Ghana, July–Aug, n ¼ 81) 112–205 (147), X (Ghana, July–Sept, n ¼ 41) 110–170 (142); unsexed (n ¼ 44) 122–162 (139), (Nigeria, May–June, n ¼ 19) 138–167 (152), (Senegal) 143– 160 (Schifter 1986). IMMATURE: like X; juvenile before post-juv. moult has broad buffy edges to feathers, notably wing-coverts. NESTLING: not described. E. a. strictus Ru ¨ ppell: highlands of Ethiopia. Breeding Y has narrow black band on nape, and yellow on sides of neck extends to a point behind the lores; X more buffy on underparts, which are more streaked than in other races. Larger. SIZE: wing, Y (n ¼ 31) 58–72 (664), X (n ¼ 6) 58–64 (612). WEIGHT: Y (n ¼ 8) 126–178 (155), X 111, 117, 119. E. a. taha Smith: from S Sudan, S Ethiopia and Somalia to South Africa. Breeding Y has prominent black band on nape, underparts from chin to vent black, with some yellow feathers at sides of breast level with the wings, and on flanks. SIZE: wing, Y (n ¼ 51) 59–66 (633), X (n ¼ 43) 56–63 (603). WEIGHT: Y (Zaı¨re) 10, 12, 14, (South Africa, n ¼ 11) 128–198 (165); X (Zimbabwe) 13, 152, 14, (Tanzania) 10, (South Africa, n ¼ 5) 135–166 (149); unsexed (South Africa, n ¼ 12) 140–197 (162).

Field Characters. Length 95–12 cm. Very small and short-tailed. Yellow and black breeding Y readily identified by golden crown; underparts mainly black, small yellow patch at sides of breast in southern Africa (A, B), yellow breast-band (tinged chestnut in centre) in nominate race. X and non-breeding Y have broad yellowish white supercilium, dark line through eye, and blackish earcoverts which contrast with white face; whiter below than other small bishops, more boldly streaked. Voice. Tape-recorded (20, 88, 99, 104, B, F, BRU LEM). 3-part song (southern Africa), given on perch and in courtship flight, begins with introductory gargle or rustling sound, then high-pitched, buzzy trills ‘zzzeeet, zzzeeet, zzzeeet . . . ’, ending with chipping notes, ‘rik-rik’. Bird recorded in Chad (tape 104) had different song, two 3-note phrases of dry notes, ‘wee-ta-tuck, wee-ta-tuck’ followed either by ‘wee’ or by a slow, low-pitched trill. Disputing YY give rapid ‘ha-ha-ha’ and harsh ‘zeep’. Y once landed near X, fluffed feathers and approached her with buzzing call. Contact call a short ‘chuk’; alarm similar but louder. Loud rattling call given in flight or when other birds fly over; another flight call is ‘sip-sip-sip’. Sonagrams of nominate race in Craig (1976). General Habits. Inhabits open grassy valleys, usually at low altitude, but in Ethiopia up to 1800 m; generally closely associated with marshes or seasonally flooded areas;

also in rice fields and occasional on coastal saltflats (Gambia), even in papyrus swamps (Zaı¨re). Breeds in very wet areas such as swamps and seasonally flooded pans. Occurs in small mixed flocks with other bishops and queleas, sometimes in relatively dry country. Feeds in flocks away from breeding territory. On the ground the birds move primarily by hopping. In many areas appears to be nomadic, and may appear following heavy rains, nest, then not return to the same locality for several seasons (pers. obs. and Skead 1956a).

229

230

PLOCEIDAE

There may be regular seasonal movements in Zambezi valley (Benson et al. 1971). Erratic in Zimbabwe, and birds in breeding plumage may appear and leave without nesting (Irwin 1981); in Botswana, also few records in years with poor rains (Beesley and Irving 1976). At Molai, NE Nigeria, moves southwards at start of rainy season, when birds have up to 29 g of lipid, suggesting pre-migratory fattening in preparation for a flight of up to 600 km (Jones and Ward 1977). Categorized as ‘migrant’ in Central African Republic (Carroll 1988). Anting reported in captive birds (Simmons 1961). On hot days, bathes frequently in shallow water of streams near sandbanks (Heinrich 1958). Food. Primarily grass seeds; 5 stomachs contained only grass seeds, 1 also grit. In Nigeria eats seeds of grasses Brachiaria brachylopha, B. julata, B. lata, Cymbogon sp., Dactyloctenium aegyptium, Digitaria ciliaris, Hyperthelia dissolute, Sporobolus pyramidalis; sedges Cyperus dilatatus and Mariscus alternifolius and of herbs Boerhavia erecta, B. diffusa and Tridax procumbens (Davidson 1975). Echinochloa seeds in Kenya (Cunningham-van Someren 1949). In Free State, South Africa, eats seeds mainly of Setaria flabellate and Panicum maximum but takes greater variety of seeds in winter, mostly small, including young maize (Roos and Kok 1978). Flowers of Rhynchelytrum repens (Skead 1975). Captive birds take animal food, including mealworms, termites, caterpillars, wasps and fly larvae. Breeding Habits. Polygynous, territorial; 2 XX in territory simultaneously. Nests in small colonies. Territory c. 28 m in diam. (Cheesman and Sclater 1935–1936); 2–3 nests per ha in grasses standing in lagoon (de Naurois 1969). Interspecifically territorial with Southern Red Bishop E. orix in South Africa (Farkas 1966). YY expelled Fiscal Shrike Lanius collaris and weavers Ploceus spp. from territories (Skead 1995). Y fluffs out plumage (B), resembling bumble-bee; flies out after passing X, approaching in this posture if she alights in territory. Fluffedplumage flights also directed at rival YY. In perched display on grass stalks, Y flutters wings for several s. NEST: oval ball, built of grass strips; lined with fine grasses with seedheads attached. Nest supported by vertical grass stalks; side entrance near top, often with grass seedheads of lining protruding; tops of grass stalks may be twisted down over it, but bird not known to strip leaves off surrounding vegetation. Nest well hidden in a tuft of grass, also in bushes of Rhus lancea, tamarisk bushes or rushes and sedges by flooded rice fields. Built by Y, with X

bringing lining material during incubation, but X may also contribute to nest construction in its early stages (Skead 1995). Nests are generally in waterlogged or flooded areas, 9 years period (Peach et al. 2001). Oldest birds in Malaw (Hanmer 1989). Worms under skin of some Zambian birds were larval Acanthocephala, final hosts of which are predatory animals (White 1943). Key References (1956).

Mouritz, L. B. (1913), van Someren, V. G. L.

Euplectes albonotatus (Cassin). White-winged Widowbird; White-winged Bishop. Euplecte a` Plate 14 ´paules blanches. e (Opp. p. 203) Vidua albonotata Cassin, 1848. Proc. Acad. Nat. Sci. Philadelphia, p. 65; Durban, South Africa. Range and Status. Endemic resident. Common in E and SE Africa. Introduced unsuccessfully to St Helena; and in New South Wales, Australia, in 1931 but extinct by 1968 (Long 1981). Central African Republic, Bamingui, Lobaye Pre´f. Sa˜o Tome´, probably introduced shortly before 1900. Gabon, Cap Lopez. Congo, Bateke Plateau north to Le´fini. Zaı¨re, Kinshasa to coast on west; in SE, Katanga and Marungu Mts; in E along Albertine Rift. Sudan, summer breeding in Darfur, occasional on Ugandan border. Ethiopia, W highlands south of Addis Ababa, SE highlands, southern region. Uganda, north to Mubende, Lango, Elgon and Kidepo Nat. Park. Rwanda. Burundi. Kenya, central highlands, occasional to Marsabit, Tsavo, rarely on coast. Tanzania, south to Kibondo, Tabora and Ulugurus, Songea, Mpanda and Iringa to Mkindani. Angola, central Huı´la and coastal Benguela to southern Cuanza Norte and Malanje, Cabinda. Zambia, mainly east of 26
E, rare ˆ i, throughout, especially common in in Luapula. Malaw valleys and lake shore areas. Mozambique, localized in Zambezi valley, Zumbo, Gorongosa, Limpopo valley south to Maputo district. Namibia, restricted to Caprivi. Botswana, Okavango, Chobe R. and local in SW: Zimbabwe, widespread, with some gaps in E. South Africa,

Euplectes albonotatus

?

Euplectes albonotatus 1995). Nest attached to upright stems on either side of entrance, 05–17 m above ground; some of surrounding grass may be woven into outer shell (A). Tops of many herbs are nipped off within 60 cm radius of nest. Frame built by Y, starting with entrance ring, X adding to it and lining nest, even during incubation. One nest 178 high, 127 wide. Sited in grass or small shrubs; often in standing wheat along streamlines in W Cape (Vincent 1949). Captive birds made nest of dry grass in clump of pampas grass. EGGS: 2–4. Very variable: bluish white, heavily speckled with reddish brown; light brown tinged with greenish, heavily blotched and streaked all over with dark brown and grey; olive-brown with ash-brown spots; cream or very pale greenish, covered evenly in bold streaky freckling of olive, sepia and shades of grey; pale bluish grey, with numerous blotches and spots of reddish brown and olivebrown. SIZE: (capensis, n ¼ 25) 199–234  140–160, (213  150), (approximans, n ¼ 3) (207  148), (crassirostris, n ¼ 27) (196  139), (phoenicomerus, n ¼ 5) 200–215  150–137 (207  145). LAYING DATES: Cameroon, Nov (breeding Sept–Jan); Nigeria, (breeding plumage Sept); Ethiopia July–Oct (prenuptial moult in May, X feeding fledged young Oct); Zaı¨re, Uele, July–Nov (prenuptial moult follows rains in Apr: Chapin 1932), Ruwenzori, (breeding plumage in Jan), Itombwe, Jan–Mar, Katanga, Dec, Lubumbashi, Feb–Apr; Uganda, May–Aug, Oct–Nov; Kenya, primarily Mar–June,

235

also in July, Nov–Feb; Tanzania, Apr–May; Angola, Mar– ˆ i, May; Zambia, Dec–Mar (breeding plumage Nov); Malaw Feb; Zimbabwe, Dec–Mar, rarely Apr; Mozambique, Dec– Feb (breeding condition from Dec to May); South Africa, SW Cape, Aug–Dec, Eastern Cape and KwaZulu-Natal, Dec–Mar, Transvaal, Nov–Mar. INCUBATION: by X only; X leaving nest is regularly chivvied by Y. Period: 15–16 days (Mouritz 1913) or 13–14 days; in captivity, 13 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only, by regurgitation. Period: 16 days, rarely 20; in captivity, young fed by X with mealworms and left nest after 14 days. BREEDING SUCCESS/SURVIVAL: X liable to desert if nest interfered with before laying. Many eggs and young taken by predators. Y pursued Diederik Cuckoo Chrysococcyx caprius although no records of brood parasitism (Brooke 1953). In Zimbabwe, 5 birds recaptured at ringing site after 25–4 years (Irwin 1981); in Zaı¨re of 23 birds ringed, 9 were recaptured at the same site, some several times, up to 20 ˆ i annual months after ringing (Bont et al. 1965); in Malaw survival rate estimated at 54% from recaptures over 17-year ˆ i >9 years period (Peach et al. 2001). Oldest birds in Malaw (Hanmer 1989). Worms under skin of some Zambian birds were larval Acanthocephala, final hosts of which are predatory animals (White 1943). Key References (1956).

Mouritz, L. B. (1913), van Someren, V. G. L.

Euplectes albonotatus (Cassin). White-winged Widowbird; White-winged Bishop. Euplecte a` Plate 14 ´paules blanches. e (Opp. p. 203) Vidua albonotata Cassin, 1848. Proc. Acad. Nat. Sci. Philadelphia, p. 65; Durban, South Africa. Range and Status. Endemic resident. Common in E and SE Africa. Introduced unsuccessfully to St Helena; and in New South Wales, Australia, in 1931 but extinct by 1968 (Long 1981). Central African Republic, Bamingui, Lobaye Pre´f. Sa˜o Tome´, probably introduced shortly before 1900. Gabon, Cap Lopez. Congo, Bateke Plateau north to Le´fini. Zaı¨re, Kinshasa to coast on west; in SE, Katanga and Marungu Mts; in E along Albertine Rift. Sudan, summer breeding in Darfur, occasional on Ugandan border. Ethiopia, W highlands south of Addis Ababa, SE highlands, southern region. Uganda, north to Mubende, Lango, Elgon and Kidepo Nat. Park. Rwanda. Burundi. Kenya, central highlands, occasional to Marsabit, Tsavo, rarely on coast. Tanzania, south to Kibondo, Tabora and Ulugurus, Songea, Mpanda and Iringa to Mkindani. Angola, central Huı´la and coastal Benguela to southern Cuanza Norte and Malanje, Cabinda. Zambia, mainly east of 26
E, rare ˆ i, throughout, especially common in in Luapula. Malaw valleys and lake shore areas. Mozambique, localized in Zambezi valley, Zumbo, Gorongosa, Limpopo valley south to Maputo district. Namibia, restricted to Caprivi. Botswana, Okavango, Chobe R. and local in SW: Zimbabwe, widespread, with some gaps in E. South Africa,

Euplectes albonotatus

?

236

PLOCEIDAE

throughout Northern Province except arid Limpopo valley; Mpumalanga, Gauteng, KwaZulu-Natal, NE Free State. Swaziland, throughout. Density of up to 20 birds per km on transects near Gaborone, Botswana (Tyler 2001) and of 77, 108, 110 and 193 birds per 100 ha in Acacia savannas, Swaziland (Monadjem 2002).

Voice. Tape-recorded (88, 99, 104, B, GREG, LEM, McVIC, PAY). Sequence of song components: 3 dry paper-tearing noises, brief dry rattle, 1–3 nasal ‘zem’ notes, high thin sewing machine trill lasting 5–10 s, ‘zzhannzzha-zzhann’ (or ‘squee-squi-squeege’), twirrr, zem-zemzem, tisisisisisisisi . . . ’. Twittering contact calls; in flight and display, ‘squi-squi-squeege-squeege’.

Description. E. a. albonotatus (Cassin): S Tanzania (intergrading with eques in Iringa and Dodoma districts) to NE Botswana, and South Africa. ADULT Y (breeding): wholly black except for wings; tail elongated, rounded and black; primary feathers black with white bases, secondaries black with narrow buff edges and white bases, lesser coverts golden yellow, median coverts black with buffy edges, greater coverts white; leading edge of wing yellow, underwing white. Bill dark blue-grey; eyes brown; legs dark blackish brown. (a captive Y was wholly black: van Someren 1918). ADULT Y (non-breeding): forehead to uppertail-coverts brown with dark central streaks; tail short, brown with paler edges; broad yellowish white superciliary stripe, yellow patch at base of bill; lores, cheeks and ear-coverts buffy brown; chin and throat white; breast, flanks and thighs buffy brown; belly and undertail-coverts white; wings as in breeding Y. Bill blue-black, often blotchy; eyes brown; legs light brown. SUB-ADULT Y: resembles non-breeding Y, but much whiter ventrally; wing feathers brown without white bases, lesser coverts brown with yellow fringes. Upper mandible brown, lower flesh-coloured; eyes brown; legs brown. ADULT X: like sub-ad. Y but smaller. (1 white with yellow and brown streaking; Dowsett 1974). SIZE (32 YY, 22 XX): wing, Y 71–81 (760), X 63–71 (661); tail, breeding Y (n ¼ 28) 77–93 (845), non-breeding Y (n ¼ 10) 47–58 (527), X 41–51 (450); bill, Y 152–166 (158), X 140–157 (148); tarsus, Y 212–236 (220), X 185–214 (199). Can be sexed by winglength (Craig and Manson 1981). WEIGHT: (KwaZulu-Natal) Y, (n ¼ 31) 200–265 (232), X (n ¼ 21) 160–240 (191). IMMATURE: resembles X; some have yellow wash on underparts. Y acquires ad. wing pattern at about 13 months, during second complete moult (Lynes 1924–1925). NESTLING: hatches naked, skin pink. E. a. asymmetrurus (Reichenow): Sa˜o Tome´, W Gabon to W Angola. Tail of breeding Y much larger. SIZE: wing, Y (n ¼ 14) 70–79 (734); tail, breeding Y (n ¼ 12) 117–152 (132), nonbreeding Y (n ¼ 14) 51–69 (579). E. a. eques (Hartlaub): Central African Republic, Sudan and Ethiopia to S Tanzania where intergrades with nominate race. Breeding Y has cinnamon-brown (not yellow) on epaulet, X and sub-ad. Y with lesser coverts edged with cinnamon. SIZE: wing, Y (n ¼ 26) 73–85 (771); tail, breeding Y (n ¼ 24) 56–99 (835), nonbreeding Y (n ¼ 6) 41–58 (525). WEIGHT: (Kenya) Y (n ¼ 10) 187–237 (221), X 165, 215.

General Habits. Inhabits tall bushed grassland, generally below 2000 m. On Sa˜o Tome´, common in northern savannas near farmland and towns. Favours rather fine grass, knee to waist high in E Zaı¨re, and not usually found alongside Red-collared Widowbird E. ardens (Chapin 1954). Large flocks of non-breeding birds mix with bishops E. hordeaceus, E. ardens, E. capensis, E. orix, E. franciscanus and with Red-headed Queleas Q. erythrops; in S Zaı¨re, moves from lake shores and flooded savanna to cultivated areas and, after breeding, to permanent swamps. Up to 600 birds roost gregariously in dry season. Catches termites on ground and hawks them in air, returning to perch to eat them. Some local movements; seasonal visitor to Darfur, absent in dry season; in Kenya, influx into RuarakaKahawa-Ruiru region in May–Aug breeding season; wet season visitor in Tsavo region.

Field Characters. Length, breeding Y 16 cm, non-breeding Y 14 cm, X 12 cm. A small widowbird with fan-shaped tail intermediate in length between bishop and typical widow. Breeding Y black with white wing-patch and wing edgings, white underwing; southern African birds have yellow wing shoulder but do not overlap ‘Yellow-shouldered Widowbird’ E. macroura macrocerca of E Africa; in E Africa epaulet is cinnamon-rufous. Streaky non-breeding Y readily identified in flight by longish tail, white and yellow/rufous wing patches, white underwing; X is whiter below than other Euplectes spp., breast unstreaked, has pronounced white eyebrow, bend of wing yellow or cinnamon, underwing white; buffy white margins to primary coverts show as pale flash in flight (Maclean 1993).

Food. Seeds of Brachiaria bizantha, Chloris virgata, Hyparrhenia spp., Pennisetum sp., Rhynchelytrum repens, Rottboellia exaltata, Setaria sphacelata, Urochloa pullulans. Flocks in millet fields in Ethiopia. Nectar of Aloe marlothii. Insects, including termite alates; in captivity, animal food includes mealworms, earwigs, termites. Breeding Habits. Polygynous, territorial; defends area of up to 1900 m2; sometimes YY 50–100 m apart, with territories of 600–1200 m2. Y perches on grass stems, also higher sites such as dead trees; fans and lowers tail in display, sings with erect posture and ruffled nape feathers, beak horizontal. In S Zaı¨re, Y sets up territory in bushy savanna with Pterocarpus and Acacia trees. On Sa˜o Tome´ nests in grasslands, often near bishops E. hordeaceus and E. aureus; interspecifically territorial with them (de Naurois 1983); in Zimbabwe, Y chased Y E. orix and defended territory against E. ardens. In display, flies slowly but with rapid wing-beats, tail spread conspicuously, and epaulets obvious. If X lands in territory, Y drops into grass, approaches her with beak depressed, wings slightly hunched, epaulets and ruff on nape erect, tail spread and depressed; bounces up and down calling ‘squee-squeesqueege-squeege’. In nest invitation display, Y flies to nest frame, hangs at entrance with head raised, beak pointed upward, tail spread and wings slightly opened and quivered rapidly, and calls as X approaches. X solicits copulation by raising wings slightly, quivering tail and giving series of thin piping calls; Y mounts briefly with much wing-fluttering. Y remain close to nest in presence of intruder, once chased off returning X (Vincent 1936). Not double-brooded (in one intensive study, Zimbabwe). NEST: oval with large side entrance; framework of dry and semi-green grass, supported by upright grass stems incorporated into weaving on either side of entrance; built

Euplectes macroura by Y. Within this a stouter weaving of finer dry grass, usually Sporobolus, contributed by X. No dome of living grass; very like nest of Black-winged Bishop E. hordeaceus and Red-collared Widowbird E. ardens. Size, 130 high, 70 deep; some nests have small porch of projecting grassheads. Generally sited below 15 m. Eggs may be laid before nest is lined. EGGS: 2–4, (26, n ¼ 20, Zimbabwe). Pale greenish blue, marked closely all over with blurred or clear olive-green speckles. SIZE: (n ¼ 6, Tanzania). 175–198  140–148 (187  143); (n ¼ 7, Kenya) av. 188  141; (Darfur, Sudan) av. 185  132. LAYING DATES: Sa ˜o Tome´, (Dec–Apr); Sudan, Sept–Oct (Aug–Oct); Ethiopia, June; Zaı¨re, Feb–Apr in Katanga (Mar–May); Rwanda, Feb–Apr; Kenya, Mar–June peak, also Sept, Dec–Feb; Tanzania, Dec–Mar; Angola, Jan–Apr (pre-nuptial moult Nov, breeding YY Jan, juvs Jan); ˆ i, Jan–Apr; Botswana, Dec–Feb; Zambia, Jan–Mar; Malaw

237

Zimbabwe, Dec–Apr; Mozambique, Mar; South Africa, Oct–Apr. INCUBATION: by X only. Period 12–14 days; in captivity 12–15 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only, primarily by regurgitation; nestling period 11–14 days, 11– 18 days in captivity; young independent at 36 days. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius; of 167 nests only 18% parasitized (Payne and Payne 1967). Eggs and newlyhatched chicks eaten by multimammate mouse Mastomys natalensis. Of 21 nests in Zimbabwe, 14 destroyed by people (5) and predators (9). Oldest bird recaptured in Zimbabwe, >7 years (Manson 1982). Key References Hornby, H. E. (1967, 1970, 1977), Moreau, R. E. and Moreau, W. M. (1937).

Euplectes macroura (Gmelin). Yellow-mantled Widowbird; Yellow-backed Widow. Euplecte `a Plate 14 dos d’or. (Opp. p. 203) Loxia macroura Gmelin, 1789. Syst. Nat., 1, pt 2, p. 845; Dahomey (¼ Benin). Range and Status. Endemic resident. Common at least locally; in parts of W Africa commoner than bishops. Senegal, Dakar, Basse Casamance, Ziguinchor. Gambia, in S. Guinea-Bissau. Guinea, Kindia, Macenta. Sierra Leone, Bo, Tingi Mts, Bendugu, Kakunda, Saiama, Kurrabondo, Nerekoro, Kankordu, Kwendu, Balemu, Rokupr. Liberia, locally common outside forest. Ivory Coast, abundant from Lamto northward. Burkina Faso, widespread in SW, north as far as Ouagadougou. Mali, rare in Boucle de Baoule´ reserve, old record from San. Ghana and Togo, widespread except near coast. Benin, widespread, especially in savanna. Nigeria, near coast from Lagos to Calabar, inland to Zaria, Zonkwa, Jos Plateau and Birnin Kudu. Niger, occasional in SW, near Niger R. and east to Dosso. Chad, local in SW. Cameroon, widespread south of Benue Plain. Central African Republic, Manovo-Gounda-Saint Floris Nat. Park, Lobaye Pre´f., Bangui, Haute Sangha Pre´f., Bamingui-Bangoran Nat. Park, Dzanga Reserves. Sudan, SW Nile basin, E highlands. Ethiopia, W highlands, NE, SE highlands, S region, rift valley. Eritrea, in highlands. Gabon, Lope´ to Tchibanga. Congo, Djambala, Gamboma, Odzala. Zaı¨re, savanna from Duma to Mahagi on L. Albert; south of forest, from coast to Manyema and Katanga, east of L. Tanganyika south to Ufipa and Marungu. Rwanda. Uganda, north to West Nile, Lango, Acholi, Teso and SW Karamoja. W Kenya and N Tanzania: see NOTE, below. Elsewhere in Tanzania, Ukerewe I., Kibondo and Tabora to Ufipa Plateau, Njombe highlands; absent from central plateau, coastal areas and W-central miombo. Angola, Cabinda south to Cuanza Norte, Malanje and Cuanza Sul, east to Lunda Norte, Lunda Sul and Moxico. Zambia, throughout, except for parts of Northern and Western Provs, Luangwa and ˆ i, west of rift, from Dedza Middle Zambesi valleys. Malaw

Euplectes macroura

northwards. Zimbabwe, local on Mashonaland plateau. N Mozambique, near Vila Coutinho, Zumbo. At Lamto, Ivory Coast, density of 64 birds per 50 ha in dry season, 75 per 50 ha in rainy season (Thiollay 1973). Description. E. m. macroura (Gmelin) (includes ‘soror’, ‘intermedia’): range of species except for Ethiopian highlands and Ukerewe I., L. Victoria. ADULT Y (breeding): black, except for golden yellow mantle and yellow patch on wing. Nape feathers

Euplectes macroura by Y. Within this a stouter weaving of finer dry grass, usually Sporobolus, contributed by X. No dome of living grass; very like nest of Black-winged Bishop E. hordeaceus and Red-collared Widowbird E. ardens. Size, 130 high, 70 deep; some nests have small porch of projecting grassheads. Generally sited below 15 m. Eggs may be laid before nest is lined. EGGS: 2–4, (26, n ¼ 20, Zimbabwe). Pale greenish blue, marked closely all over with blurred or clear olive-green speckles. SIZE: (n ¼ 6, Tanzania). 175–198  140–148 (187  143); (n ¼ 7, Kenya) av. 188  141; (Darfur, Sudan) av. 185  132. LAYING DATES: Sa ˜o Tome´, (Dec–Apr); Sudan, Sept–Oct (Aug–Oct); Ethiopia, June; Zaı¨re, Feb–Apr in Katanga (Mar–May); Rwanda, Feb–Apr; Kenya, Mar–June peak, also Sept, Dec–Feb; Tanzania, Dec–Mar; Angola, Jan–Apr (pre-nuptial moult Nov, breeding YY Jan, juvs Jan); ˆ i, Jan–Apr; Botswana, Dec–Feb; Zambia, Jan–Mar; Malaw

237

Zimbabwe, Dec–Apr; Mozambique, Mar; South Africa, Oct–Apr. INCUBATION: by X only. Period 12–14 days; in captivity 12–15 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only, primarily by regurgitation; nestling period 11–14 days, 11– 18 days in captivity; young independent at 36 days. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius; of 167 nests only 18% parasitized (Payne and Payne 1967). Eggs and newlyhatched chicks eaten by multimammate mouse Mastomys natalensis. Of 21 nests in Zimbabwe, 14 destroyed by people (5) and predators (9). Oldest bird recaptured in Zimbabwe, >7 years (Manson 1982). Key References Hornby, H. E. (1967, 1970, 1977), Moreau, R. E. and Moreau, W. M. (1937).

Euplectes macroura (Gmelin). Yellow-mantled Widowbird; Yellow-backed Widow. Euplecte `a Plate 14 dos d’or. (Opp. p. 203) Loxia macroura Gmelin, 1789. Syst. Nat., 1, pt 2, p. 845; Dahomey (¼ Benin). Range and Status. Endemic resident. Common at least locally; in parts of W Africa commoner than bishops. Senegal, Dakar, Basse Casamance, Ziguinchor. Gambia, in S. Guinea-Bissau. Guinea, Kindia, Macenta. Sierra Leone, Bo, Tingi Mts, Bendugu, Kakunda, Saiama, Kurrabondo, Nerekoro, Kankordu, Kwendu, Balemu, Rokupr. Liberia, locally common outside forest. Ivory Coast, abundant from Lamto northward. Burkina Faso, widespread in SW, north as far as Ouagadougou. Mali, rare in Boucle de Baoule´ reserve, old record from San. Ghana and Togo, widespread except near coast. Benin, widespread, especially in savanna. Nigeria, near coast from Lagos to Calabar, inland to Zaria, Zonkwa, Jos Plateau and Birnin Kudu. Niger, occasional in SW, near Niger R. and east to Dosso. Chad, local in SW. Cameroon, widespread south of Benue Plain. Central African Republic, Manovo-Gounda-Saint Floris Nat. Park, Lobaye Pre´f., Bangui, Haute Sangha Pre´f., Bamingui-Bangoran Nat. Park, Dzanga Reserves. Sudan, SW Nile basin, E highlands. Ethiopia, W highlands, NE, SE highlands, S region, rift valley. Eritrea, in highlands. Gabon, Lope´ to Tchibanga. Congo, Djambala, Gamboma, Odzala. Zaı¨re, savanna from Duma to Mahagi on L. Albert; south of forest, from coast to Manyema and Katanga, east of L. Tanganyika south to Ufipa and Marungu. Rwanda. Uganda, north to West Nile, Lango, Acholi, Teso and SW Karamoja. W Kenya and N Tanzania: see NOTE, below. Elsewhere in Tanzania, Ukerewe I., Kibondo and Tabora to Ufipa Plateau, Njombe highlands; absent from central plateau, coastal areas and W-central miombo. Angola, Cabinda south to Cuanza Norte, Malanje and Cuanza Sul, east to Lunda Norte, Lunda Sul and Moxico. Zambia, throughout, except for parts of Northern and Western Provs, Luangwa and ˆ i, west of rift, from Dedza Middle Zambesi valleys. Malaw

Euplectes macroura

northwards. Zimbabwe, local on Mashonaland plateau. N Mozambique, near Vila Coutinho, Zumbo. At Lamto, Ivory Coast, density of 64 birds per 50 ha in dry season, 75 per 50 ha in rainy season (Thiollay 1973). Description. E. m. macroura (Gmelin) (includes ‘soror’, ‘intermedia’): range of species except for Ethiopian highlands and Ukerewe I., L. Victoria. ADULT Y (breeding): black, except for golden yellow mantle and yellow patch on wing. Nape feathers

238

PLOCEIDAE

long and square-tipped, forming a ruff; some concealed white feathers in midline of breast; tail long, black, graduated. Wings black, lesser coverts yellow, median and secondary coverts, secondaries and tertials buff-edged. Bill black except for bluegrey tip to lower mandible; eyes dark brown; legs blackish. ADULT Y (non-breeding): forehead, crown, nape and mantle brown with dark central streaks on feathers; rump and uppertailcoverts light brown, tail short, brown; dull white superciliary stripe; lores, cheeks and ear-coverts light brown; chin whitish, throat and breast light brown with faint central streaks; belly white; thighs and undertail-coverts light brown; wings brown, underwing buff. Bill brown; eyes brown; legs brown. SUB-ADULT Y: like non-breeding Y, but wing feathers brown with pale edges, yellow margins to lesser coverts; yellowish supercilium. Bill brown, lower mandible much paler. ADULT X: like sub-ad. Y, but generally paler ventrally. SIZE (50 YY, 32 sub-ad. Y, 59 XX): wing, Y 76–87 (803), sub-ad. Y 71–79 (751), X 61–72 (676); tail, Y (breeding) 85–118 (107), (non-breeding, n ¼ 16) 57–69 (625), sub-ad. Y 50–67 (586) X 46–67 (508); bill, Y 157–184 (169), sub-ad. Y 153–179 (166), X 145–168 (156); tarsus, Y 224–248 (235), sub-ad. Y 210–235 (232), X 197–221 (210). WEIGHT: Y (Ghana, July–Sept, n ¼ 8) 210–240 (220), (Nigeria, Nov–July, n ¼ 26) 223–285 (251), (Zaı¨re, n ¼ 7) 230–280 (251), (Kenya, n ¼ 46) (av. 238); sub-ad. Y (Kenya, n ¼ 39) (av. 230); X (Ghana, July–Sept, n ¼ 8) 160–220 (197), (Nigeria, Nov–May, n ¼ 18) 171–202 (187), (Nigeria, June–July, n ¼ 4) 187–226 (208), (Kenya, n ¼ 99) (av. 193); unsexed, non-breeding birds (Ghana, n ¼ 6) 160–188 (174), (Nigeria, n ¼ 23) 179–265 (225). IMMATURE: like ad. X but underparts often washed with yellow; young Y more heavily streaked on flanks and belly. NESTLING: not described. E. m. conradsi (Berger): Ukerewe I., L. Victoria. The only specimen has yellow back and very long tail; current status of population not known. SIZE (1 Y): wing 87, tail 143, bill 169, tarsus 235. E. m. macrocerca (Lichtenstein) ‘Yellow-shouldered Widowbird’: Ethiopian highlands; Uganda and W Kenya (see below). Breeding Y larger and longer-tailed than other races, lacks yellow on mantle. Non-breeding Y and X have very dark streaked upperparts, dark tips to wing feathers (pale in other populations). SIZE (29 YY, 16 XX): wing, Y 80–93 (880), X 70–77 (731); tail, Y (breeding, n ¼ 17) 121–167 (138), Y (non-breeding, n ¼ 11) 57– 71 (646), X 51–59 (541); bill, Y 163–181 (171), X 146–168 (159); tarsus, Y 239–260 (248), X 205–232 (221). TAXONOMIC NOTE: macrocerca is locally common in W Kenya from Kapenguria, Kitale, Malaba, Bungoma and Webuye Districts to Busia, Mumias, Ng’iya, Maseno and Ahero; 50 km to south, nominate macroura is local and uncommon from Mara Game Res. and Loita Hills, and across border into Serengeti Nat. Park and Loliondo district, Tanzania (Zimmerman et al. 1996). Black-mantled macrocerca occurs spasmodically in this range of yellow–mantled macroura and in Uganda, YY of the 2 sometimes together (XX are indistinguishable in the field); intermediates occur. ‘Exact ranges and overlap between the two’ remains unclear (Stevenson and Fanshawe 2002) and so does their taxonomic relationship.

Field Characters. Length, breeding Y 18–22 cm, X 13–14 cm. Breeding Y black with fairly long rounded tail, yellow mantle and wing shoulder. In parts of E Africa yellow only on shoulder (‘Yellow-shouldered Widowbird’ E. m. macrocerca), but these do not meet yellow-shouldered races of White-winged Widowbird E. albonotatus. Yellow Bishop E. capensis has short tail, yellow rump but black mantle. Streaked non-breeding Y has dark brown wings, small yellow epaulets, broad, not pointed rectrices, yellowish wash on underparts, underwing buff, tail rather longer

than bishops in similar plumage; X lacks epaulets but has yellow margins to lesser and median coverts. Voice. Tape-recorded (86, 99, 104, B, C, PAY, STJ). Song from perch a low grating noise like paper being torn, ‘zzher-tsi-zzher-zzhyer’; given alone or preceded by repeated ‘tzip . . . tzip . . . ’ or ‘tsee-piti, tsee-piti . . . ’. Another song a dry rattle followed by a shrill trill, possibly the ‘high-pitched twittering song often alternated with hissing trills in boundary displays’ of Savalli (1993a). Mournful whistling call in flight; Y on perch calls ‘tsinktsink’; other calls: buzzy ‘tzik’ or ‘dzip’, nasal ‘bew’, highpitched ‘tswip’ and ‘tsweep’. General Habits. Inhabits moist grassland, marshy areas with scrub and trees, also ricefields in W Africa. Most abundant in steppes dominated by Loudetia simplex in S Zaı¨re. In Bamenda area of Cameroon up to 1700 m, in Kenya, at 1000–1800 m, in areas with >1000 mm rainfall; at 1000–2100 m in Ethiopia; mostly above 1200 m in Zimbabwe. Forms flocks of >100 birds after breeding, joining with flocks of E. hartlaubi (Savalli 1993a) and in Ethiopia with E. ardens laticauda, sometimes damaging crops; also joins large mixed flocks of weavers. Roosts in papyrus or reeds in swamps, and in thickets of Acanthus pubescens. Y joins communal roost during breeding season and does not remain on territory. Small numbers of birds join mixed-species flocks in savanna woodland in Ghana, and breeding birds may join flocks while they are moving within their territories (Greig-Smith 1978). Smaller feeding groups of about 20 birds form in dry conditions or after grass has been burnt; occurrence clearly seasonal in some parts of Ghana. Hawks winged termites, returning to perch to eat them (Brooke 1970). Both sexes have partial prenuptial moult of body plumage and tail, in Mar–Apr in Kenya. Complete moult after breeding; non-breeding (sub-ad.) may start earlier than ads; YY still in breeding plumage in Sept, all in eclipse plumage by Nov (Savalli 1993b). Anting noted in captive birds (Poulsen 1956). Food. Seeds, particularly of sedges Cyperaceae; grass seeds and a few insects including winged termites. In Ivory Coast savanna, diet estimated at 80% seeds, 20% arthropods (Thiollay 1973). Breeding Habits. Polygynous, with up to 5 XX per Y, and highly territorial. Where very common, breeding is almost colonial (Thiollay 1985). Territory-holder displaces any other Y which approaches his area. However, appears not to interact with Hartlaub’s Marsh Widowbird E. hartlaubi where territories overlap (Savalli 1993a). Territory size (n ¼ 42, Kenya) 017–27 ha (095); in Benin, 5 YY in area of 4 ha. On approach of intruder, Y drops down to grass near nest. XX remain in flocks when Y first take up its territory. Sex ratio, Kenya, at parity (but almost half YY immature: Savalli 1994a). Y often perches conspicuously, calling with jerky movements of wings, tail slightly spread. When he ‘sizzles’, nape feathers are ruffled and body drawn up with legs stretched; he sometimes flies up vertically, with ruffled nape. In territorial flight Y jerks or flicks body

Euplectes ardens and wags tail up and down; on landing, he raises and spreads tail. During perched courtship, Y approaches X with plumage ruffled, body stretched upwards, and ‘sizzling’. When body plumage is ruffled, silvery white under-feathers are exposed on lower breast and belly. Experimental manipulation of Y tail-length showed that YY with shortened tails at start of breeding season lost their territories; but changing tail-length later had no apparent effect in terms of success in attracting XX (Savalli 1994b). YY in Kenya each had up to 5 nests occupied by X simultaneously on territory; number of nest frames made by Y was best predictor of success in attracting XX (Savalli 1994c); up to 27 frames on 1 territory. No evidence of second broads (Kenya, Savalli 1997). NEST: shape like nests of congeners but larger. Built by Y; blades of living grass drawn over and woven in; no porch over entrance. X continues to add lining of dry Sporobolus grass during incubation. Nest 140 deep, 90 wide, entrance diam. 45. Nests are in damp or waterlogged areas, 15–60 cm above ground, in grass no more than 90 cm tall (A). In Kenya, 80 nests in Loudetia kagerensis (72%), Hyparrhenia diplandra (10%), H. filipendula (8%), Setaria sphacelata (8%) and Digitaria mailandia (2%). EGGS: 2–3, (n ¼ 28, Kenya, av. 19). Greenish blue with irregular streaks and spots of grey, or evenly-spaced grey or brown spots; often peppered. SIZE: (n ¼ 13, Zaı¨re) 169–210  135–147 (192  141). LAYING DATES: (in parentheses, YY in breeding plumage): Sierra Leone, Oct; Liberia, Sept–Oct (July–Nov); Ghana, June–Aug (Apr–Oct); Togo, (prenuptial moult, June); Benin, June–Nov; Nigeria, July–Oct (May–June to Nov); Burkina Faso, Aug–Sept (July–Nov); Nigeria, July– Oct; Cameroon, Sept–Nov (July–Dec); Central African Republic (June–Nov); Congo, Oct (prenuptial moult Oct–Jan); Gabon, Mar–May; Sudan, Aug–Oct; Ethiopia, July–Aug, Oct (July–Oct); Uganda, June–Aug; Kenya, May–Sept; Tanzania, June; N Zaı¨re, July–Oct; S Zaı¨re, Jan–Apr; Angola, Mar–May; Zambia, Dec–Mar (Dec); ˆ i, Jan–Feb; Mozambique, (Mar); Zimbabwe, Malaw Dec–Mar. INCUBATION: by X only. In captivity, period 12–14 days (Hermann 1978).

239

DEVELOPMENT AND CARE OF YOUNG: young fed by X only; captive young initially fed insects, later bread, milk, seeds. Nestling period: 15 days (Hermann 1978). Y once fed fledgling (Savalli 1997). BREEDING SUCCESS/SURVIVAL: of 28 nests, 27% fledged 1 or more young; 24% of eggs produced flying young. Most failures are during egg stage; nests destroyed by cattle, people cutting grass, or nests abandoned. In captivity, can hybridize with E. ardens and E. axillaris (Colahan and Craig 1981, Hopkinson 1938).

Key References

Savalli, U. M. (1993a, b, 1994a–c, 1997).

Euplectes ardens (Boddaert). Red-collared Widowbird. Euplecte veuve-noire.

Plate 14

Fringilla ardens Boddaert, 1783. Tabl. Pl. enlum., p. 39; Cape of Good Hope.

(Opp. p. 203)

Range and Status. Endemic resident. Common to abundant in SE Africa, more local and less common elsewhere. Gambia, several old records but none since 1965. Senegal, 1 recent record, Niokolo-Koba Nat. Park, 1992 (Sauvage and Rodwell 1998). Mali, Niger R. valley at Kangaba and Koure´male´, and Mandingo Mts near Sagabari. Guinea, Macenta on inland plateau. Sierra Leone, local along rivers, near Tingi Mts. Liberia, locally common in N. Ivory Coast, Beoumi-Bouake´ to Comoe´ and Korhogo, Goue´ssesso, Marahoue´ Nat. Park. Ghana, 1, Cape Coast, 1977 (Grimes 1987). Togo, rare: Misaho ¨he, Pagala, Djodji.

Benin, no recent records. Nigeria, locally not uncommon on Jos, Obudu and Mambilla Plateaux. Niger, 1988 record in Parc du W requires confirmation. Cameroon, Wum, Tchabal Mbalo, Adamawa Plateau, north to about 7
N. Central African Republic, Bamingui, Lobaye, BaminguiBangoran Nat. Park. Sudan, along S border, Torit, Yei and Boma hills. Ethiopia, W highlands, Rift Valley, S and SE regions. Eritrea, uncommon in highlands. Gabon, Tchibanga. Congo, in S on R. Congo. Zaı¨re, savannas of Uele and Ituri districts, lower Congo valley and Kasai, S savannas and E grasslands. Rwanda. Burundi. Uganda,

Euplectes ardens and wags tail up and down; on landing, he raises and spreads tail. During perched courtship, Y approaches X with plumage ruffled, body stretched upwards, and ‘sizzling’. When body plumage is ruffled, silvery white under-feathers are exposed on lower breast and belly. Experimental manipulation of Y tail-length showed that YY with shortened tails at start of breeding season lost their territories; but changing tail-length later had no apparent effect in terms of success in attracting XX (Savalli 1994b). YY in Kenya each had up to 5 nests occupied by X simultaneously on territory; number of nest frames made by Y was best predictor of success in attracting XX (Savalli 1994c); up to 27 frames on 1 territory. No evidence of second broads (Kenya, Savalli 1997). NEST: shape like nests of congeners but larger. Built by Y; blades of living grass drawn over and woven in; no porch over entrance. X continues to add lining of dry Sporobolus grass during incubation. Nest 140 deep, 90 wide, entrance diam. 45. Nests are in damp or waterlogged areas, 15–60 cm above ground, in grass no more than 90 cm tall (A). In Kenya, 80 nests in Loudetia kagerensis (72%), Hyparrhenia diplandra (10%), H. filipendula (8%), Setaria sphacelata (8%) and Digitaria mailandia (2%). EGGS: 2–3, (n ¼ 28, Kenya, av. 19). Greenish blue with irregular streaks and spots of grey, or evenly-spaced grey or brown spots; often peppered. SIZE: (n ¼ 13, Zaı¨re) 169–210  135–147 (192  141). LAYING DATES: (in parentheses, YY in breeding plumage): Sierra Leone, Oct; Liberia, Sept–Oct (July–Nov); Ghana, June–Aug (Apr–Oct); Togo, (prenuptial moult, June); Benin, June–Nov; Nigeria, July–Oct (May–June to Nov); Burkina Faso, Aug–Sept (July–Nov); Nigeria, July– Oct; Cameroon, Sept–Nov (July–Dec); Central African Republic (June–Nov); Congo, Oct (prenuptial moult Oct–Jan); Gabon, Mar–May; Sudan, Aug–Oct; Ethiopia, July–Aug, Oct (July–Oct); Uganda, June–Aug; Kenya, May–Sept; Tanzania, June; N Zaı¨re, July–Oct; S Zaı¨re, Jan–Apr; Angola, Mar–May; Zambia, Dec–Mar (Dec); ˆ i, Jan–Feb; Mozambique, (Mar); Zimbabwe, Malaw Dec–Mar. INCUBATION: by X only. In captivity, period 12–14 days (Hermann 1978).

239

DEVELOPMENT AND CARE OF YOUNG: young fed by X only; captive young initially fed insects, later bread, milk, seeds. Nestling period: 15 days (Hermann 1978). Y once fed fledgling (Savalli 1997). BREEDING SUCCESS/SURVIVAL: of 28 nests, 27% fledged 1 or more young; 24% of eggs produced flying young. Most failures are during egg stage; nests destroyed by cattle, people cutting grass, or nests abandoned. In captivity, can hybridize with E. ardens and E. axillaris (Colahan and Craig 1981, Hopkinson 1938).

Key References

Savalli, U. M. (1993a, b, 1994a–c, 1997).

Euplectes ardens (Boddaert). Red-collared Widowbird. Euplecte veuve-noire.

Plate 14

Fringilla ardens Boddaert, 1783. Tabl. Pl. enlum., p. 39; Cape of Good Hope.

(Opp. p. 203)

Range and Status. Endemic resident. Common to abundant in SE Africa, more local and less common elsewhere. Gambia, several old records but none since 1965. Senegal, 1 recent record, Niokolo-Koba Nat. Park, 1992 (Sauvage and Rodwell 1998). Mali, Niger R. valley at Kangaba and Koure´male´, and Mandingo Mts near Sagabari. Guinea, Macenta on inland plateau. Sierra Leone, local along rivers, near Tingi Mts. Liberia, locally common in N. Ivory Coast, Beoumi-Bouake´ to Comoe´ and Korhogo, Goue´ssesso, Marahoue´ Nat. Park. Ghana, 1, Cape Coast, 1977 (Grimes 1987). Togo, rare: Misaho ¨he, Pagala, Djodji.

Benin, no recent records. Nigeria, locally not uncommon on Jos, Obudu and Mambilla Plateaux. Niger, 1988 record in Parc du W requires confirmation. Cameroon, Wum, Tchabal Mbalo, Adamawa Plateau, north to about 7
N. Central African Republic, Bamingui, Lobaye, BaminguiBangoran Nat. Park. Sudan, along S border, Torit, Yei and Boma hills. Ethiopia, W highlands, Rift Valley, S and SE regions. Eritrea, uncommon in highlands. Gabon, Tchibanga. Congo, in S on R. Congo. Zaı¨re, savannas of Uele and Ituri districts, lower Congo valley and Kasai, S savannas and E grasslands. Rwanda. Burundi. Uganda,

240

PLOCEIDAE

Euplectes ardens

widespread in W, east to Acholi, but now apparently scarce around L. Victoria. Kenya, Mt Elgon, Cheranganis and Mt Kenya south to Mau, Mara Game Res., central Rift Valley, Aberdares, Nairobi and Machakos Dist., lower R. Tana. Tanzania, Loliondo, Crater Highlands, Arusha Nat. Park, Mts Meru and Kilimanjaro; widespread in interior to Soga, Utete; at coast at Dar es Salaam and near 10
S. Angola, N Huı´la to lower Congo R., east to Lunda Norte, Lunda Sul and Moxico. Zambia, north of 15
S, western Lusaka Prov., S Central Prov., E sector of Southern and Western Provs. ˆ i, widespread except in lake littoral areas and high Malaw plateaux. Mozambique, Mocuba, Gurue, Ile, Vila Coutinho, Zobue, Namwera, Inhambane, Palmeira, Lebombo Mts. Botswana, once, near Gaborone (Penry 1994). Zimbabwe, central plateau and E highlands. South Africa, plateau areas of Northern Prov., Gauteng, Mpumalanga, N and E Free State, throughout KwaZulu-Natal, down E coast to Port Elizabeth. Swaziland, throughout. Lesotho, mountain slopes and valleys, not on highest areas. Description. E. a. ardens (Boddaert) (includes ‘concolor’): Sierra Leone to Uganda, SW Sudan, NW and S Tanzania, to South Africa. ADULT Y (breeding): black, except for narrow red, sickleshaped collar separating throat and breast (collar rarely orange or yellow; absent in some populations, narrow in intermediates or possible hybrids); black wing feathers edged with pale brown; undertail-coverts edged with white. Tail very long, outermost feathers longest; in South Africa once with white central rectrices. Bill black; eyes brown; legs blackish. ADULT Y (nonbreeding): forehead to rump brown, feathers with black central streaks; uppertail-coverts black with whitish edges; tail short, brown feathers with paler margins; prominent yellowish supercilium; lores, cheeks and ear-coverts brown; chin yellowish white; throat and breast yellow-buff; flanks buff, belly white; thighs buff with black flecks; wings black with buff margins, underwing sooty grey. Bill brown, lower mandible much lighter; eyes brown; legs brown. SUB-ADULT Y: like non-breeding Y, but

dorsal streaking dark brown, not black; underparts with distinct buff band above white belly; undertail-coverts brown with whitish edges; wings brown with pale edges; underwing with some buffy feathers. ADULT X: like sub-ad. Y. E. a. ‘concolor’ is a melanistic variant, from Sierra Leone to Congo R. and Uganda; breeding YY have no red collar. In Angola, S Zaı¨re, adjacent parts ˆ i and S Tanzania, intergrades have very narrow of Zambia, Malaw red collar. SIZE (77 YY, 69 XX): wing, Y 68–85 (744), X 61–69 (646); tail, Y (breeding, n ¼ 55) 183–280 (228), Y (non-breeding, n ¼ 12) 51–62 (550), X 40–49 (442); bill, Y 142–173 (154), X 136–160 (148); tarsus, Y 212–240 (224), X 188–220 (206). WEIGHT: Y (KwaZulu-Natal, n ¼ 187) 200–255 (227), (Zimbabwe, n ¼ 76) 156–233 (186), (Zaı¨re, n ¼ 9) 180–210 (193); X (KwaZulu-Natal, n ¼ 211) 160–215 (197), (Zimbabwe, n ¼ 100) 131–187 (160), (Zaı¨re) 16. IMMATURE: like X but broader buffy margins to feathers of upperparts, underparts with stronger buff wash. NESTLING: sparse covering of pale grey down. E. a. laticauda (Lichtenstein): Ethiopia, Eritrea and SE Sudan. Breeding Y with crown and nape deep red, linked to collar on throat; forehead always black, some crown feathers black-tipped. Much shorter black tail, retained in non-breeding Y. Y has some streaking on breast. SIZE (41 YY, 28 XX): wing, Y 77–88 (832), X 65–71 (676); tail, Y (breeding, n ¼ 37) 121–156 (141), X 43–65 (563); bill, Y 154–172 (161), X 143–158 (149); tarsus, Y 230– 254 (240), X 197–224 (216). WEIGHT: Y 267, 270; X (n ¼ 4) 180–200 (191). E. a. suahelica (van Someren): highlands of Kenya and Tanzania. Breeding Y like laticauda but extent of red on head variable, sometimes forehead red almost to base of bill. Nonbreeding Y like non-breeding Y laticauda. SIZE (35 YY, 32 XX): wing, Y 75–91 (792), X 63–69 (671); tail, Y (breeding, n ¼ 27) 145–204 (170), X 47–67 (564); bill, Y 144–164 (156), X 132– 155 (147); tarsus, Y 224–245 (232), X 200–220 (213). WEIGHT: (Kenya) Y (n ¼ 12) 219–259 (237), X (n ¼ 14) 170– 230 (197). TAXONOMIC NOTE: continuously distributed lowland populations intergrade; highland suahelica and laticauda are isolated from lowland birds and from each other, and in terms of the phylogenetic species concept they could be treated as a separate species.

Field Characters. Length 12–13 cm (breeding Y 25– 30 cm). Breeding Y has small body and long, thin floppy tail (A); colour variable, either all black (W Africa) or with crescent-shaped red collar on upper breast, head black (southern Africa) or with varying amounts of red on crown and nape (E Africa). Streaky brown X and non-breeding Y have prominent yellowish supercilium, unstreaked buffy breast sharply demarcated from white belly; Y retains black wings and sometimes variably elongated black tail; Black-winged Bishop E. hordeaceus has black wings but much larger bill, streaked breast. Tail feathers narrow and pointed, like much larger Long-tailed and Jackson’s Widowbirds E. progne and E. jacksoni; X and sub-ads have tapered undertail-coverts with pale edges and dark centres, sometimes visible in the field, obvious in the hand. Occurs with all other members of the genus; most misidentified museum specimens belong to this species. Voice. Tape-recorded (86, 88, 99, 104, B, C, F, LEM, MOY, PAY). Song from perch high, sibilant sewing machine or insect-like trill, either continuous or in short repeated bursts, ‘chisisisisi-chisisisisisi-chisisisisi . . . ’; in other songs, slower sewing machine alternates with scraping ‘jaar’ (from second bird?), ‘chi-si-si-si-si-si-si-si- . . . jaar . . . chi-si-si-si-si-si-si . . . jaar . . . chi-si-si-si-si-si-si’.

Euplectes ardens changes seasonally; in Zimbabwe XX predominate in Nov–Jan, YY in June–Oct; in KwaZulu-Natal, more XX in June–Aug than earlier. Sex ratio of juveniles does not change (Craig and Manson 1979b). Anting recorded (in captive birds: Poulsen 1956). In South Africa, complete post-nuptial moult from Feb to June, when both ads and juvs replace entire plumage (Craig and Manson 1979a); in Liberia, post-nuptial moult begins in Feb. Y acquires black wing-feathers at 2nd complete moult and comes into breeding plumage during following pre-nuptial moult period. Mainly sedentary, but can move locally during nonbreeding season. 1 bird in Zimbabwe recovered 100 km from ringing site. Food. Seeds, insects, some nectar. Seeds of sorghum, Panicum maximum (Skead 1997). Nectar of Leonotis (Swynnerton 1908) and Aloe candelabrum (Oatley and Skead 1972). In 12 crops and stomachs, 11 held grass seed, some still green; and 3, insects including ants, beetles and a small caterpillar. Of 9 stomachs, 6 held grass seeds only, 3 contained beetles and a caterpillar. Seeds, small berries and grit in stomachs (Sjo ¨stedt 1910).

Wide vocabulary includes hissing, drawn-out ‘sscherz’ or ‘zzhooy’ like tearing paper, sometimes followed by rapid stuttering ‘hitza hitza’, both perched and in flight; soft husky ‘hizz zizz zizz’ and rapid ‘screep-screep’ in flight; also low buzzy ‘dzer’, harsh forceful ‘chrrt chrrt’, rapid staccato ‘chekety chekety chekety’ or ‘wit-zitti-wit-zittiwit’, and grating rattle. Alarm a husky ‘sskip sskip’ accompanied by wing-flicking (Skead 1995). General Habits. Uses wider range of habitats than congeners, often away from water, and on slopes above wet areas: tall open or bushed grassland, rank herbaceous growth, scrub and cultivated areas, clearings and degraded land in forest. In L. Lufira region of Zaı¨re moves to permanent swamps after burning of grasslands and favours areas of Hyparrhenia rufa or Pennisetum sp. In Matopos Nat. Park, Zimbabwe, colonized Pennisetum glaucocladum reedbed following changes in land management. In E Africa, ardens occurs usually at 100–200 m, occasionally at coast; in Ruwenzoris not above 1500 m; in Cameroon up to 1900 m. E. a. laticauda occurs in Ethiopia usually above 1800 m (often associated with Yellow-mantled Widowbird E. macroura); suahelica in E Africa at 1500–3000 m. Forages on ground, often in flocks of 200 birds or more; winged termites caught and eaten on ground, hawked in air and then eaten at a perch or on ground (Brooke 1970). Normal flight of Y is rapid, with tail stretched out straight behind. Forms large roosts, even during breeding season, sometimes with Red-billed Quelea Quelea quelea or Redshouldered Widowbirds E. axillaris. Forms large mixed flocks in Angola with E. ardens, White-winged Widowbird E. albonotatus, Yellow Bishop E. capensis, Southern Red Bishop E. orix (Braun 1930). Sex ratio of ads in roosts

Breeding Habits. Polygynous, territorial. In Ethiopia, 6 YY held territories on a 500 m stretch of grassy hill-slope; territory size in E Cape, South Africa, c. 7500 m2. From conspicuous perch, Y flies at intruders which enter territory; may displace Y Southern Red Bishop and Y White-winged Widowbird, but does not interact with Longtailed Widowbird where respective territories overlap. Once dropped into grass, locked in fight, but usually Y on ‘patrol’ turns back before crossing boundary of territory. Y makes long (20–30 m), low flights with exaggerated wing-beats, during which tail is spread but shorter central feathers also curved downwards, and head feathers erected into a hood; he may hover with quivering wings. Display flights directed at non-breeding-plumaged birds and at other YY. When landing, Y flops onto perch with tail askew, wings outspread. He sings from high perches and low in grass. In display at perch, he quivers wings, fluffs body feathers, gives insect-like song; he bounces up and down, facing X, throat inflated so that red collar is prominent. Sparrowyplumaged sub-ad. Y carries out typical display flight even before acquiring long tail. NEST: made of fine wiry grass woven onto grass stems (B), with green blades of grass bent over the top so that there is a living grass dome; outer shell appears flimsy; lined with grass heads stripped of seeds (Panicum maximum, Eragrostis curvula), which may project from side entrance to form a porch. 1 nest contained 518 seed heads (Skead 1995). Placed in tall grass, particularly elephant grass, at 10–35 m above ground. Built mainly by X (van Someren 1956); however, Y once brought material from 300 m away and wove it into nest (Skead 1995). X continues to add to lining during incubation, and eggs may be laid before much lining has been added. ˆ i); grey or blue-green EGGS: 2–3 (22, n ¼ 74, Malaw heavily speckled with brown, often forming ring near thick end. SIZE: (n ¼ 16, Kenya) 177–219  130–150

241

242

PLOCEIDAE

LAYING DATES: Liberia, Sept–Oct (breeding plumage from Aug); Nigeria, (breeding plumage Sept, copulation Nov, fledged young Oct); Cameroon, (breeding plumage Aug–Dec); Gabon, (breeding plumage Apr); Ethiopia, Apr–May, July–Sept (possible Oct; breeding plumage June–Nov); Zaı¨re, Sept–Nov in NE, Jan–May in SE; Rwanda, Mar–Apr; Uganda, Apr–June, Sept; Kenya, Apr– July, Dec–Jan; Tanzania, Jan–May (Arusha Nat. Park); Angola, (breeding plumage Mar–May); Zambia, Dec–Apr; ˆ i, Jan–May; Mozambique, Dec–May in N, Oct–Mar Malaw in S; Zimbabwe, Dec–Mar, rarely Apr; South Africa, Oct–Mar. INCUBATION: by X only, period 12–15 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only, primarily by regurgitation; nestling period 14–17 days. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius. Eggs and newly hatched young taken by rats and snakes. Fledglings, on leaving nest but unable to fly far, shelter in vegetation where sometimes taken by raptors or mongooses. Of 195 birds ringed in southern Zaı¨re, 9 recaptured, only 1 >1 year after ringing (Bont et al. 1965). In Zimbabwe, recaptured up to 4 years after ringing; annual ad. mortality c. 46% (Haldane’s method: Craig and Manson 1979b). Oldest bird recaptured after 75 years (Manson 1982). Worms under skin of some Zambian birds were larval Acanthocephala, final hosts of which are predatory animals (White 1943).

(198  139), (n ¼ 55, South Africa) 173–204  127–144 (189  137).

Plate 14 (Opp. p. 203)

Key References (1956).

Skead, C. J. (1995), van Someren, V. G. L.

Euplectes axillaris (A. Smith). Fan-tailed Widowbird; Red-shouldered Widow. Euplecte a` ´epaules orange´es. Vidua axillaris Smith, 1838. Ill. Zool. S. Afr., p. 17; Eastern Cape. Range and Status. Endemic resident. Mali, isolated populations on Niger R. between 14
and 17
N; Niger, local along Niger R. upstream from Tillabe´ri and in ‘W’ Nat. Park (Crisler et al. 2003). Nigeria, known only from Malamfatori and Logomani, west of L. Chad; Cameroon, highlands at Foumban, Ndop, Djuttitsa south to Dschang, in north at Baga Kawa and on Logone Gana R.; Chad, close to L. Chad; Central African Republic, MonovoGounda-Saint Floris Nat. Park; Sudan, south of 14
N, primarily along Nile and its tributaries; Ethiopia, western highlands south of Addis Ababa, rift valley, and southern region; Somalia, lower and mid-sections of Jubba and Shabeelle Rivers; Congo; Zaı¨re, Katanga, Maniema, Marungu Mts, Kananga and in the east from L. Albert to north of L. Tanganyika; Rwanda and Burundi, local near lakes; Uganda, L. Victoria basin, north to Bunyoro, Lango, Teso; Kenya, on the coast from Lamu south, inland to Kilosa and Tana R., L. Victoria basin east to Nandi, up to 2300 m at Uasin Gishu; Tanzania, on coast south to Rufiji, Mafia Island, typically below 1500 m, up to 2000 m at ˆ i, generally below Mjombe and Mbulu; Zambia and Malaw

900 m; Angola, from Cuando Cubango west to W and N Huı´la, across Huambo to Cuanza Sul, east to Malanje, N Lunda Norte, Lunda Sul and NE Moxico, along Cuanza R. in southern Bengo; Namibia, restricted to Caprivi region; Botswana; in Okavango basin; Zimbabawe, along Zambezi R. west of Victoria Falls; Mozambique, Mixixine, L. Amaramba in the north, along coast south of Save R. and in southern sector; South Africa, from Algoa Bay area in a broad belt between the escarpment and coast, northeast to Gauteng and Mpumalanga; Lesotho, 1 old record; Swaziland, widespread throughout. Common. ˆ i south Description. E. a. axillaris (Smith): Mozambique, Malaw to E South Africa. ADULT Y (breeding): plumage black except for wing; tail long and fan-shaped; wing with flight feathers black, lesser coverts red, greater coverts cinnamon brown, buff or light brown edges to tertials and secondary coverts; wing edge and underwing cinnamon-brown. Bill metallic blue; eyes brown; legs black. ADULT Y (non-breeding): forehead and crown black with narrow brown edges to feathers; nape, mantle and rump feathers brown with broad blackish brown central streaks; tail dark brown; chin and throat whitish; breast and flanks buff with faint

242

PLOCEIDAE

LAYING DATES: Liberia, Sept–Oct (breeding plumage from Aug); Nigeria, (breeding plumage Sept, copulation Nov, fledged young Oct); Cameroon, (breeding plumage Aug–Dec); Gabon, (breeding plumage Apr); Ethiopia, Apr–May, July–Sept (possible Oct; breeding plumage June–Nov); Zaı¨re, Sept–Nov in NE, Jan–May in SE; Rwanda, Mar–Apr; Uganda, Apr–June, Sept; Kenya, Apr– July, Dec–Jan; Tanzania, Jan–May (Arusha Nat. Park); Angola, (breeding plumage Mar–May); Zambia, Dec–Apr; ˆ i, Jan–May; Mozambique, Dec–May in N, Oct–Mar Malaw in S; Zimbabwe, Dec–Mar, rarely Apr; South Africa, Oct–Mar. INCUBATION: by X only, period 12–15 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only, primarily by regurgitation; nestling period 14–17 days. BREEDING SUCCESS/SURVIVAL: parasitized by Diederik Cuckoo Chrysococcyx caprius. Eggs and newly hatched young taken by rats and snakes. Fledglings, on leaving nest but unable to fly far, shelter in vegetation where sometimes taken by raptors or mongooses. Of 195 birds ringed in southern Zaı¨re, 9 recaptured, only 1 >1 year after ringing (Bont et al. 1965). In Zimbabwe, recaptured up to 4 years after ringing; annual ad. mortality c. 46% (Haldane’s method: Craig and Manson 1979b). Oldest bird recaptured after 75 years (Manson 1982). Worms under skin of some Zambian birds were larval Acanthocephala, final hosts of which are predatory animals (White 1943).

(198  139), (n ¼ 55, South Africa) 173–204  127–144 (189  137).

Plate 14 (Opp. p. 203)

Key References (1956).

Skead, C. J. (1995), van Someren, V. G. L.

Euplectes axillaris (A. Smith). Fan-tailed Widowbird; Red-shouldered Widow. Euplecte a` ´epaules orange´es. Vidua axillaris Smith, 1838. Ill. Zool. S. Afr., p. 17; Eastern Cape. Range and Status. Endemic resident. Mali, isolated populations on Niger R. between 14
and 17
N; Niger, local along Niger R. upstream from Tillabe´ri and in ‘W’ Nat. Park (Crisler et al. 2003). Nigeria, known only from Malamfatori and Logomani, west of L. Chad; Cameroon, highlands at Foumban, Ndop, Djuttitsa south to Dschang, in north at Baga Kawa and on Logone Gana R.; Chad, close to L. Chad; Central African Republic, MonovoGounda-Saint Floris Nat. Park; Sudan, south of 14
N, primarily along Nile and its tributaries; Ethiopia, western highlands south of Addis Ababa, rift valley, and southern region; Somalia, lower and mid-sections of Jubba and Shabeelle Rivers; Congo; Zaı¨re, Katanga, Maniema, Marungu Mts, Kananga and in the east from L. Albert to north of L. Tanganyika; Rwanda and Burundi, local near lakes; Uganda, L. Victoria basin, north to Bunyoro, Lango, Teso; Kenya, on the coast from Lamu south, inland to Kilosa and Tana R., L. Victoria basin east to Nandi, up to 2300 m at Uasin Gishu; Tanzania, on coast south to Rufiji, Mafia Island, typically below 1500 m, up to 2000 m at ˆ i, generally below Mjombe and Mbulu; Zambia and Malaw

900 m; Angola, from Cuando Cubango west to W and N Huı´la, across Huambo to Cuanza Sul, east to Malanje, N Lunda Norte, Lunda Sul and NE Moxico, along Cuanza R. in southern Bengo; Namibia, restricted to Caprivi region; Botswana; in Okavango basin; Zimbabawe, along Zambezi R. west of Victoria Falls; Mozambique, Mixixine, L. Amaramba in the north, along coast south of Save R. and in southern sector; South Africa, from Algoa Bay area in a broad belt between the escarpment and coast, northeast to Gauteng and Mpumalanga; Lesotho, 1 old record; Swaziland, widespread throughout. Common. ˆ i south Description. E. a. axillaris (Smith): Mozambique, Malaw to E South Africa. ADULT Y (breeding): plumage black except for wing; tail long and fan-shaped; wing with flight feathers black, lesser coverts red, greater coverts cinnamon brown, buff or light brown edges to tertials and secondary coverts; wing edge and underwing cinnamon-brown. Bill metallic blue; eyes brown; legs black. ADULT Y (non-breeding): forehead and crown black with narrow brown edges to feathers; nape, mantle and rump feathers brown with broad blackish brown central streaks; tail dark brown; chin and throat whitish; breast and flanks buff with faint

Euplectes axillaris

Euplectes axillaris

?

dark streaks; belly whitish; thighs and undertail-coverts buff. Bill brown in birds which have just acquired black wing feathers, blue colour retained in older birds. Legs dark brown. SUB-ADULT Y: like non-breeding Y, but wings brown with pale edges, lesser coverts edged with cinnamon to orange-brown but no red on epaulet, underwing cinnamon. Upper mandible brown, lower paler brown; eyes brown; legs pale brown to flesh-coloured. ADULT X: like sub-ad. Y, but smaller. SIZE (38 YY, 10 sub-ad. YY, 16 XX): wing, Y 81–94 (888), sub-ad. Y 69–81 (764), X 64–72 (680); tail, Y (breeding) 64–86 (755), (non-breeding, n ¼ 18) 54– 70 (625), sub-ad. Y 46–52 (496), X 42–49 (449); bill, Y 172–200 (182), sub-ad. Y 146–180 (170), X 155–173 (165); tarsus, Y 234–266 (250), sub-ad. Y 228–260 (246), X 206–224 (216). WEIGHT: Y (KwaZulu-Natal, South Africa, n ¼ 268) 265–320 ˆ i, Mozambique, n ¼ 16) 22–28 (251); sub-ad. Y (295), (Malaw (KwaZulu-Natal, n ¼ 226) 255–315 (280); X (KwaZulu-Natal, ˆ i, n ¼ 21) 17–23 (187). n ¼ 406) 200–250 (220), (Malaw IMMATURE: like ad. X, but very broad buffy edges to feathers of the upperparts; bill uniform pale brown. NESTLING: not described. TAXONOMIC NOTE: populations in coastal E Africa and near coast in Angola are particularly heavy-billed. This may have an ecological explanation, related to differences in diet (Craig 1993c). E. a. bocagei (Sharpe) (includes ‘mechowi’ and ‘quanzae’): W Africa south to Angola, SW Zaı¨re, N Botswana and Namibia. Breeding Y has lesser coverts orange-yellow, greater coverts and primary coverts cinnamon-brown, making a much broader epaulet than in other populations. Large-billed, particularly in coastal Angola. SIZE (52 YY, 22 XX): wing, Y 82–97 (881), X 68–78 (739); tail, Y (breeding) 64–83 (742), (non-breeding, n ¼ 18) 59–74 (653), X 45–56 (504); bill, Y 164–191 (178), X 154–172 (161); tarsus, Y 242–282 (263), X 210–235 (225). WEIGHT: 1 Y 30. E. a. phoeniceus (Heuglin): S Sudan through E Africa to E Zambia. Breeding Y has lesser coverts orange, not red; greater coverts and primary coverts cinnamon-brown. SIZE (26 YY, 18 XX): wing, Y 80–90 (857), X 65–72 (687); tail, Y (breeding) 59– 70 (636), (non-breeding, n ¼ 25) 50–64 (568), X 41–48 (440);

bill, Y 162–181 (172), X 148–160 (156); tarsus, Y 234–261 (244), X 203–228 (218). WEIGHT: Y (Kenya, n ¼ 69) 230–292 (265); X (Kenya, n ¼ 84) 182–249 (209) (Britton 1977); Y (Nandi, Kenya) 262–300, X 178–215 (Schifter and Cunninghamvan Someren 1998). E. a. traversii (Salvadori): Ethiopian highlands. Breeding Y proportionately longer-tail; like phoeniceus in plumage, but on underwing, coloured area at base of primaries extends beyond coverts, and alula usually wholly cinnamon-brown. SIZE (21 YY, 25 XX): wing, Y 84–94 (899), X 66–78 (730); tail, Y (breeding) 76–98 (855), (non-breeding, n ¼ 17) 57–73 (662), X 42–54 (484); bill, Y 160–182 (171), X 141–170 (156); tarsus, Y 243–275 (259), X 210–241 (224). E. a. zanzibaricus (Shelley): coastal Kenya, Tanzania and Somalia, and offshore islands. Breeding Y like phoeniceus, but dark spots on cinnamon wing-coverts, and heavy bill. SIZE (28 YY, 6 XX): wing, Y 81–95 (881), X 70–75 (725); tail, Y (breeding) 58–80 (688), (non-breeding, n ¼ 2) 56, 58, X 44–49 (468); bill, Y 168–200 (188), X 164–187 (172); tarsus, Y 243– 270 (255), X 213–228 (220). WEIGHT: Y (Kenya) 275, 290, 308; X (Kenya) 220, 228 (Britton 1977).

Field Characters. Length 15 cm. Breeding Y black with pale blue-grey bill, orange-red and cinnamon shoulder patch; broad fan-shaped tail with rounded tips to rectrices much shorter than Hartlaub’s Widowbird E. hartlaubi. Non-breeding Y heavily streaked black above, with dark crown, conspicuous pale superciliary stripe, red shoulders, black primaries and cinnamon underwing; X similar but primaries brown, shoulder duller and often partly concealed. Flight undulating, with broad wings. Voice. Tape-recorded (88, 99, 104, B, C, F, PAY). Song, given in flight and from prominent perch, weak, highpitched and scratchy, shrill ‘zeep’ repeated at 06–08 s intervals while second bird gives sibilant ‘sisisisisisi . . . ’; or ‘zeep’ may be followed by almost liquid chatter and thin ‘zip’ notes; rendered ‘skreep skrik skrik wirra skreek skreek wirrily wirrily wirrily chink chink’ (Maclean 1993). In patrol flight Y gives 1–2 single notes followed by rolling trill, ‘jerp-tirrrrrr, jerp-tirrrrrr’, ‘chuk chuk turrrr, chuk chuk turrrr’ or ‘tseek-wirra-wirra tseek-wirra-wirra’; when perched, a low grating swizzle like the sound of paper being torn, ‘jerzzh . . . jerzzh . . . jerzzh’. Contact call ‘zipzip’; quiet rolling call heard in feeding flocks during breeding season (Skead 1959). General Habits. Inhabits tall moist grassland and swamp edges in much of its range, but at the coast occurs also in dry bushed grassland and cultivation including sugarcane fields. Seasonal shift in L. Lufira region, Zaı¨re, from permanent swamps with Typha to periodically flooded margins of plains dominated by grasses such as Hyparrhenia rufa and Themeda triandra, where YY establish territories at start of breeding season. In Okavango delta, in reedbeds and papyrus; in Zambia, confined to swamps and adjacent wet grasslands. Typically below 1500 m but in Tanzania and Kenya also at 2000–2300 m, and in E Zaı¨re from lake shores up to plateaux at 2000 m. Gregarious, forming large post-breeding flocks; roosts in association with other Euplectes and Ploceus species. Pure flocks of 5–30 birds in Botswana, >100 in South Africa and Mozambique. During breeding season, Y feeds inside and outside territory, and

243

244

PLOCEIDAE

joins flocks for feeding and roosting. Ringing studies in KwaZulu-Natal, South Africa, suggest that birds are resident. Sex ratios showed excess of YY in 5 months and of XX in 1 month only; overall ratio showed significant imbalance in favour of YY (Craig and Manson 1979a). Food. Seeds of grasses and herbs; also insects, including termites and caterpillars. Seeds of rice, Setaria, Digitaria, Paspalum (Skead 1997), Polygonum senegalense, Echinochloa colonum. Non-breeding flocks in maize fields (Granvik 1923). Breeding Habits. Territorial, polygynous. Up to 8 nests in a territory of 06 ha. Y sometimes escorts Y Southern Red Bishops E. orix and Long-tailed Widowbirds through territory. Interspecific territorial behaviour occurs with Y Hartlaub’s Marsh Widowbird in Katanga, where territory covers 05 ha (Ruwet 1964b, 1965). In boundary display, Y tenses body, puffs out feathers, flicks wings and sways from side to side. Y flies over territory in slow, undulating flight on rounded wings, just above grass tops, then suddently twists backwards to alight on tall grass stem. Sings from these prominent perches, with epaulets conspicuous, tail spread and hindneck ruff erected. If X lands in territory, Y from patrol flight dives into grass, postures with wings arched and spread, leaning forward. He erects epaulet feathers and ruff, depresses bill, and apparently mandibulates, with audible ticking sound; bounces up and down in front of X, singing loudly. Y also sings at nest frame with tail spread, beak depressed and wings lifted slightly and quivered; sways from side to side. X may then enter nest. NEST: oval ball of thin woven grass strips, with dense lining of seedheads projecting from entrance, attached to 2 or more stems of grass or other plants. 1 nest was in territory of Long-tailed Widowbird, close to active nests; in tall rank grass of swampy areas, also in rice fields, only

5–10 cm off ground in grass tuft; elsewhere nests are 60–80 cm above ground. Size, 140 high, 65 wide, 176 deep, entrance diam. 50. Y builds frame, weaving long grass strands into bower of living green grass; X packs lining of up to 540 grass seedheads into it. EGGS: 2–3, rarely 4, av. 28 (South Africa, n ¼ 115); pale bluish green, irregularly marked with large spots of olivebrown; grey-green with large and small grey-brown or violet-brown flecks running into each other, covering blunt end; olive-green with irregular and sparse brown markings; dull grey-white, heavily overlaid by blotches and streaks of brown, some predominantly smeared with brown. SIZE: (South Africa, n ¼ 69) 177–224  133–150 ˆ i 185–195  145, Uganda, 180– (197  141); Malaw 205  130–140. LAYING DATES: Mali, (breeding plumage Sept); Niger, (breeding plumage Dec–Jan); Cameroon, (July–Sept breeding plumage; Sept–Oct breeding); Sudan, Aug; Ethiopia, (Sept–Oct); Somalia, Aug–Oct; Zaı¨re, Aug–Oct in NE, Jan–Mar in SE; Rwanda, Nov–Dec, Feb–June; Uganda, primarily Mar–July, also Oct–Jan; Kenya, Apr–July; Tanzania, Mar; Angola, Jan–Apr; Zambia, Dec–Mar; ˆ i, Dec, Feb–Apr; South Africa, Oct–Mar. Malaw INCUBATION: by X only, period 12–13 days; in captivity, 12–14 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only; nestling period 15–16 days; faecal sacs dropped up to 100 m from nest. In captivity, nestling period 14 days, young fed by X for 14 days after leaving nest. BREEDING SUCCESS/SURVIVAL: in captivity hybridizes with X E. macrourus and Y E. capensis (Hopkinson 1938).

Key References

Craig, A. J. F. K. (1980), Skead, C. J. (1959).

Plate 15

Euplectes hartlaubi (Bocage). Hartlaub’s Marsh Widowbird. Euplecte des marais.

(Opp. p. 218)

Penthetria hartlaubi Bocage, 1878. J. Lisboa, 6, p. 259; Caconda, Angola. Forms a superspecies with E. psammocromius. Range and Status. Endemic resident. E Nigeria, Mambilla Plateau. Cameroon, rare, at Ngaounde´re´, Akonolinga, Tibati, Nun R., Djuttitsa. Gabon, in savanna near Congo border. Congo, S savanna areas. Zaı¨re, Kwilu, Kwango, Kinshasha to Mbandaka, along lower reaches of Congo R., also in S Katanga. Uganda, Mengo and Busoga regions near shores of L. Victoria, and highlands east of Mt Elgon. Kenya, highlands near Mt Elgon, south to Nandi and Kakamega. Angola, northern Huı´la to Huambo, Bie´ and southern Cuanza Sul, Malanje to Lunda Norte, Lunda Sul and northern Moxico. Tanzania, Ufipa area. Zambia, Muchinga escarpment south to Kondolilo Falls, west through Northern and Luapula Provinces to Mbala, Mporokoso, Kawambwa and Samfya, Central Province to Mkushi and Luamala R., Western Province at Mwekera, Lalulushi and Luanshya, Upper Kafue south to 13
S,

Northwestern Province from Solwezi south to Balovale. Localised and generally uncommon. Description. E. h. hartlaubi (Bocage): Angola, S Zaı¨re (north to Upemba Nat. Park and Marungu Plateau), Zambia. ADULT Y (breeding): plumage wholly black except for wing-coverts; long black tail; greater wing-coverts with narrow buff margins, lesser coverts bright orange, grading into buff on posterior border; buff spot on wing margin, at base of outermost primary. Bill blueblack, edges bluish white; eyes dark brown; legs blackish. ADULT Y (non-breeding): retains orange coverts and black wing feathers, but long black tail replaced by short brown feathers with paler edges. Upperparts dark brown, feathers with buff margins; underparts dull tawny-white, some breast feathers with narrow brown central streak; short dark moustachial streaks. Bill dark brown; eyes brown; legs brown. SUB-ADULT Y: like non-breeding Y, except that wing feathers dark brown, with buff margins, lacks conspicuous epaulet but has narrow orange-yellow margins to

244

PLOCEIDAE

joins flocks for feeding and roosting. Ringing studies in KwaZulu-Natal, South Africa, suggest that birds are resident. Sex ratios showed excess of YY in 5 months and of XX in 1 month only; overall ratio showed significant imbalance in favour of YY (Craig and Manson 1979a). Food. Seeds of grasses and herbs; also insects, including termites and caterpillars. Seeds of rice, Setaria, Digitaria, Paspalum (Skead 1997), Polygonum senegalense, Echinochloa colonum. Non-breeding flocks in maize fields (Granvik 1923). Breeding Habits. Territorial, polygynous. Up to 8 nests in a territory of 06 ha. Y sometimes escorts Y Southern Red Bishops E. orix and Long-tailed Widowbirds through territory. Interspecific territorial behaviour occurs with Y Hartlaub’s Marsh Widowbird in Katanga, where territory covers 05 ha (Ruwet 1964b, 1965). In boundary display, Y tenses body, puffs out feathers, flicks wings and sways from side to side. Y flies over territory in slow, undulating flight on rounded wings, just above grass tops, then suddently twists backwards to alight on tall grass stem. Sings from these prominent perches, with epaulets conspicuous, tail spread and hindneck ruff erected. If X lands in territory, Y from patrol flight dives into grass, postures with wings arched and spread, leaning forward. He erects epaulet feathers and ruff, depresses bill, and apparently mandibulates, with audible ticking sound; bounces up and down in front of X, singing loudly. Y also sings at nest frame with tail spread, beak depressed and wings lifted slightly and quivered; sways from side to side. X may then enter nest. NEST: oval ball of thin woven grass strips, with dense lining of seedheads projecting from entrance, attached to 2 or more stems of grass or other plants. 1 nest was in territory of Long-tailed Widowbird, close to active nests; in tall rank grass of swampy areas, also in rice fields, only

5–10 cm off ground in grass tuft; elsewhere nests are 60–80 cm above ground. Size, 140 high, 65 wide, 176 deep, entrance diam. 50. Y builds frame, weaving long grass strands into bower of living green grass; X packs lining of up to 540 grass seedheads into it. EGGS: 2–3, rarely 4, av. 28 (South Africa, n ¼ 115); pale bluish green, irregularly marked with large spots of olivebrown; grey-green with large and small grey-brown or violet-brown flecks running into each other, covering blunt end; olive-green with irregular and sparse brown markings; dull grey-white, heavily overlaid by blotches and streaks of brown, some predominantly smeared with brown. SIZE: (South Africa, n ¼ 69) 177–224  133–150 ˆ i 185–195  145, Uganda, 180– (197  141); Malaw 205  130–140. LAYING DATES: Mali, (breeding plumage Sept); Niger, (breeding plumage Dec–Jan); Cameroon, (July–Sept breeding plumage; Sept–Oct breeding); Sudan, Aug; Ethiopia, (Sept–Oct); Somalia, Aug–Oct; Zaı¨re, Aug–Oct in NE, Jan–Mar in SE; Rwanda, Nov–Dec, Feb–June; Uganda, primarily Mar–July, also Oct–Jan; Kenya, Apr–July; Tanzania, Mar; Angola, Jan–Apr; Zambia, Dec–Mar; ˆ i, Dec, Feb–Apr; South Africa, Oct–Mar. Malaw INCUBATION: by X only, period 12–13 days; in captivity, 12–14 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only; nestling period 15–16 days; faecal sacs dropped up to 100 m from nest. In captivity, nestling period 14 days, young fed by X for 14 days after leaving nest. BREEDING SUCCESS/SURVIVAL: in captivity hybridizes with X E. macrourus and Y E. capensis (Hopkinson 1938).

Key References

Craig, A. J. F. K. (1980), Skead, C. J. (1959).

Plate 15

Euplectes hartlaubi (Bocage). Hartlaub’s Marsh Widowbird. Euplecte des marais.

(Opp. p. 218)

Penthetria hartlaubi Bocage, 1878. J. Lisboa, 6, p. 259; Caconda, Angola. Forms a superspecies with E. psammocromius. Range and Status. Endemic resident. E Nigeria, Mambilla Plateau. Cameroon, rare, at Ngaounde´re´, Akonolinga, Tibati, Nun R., Djuttitsa. Gabon, in savanna near Congo border. Congo, S savanna areas. Zaı¨re, Kwilu, Kwango, Kinshasha to Mbandaka, along lower reaches of Congo R., also in S Katanga. Uganda, Mengo and Busoga regions near shores of L. Victoria, and highlands east of Mt Elgon. Kenya, highlands near Mt Elgon, south to Nandi and Kakamega. Angola, northern Huı´la to Huambo, Bie´ and southern Cuanza Sul, Malanje to Lunda Norte, Lunda Sul and northern Moxico. Tanzania, Ufipa area. Zambia, Muchinga escarpment south to Kondolilo Falls, west through Northern and Luapula Provinces to Mbala, Mporokoso, Kawambwa and Samfya, Central Province to Mkushi and Luamala R., Western Province at Mwekera, Lalulushi and Luanshya, Upper Kafue south to 13
S,

Northwestern Province from Solwezi south to Balovale. Localised and generally uncommon. Description. E. h. hartlaubi (Bocage): Angola, S Zaı¨re (north to Upemba Nat. Park and Marungu Plateau), Zambia. ADULT Y (breeding): plumage wholly black except for wing-coverts; long black tail; greater wing-coverts with narrow buff margins, lesser coverts bright orange, grading into buff on posterior border; buff spot on wing margin, at base of outermost primary. Bill blueblack, edges bluish white; eyes dark brown; legs blackish. ADULT Y (non-breeding): retains orange coverts and black wing feathers, but long black tail replaced by short brown feathers with paler edges. Upperparts dark brown, feathers with buff margins; underparts dull tawny-white, some breast feathers with narrow brown central streak; short dark moustachial streaks. Bill dark brown; eyes brown; legs brown. SUB-ADULT Y: like non-breeding Y, except that wing feathers dark brown, with buff margins, lacks conspicuous epaulet but has narrow orange-yellow margins to

Euplectes hartlaubi

Euplectes hartlaubi

‘zhrree’, falling, ‘sserrraan’, or shorter note on 1 pitch, ‘zraa’, followed by 2–4 barely audible high thin lisps, ‘ps ku’, ‘tsi-tsu’, ‘su-sitisit’ or ‘see-see-peu’; repeated at 3–5 s intervals. Shorter notes may lead into rising, buzzy yet semi-musical trill with prolonged terminal phrase, ‘chrrrrrrittterweeeeeeeeeeee’ (Zimmerman et al. 1996). Song described as a simple rapid high-pitched squeaking (Dowsett and Dowsett-Lemaire 1980), like that of congeners (Aspinwall and Beel 1998). Common call of perched Y, short dry ‘drrrt’.

General Habits. Little known. Inhabits swampy grassland and nearby cultivation in valleys. In marshes may also venture into areas of sparse reeds in deep water, perching on floating aquatic plants such as lily leaves and sudd. In E Africa at 1100–1800 m. In Zambia occurs in wet dambos alongside Red-collared Widowbird E. ardens; associates also with Yellow-mantled Widowbird and Fan-tailed Widowbird. Generally in small groups of 10–15 birds.

Food. Grass seeds and insects. Grasshoppers and berries in stomachs.

anterior wing-coverts; upperparts less heavily streaked; underparts more grey than tawny. Bill pale brown. ADULT X: like subad. Y. SIZE (27 YY, 15 sub-ad. YY, 28 XX): wing, Y 101–115 (109), sub-ad. Y 90–107 (953), X 77–88 (837); tail, Y (breeding, n ¼ 13) 163–201 (177), Y (non-breeding, n ¼ 11) 76–87 (816), sub-ad. Y 64–83 (706), X 57–72 (642); bill, Y 186–207 (197), sub-ad. Y 180–204 (190), X 155–195 (179); tarsus, Y 277–303 (289), sub-ad. Y 273–306 (286), X 242–266 (255). IMMATURE: like X, but underparts browner, wing-coverts edged tawny. NESTLING: no information. TAXONOMIC NOTE: montane populations, formerly E. h. psammocromius, are now treated as a separate species (q.v.). E. h. humeralis (Sharpe): north of 5
S. Breeding Y much shorter-tailed than nominate race; tail (n ¼ 11) 98–125 (112).

Field Characters. Length: nominate race, breeding Y 36 cm, X 18 cm; race humeralis, breeding Y 20 cm, X 15 cm. Black breeding Y with white bill, yellow or orange and buff shoulder patch like Fan-tailed Widowbird E. axillaris but tail much longer; birds in Kenya sympatric with ‘Yellow-shouldered Widowbird’ E. macroura macrocerca have orange epaulet, those further south have rich yellow epaulet but sympatric E. macroura has yellow mantle. Nonbreeding Y retains shoulder-patch and dark wings; X and juv. have dusky underwing, separated from Long-tailed E. progne and Yellow-mantled Widowbirds by wholly buff underparts (underparts whitish in Long-tailed, washed yellow in Yellow-mantled). Voice. Tape-recorded (86, 102, 104, B, F, CHA, GREG, LEM). Distinctive call a single rolling, metallic, buzzy note with curious nasal quality, either rising, ‘grrreeng’ or

Breeding Habits. Polygynous, territorial; Y favours a particular perch. Displaying Y flies low, calling, with orange epaulets conspicuous. Chases any intruding Y, also Y Fan-tailed Widowbirds, out of territory. Calls from conspicuous perches within the territory. Where territories of Hartlaub’s Marsh Widowbird and Yellow-mantled Widowbird are within 9 m, Y often pursues XX of both species (Vincent 1949). NEST: spherical with side entrance, in grass tufts, generally within 20 cm of ground. Loosely woven of fine grass stems, with living grass coiled over to form substantial bower. Sited in areas of wet ground with dense growth of grass about 05 m tall. Y constructs frame, X contributes lining of dry grass, continues lining after eggs laid. EGGS: 1–3, av. 2 (n ¼ 11); of hartlaubi, pale greenish blue with spots, small blotches and patches of olive-grey, some chocolate-brown spots, mostly at broader end (Vincent 1949). [Description of eggs in Lees (1932) apparently an error for Yellow-crowned Bishop E. afer.] Eggs of E. h. humeralis pale olive-green, with irregular brown blotches and darker squiggles. SIZE: (n ¼ 16) av. 220  154. LAYING DATES: Gabon, (breeding plumage Feb); Uganda, Apr–May, July; Angola, Dec, Feb; S Zaı¨re, Jan–Mar; Tanzania, Jan–Feb; Zambia, Dec–Feb (enlarged gonads in Nov, breeding plumage Oct–Apr). No other information.

Key References

Lees, R. H. (1932), Vincent, A. W. (1949).

245

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PLOCEIDAE

Plate 15

Euplectes psammocromius (Reichenow). Mountain Marsh Widowbird. Euplecte montagnard.

(Opp. p. 218)

Penthetria psammocromia Reichenow, 1900. Orn. Monatsber., 8, p. 39; Tandala, Ukinga. Forms a superspecies with E. hartlaubi. Range and Status. Endemic resident. Restricted to Nyika ˆ i), and highlands of SW Plateau (NE Zambia, N Malaw Tanzania northeast to Njombe and Iringa, where locally common.

Euplectes psammocromius

Description. ADULT Y (breeding): wholly black except for wingcoverts; long black tail; wing has greater coverts with narrow buff margins, lesser coverts bright yellow, grading into buff on posterior border. Bill blue-black; eyes dark brown; legs blackish. ADULT Y (non-breeding): forehead to uppertail-coverts dark brown with buffy margins to feathers; tail brown. Bill dark brown; eyes brown; legs dark brown. ADULT X: like non-breeding Y, except that dark streaking on upperparts is less pronounced; below, dull white, not tawny; wing feathers dark brown with buff margins; no epaulet, but indistinct yellow margins to anterior wing-coverts. SIZE (19 YY, 9 XX): wing, Y 94–133 (103), X 76–83 (792); tail, Y (breeding, n ¼ 11) 212–294 (247), Y (non-breeding, n ¼ 2) 77, 79, X 52–62 (563); bill, Y 178–200 (187), X 163–180 (172); tarsus, Y 274– 292 (280), X 232–260 (247). IMMATURE and NESTLING: no information. TAXONOMIC NOTE: formerly treated as a race of Hartlaub’s Marsh Widowbird E. hartlaubi, but its specific status supported by habitat, geographic isolation, and distinct voices.

Field Characters. Length, breeding Y 35–50 cm, X 15– 18 cm. Black breeding Y recognized by bright yellow and buff shoulder patch, black back and long thin tail, which in display flight looks like several tails (Aspinwall and Beel 1998). Highlands, not meeting similar Hartlaub’s Marsh Widowbird E. hartlaubi. Non-breeding Y retains shoulder patches; X and sub-ad. much larger and more heavily streaked below than sympatric congeners. Voice. Tape-recorded (B, ASP, LEM). Call a whistling rattle, very different from that of Hartlaub’s Widowbird; song strikingly complicated and very high-pitched (Dowsett and Dowsett-Lemaire 1980). General Habits. Little known. Inhabits short, thick, montane grassland usually near streams; at 1800–3000 m ˆ i. In in S Tanzania, above 2100 m in Zambia and Malaw pairs or small groups, but forms large flocks in nonbreeding season.

Food. Grass seeds and insects.

Breeding Habits. Presumed to be polygynous and territorial. Y displays in slow flight, with long tail conspicuous. Nest a dome-shaped structure of fine grass, loosely woven with a side entrance; living grass bent over nest. Eggs 2, pale olive-green, densely marked with very fine brown spots, streaks and some squiggles. ˆ i, Oct–Nov; Tanzania, May; LAYING DATES: Malaw Zambia, (breeding dress Nov–Apr). Key Reference Aspinwall, D. R. and Beel, C. (1998).

Plate 15

Euplectes progne (Boddaert). Long-tailed Widowbird; Sakabula. Euplecte `a longue queue.

(Opp. p. 218)

Emberiza progne Boddaert, 1983. Tables Planch. enlum., p. 39; Cape of Good Hope. Range and Status. Endemic resident, with 3 discrete but often fragmented populations. Kenya, Tamau, Nanyuki, Naro Moru, Laikipia and Hyahururu to Nakuru, Elmenteita, Naivasha and Kinangop plateau, disjunct population around Eldoret and Kaptagat. Zambia, Bangweulu, NW Western Prov., SW North-West Prov. Angola, Huı´la, Huambo to southern Cuanza Sul, east to Lunda Norte, Lunda

Sul and Moxico; Zaı¨re, in S region, Kwango, Kwilu, Katanga, and Marungu plateau. Botswana, in SE near Lobatse, Gaborone; South Africa, from Northern and North-West Provs through KwaZulu-Natal, E Free State, Eastern Cape, with isolated western population south of Humansdorp; Swaziland, W plateau region; Lesotho, valleys and lower plateaux. Locally common.

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PLOCEIDAE

Plate 15

Euplectes psammocromius (Reichenow). Mountain Marsh Widowbird. Euplecte montagnard.

(Opp. p. 218)

Penthetria psammocromia Reichenow, 1900. Orn. Monatsber., 8, p. 39; Tandala, Ukinga. Forms a superspecies with E. hartlaubi. Range and Status. Endemic resident. Restricted to Nyika ˆ i), and highlands of SW Plateau (NE Zambia, N Malaw Tanzania northeast to Njombe and Iringa, where locally common.

Euplectes psammocromius

Description. ADULT Y (breeding): wholly black except for wingcoverts; long black tail; wing has greater coverts with narrow buff margins, lesser coverts bright yellow, grading into buff on posterior border. Bill blue-black; eyes dark brown; legs blackish. ADULT Y (non-breeding): forehead to uppertail-coverts dark brown with buffy margins to feathers; tail brown. Bill dark brown; eyes brown; legs dark brown. ADULT X: like non-breeding Y, except that dark streaking on upperparts is less pronounced; below, dull white, not tawny; wing feathers dark brown with buff margins; no epaulet, but indistinct yellow margins to anterior wing-coverts. SIZE (19 YY, 9 XX): wing, Y 94–133 (103), X 76–83 (792); tail, Y (breeding, n ¼ 11) 212–294 (247), Y (non-breeding, n ¼ 2) 77, 79, X 52–62 (563); bill, Y 178–200 (187), X 163–180 (172); tarsus, Y 274– 292 (280), X 232–260 (247). IMMATURE and NESTLING: no information. TAXONOMIC NOTE: formerly treated as a race of Hartlaub’s Marsh Widowbird E. hartlaubi, but its specific status supported by habitat, geographic isolation, and distinct voices.

Field Characters. Length, breeding Y 35–50 cm, X 15– 18 cm. Black breeding Y recognized by bright yellow and buff shoulder patch, black back and long thin tail, which in display flight looks like several tails (Aspinwall and Beel 1998). Highlands, not meeting similar Hartlaub’s Marsh Widowbird E. hartlaubi. Non-breeding Y retains shoulder patches; X and sub-ad. much larger and more heavily streaked below than sympatric congeners. Voice. Tape-recorded (B, ASP, LEM). Call a whistling rattle, very different from that of Hartlaub’s Widowbird; song strikingly complicated and very high-pitched (Dowsett and Dowsett-Lemaire 1980). General Habits. Little known. Inhabits short, thick, montane grassland usually near streams; at 1800–3000 m ˆ i. In in S Tanzania, above 2100 m in Zambia and Malaw pairs or small groups, but forms large flocks in nonbreeding season.

Food. Grass seeds and insects.

Breeding Habits. Presumed to be polygynous and territorial. Y displays in slow flight, with long tail conspicuous. Nest a dome-shaped structure of fine grass, loosely woven with a side entrance; living grass bent over nest. Eggs 2, pale olive-green, densely marked with very fine brown spots, streaks and some squiggles. ˆ i, Oct–Nov; Tanzania, May; LAYING DATES: Malaw Zambia, (breeding dress Nov–Apr). Key Reference Aspinwall, D. R. and Beel, C. (1998).

Plate 15

Euplectes progne (Boddaert). Long-tailed Widowbird; Sakabula. Euplecte `a longue queue.

(Opp. p. 218)

Emberiza progne Boddaert, 1983. Tables Planch. enlum., p. 39; Cape of Good Hope. Range and Status. Endemic resident, with 3 discrete but often fragmented populations. Kenya, Tamau, Nanyuki, Naro Moru, Laikipia and Hyahururu to Nakuru, Elmenteita, Naivasha and Kinangop plateau, disjunct population around Eldoret and Kaptagat. Zambia, Bangweulu, NW Western Prov., SW North-West Prov. Angola, Huı´la, Huambo to southern Cuanza Sul, east to Lunda Norte, Lunda

Sul and Moxico; Zaı¨re, in S region, Kwango, Kwilu, Katanga, and Marungu plateau. Botswana, in SE near Lobatse, Gaborone; South Africa, from Northern and North-West Provs through KwaZulu-Natal, E Free State, Eastern Cape, with isolated western population south of Humansdorp; Swaziland, W plateau region; Lesotho, valleys and lower plateaux. Locally common.

Euplectes progne

Euplectes progne

E. p. delacouri (Wolters) includes ‘definita’): Zaı¨re to Angola and Zambia. Breeding Y has upper mandible black rather than bluish, no white on epaulet and broader buff area, longer wing. SIZE: (breeding Y, n ¼ 26) wing 134–155 (146), tail 370–464 (425). WEIGHT: Y (Zaı ¨re, n ¼ 8) 41–49 (451); 2 XX (Zaı¨re) 32, 39; nonbreeding, 2 unsexed (Zambia) 330, 345. E. p. delamerei (Shelley): Kenya. Breeding Y longer-tailed. SIZE: (breeding Y, n ¼ 37) wing 129–152 (139), tail 416-628 (510). TAXONOMIC NOTE: application of the phylogenetic species concept would certainly lead to recognition of 3 species, since there is clearly no interchange between these populations.

Field Characters. Length, breeding Y 61–71 cm, nonbreeding Y 19–23 cm, X 15–20 cm. Breeding Y unmistakable, with tail so long it looks unreal; bill heavy and white (small and black in Red-collared Widowbird E. ardens), shoulders red and white (tawny brown in Jackson’s Widowbird E. jacksoni). Non-breeding Y like large version of Fan-tailed Widowbird E. axillaris, much the biggest of the streaked Euplectes spp., with floppy flight on broad rounded black wings, showing red and buff epaulets; tail variable in length, feathers pointed (rounded in Fan-tailed Widowbird). X and sub-ad. Y have narrower streaking on back than Fan-tailed Widowbird, whiter and more heavily streaked underparts than Jackson’s; best separated from Jackson’s and Hartlaub’s Marsh E. hartlaubi Widowbirds by uniformly dark underwing-coverts. Description. E. p. progne (Boddaert): Botswana, South Africa, Lesotho, Swaziland. ADULT Y (breeding): wholly black except for wings; feathers of crown and nape with squared tips and glossy edges, a few ventral feathers also of this type; tail very long and black; wings black but primaries with pale tips, secondaries and coverts with narrow buff margins, lesser coverts orange-red bordered with white or buff at tip, forming an epaulet; underwing sooty-black. Bill dark blue-grey; eyes brown; legs dark brown. ADULT Y (non-breeding): retains black wing feathers and red-andwhite epaulet, but much broader buff margins on secondaries and greater coverts. Forehead to uppertail-coverts brown with dark central streaks; tail short, brown with paler margins, feathers pointed; pale whitish superciliary stripe, lores, cheeks and earcoverts brown; chin and throat whitish; breast and belly buff to whitish with some central streaks on breast and flank feathers; thighs and undertail-coverts buff. Bill brown in birds which have just acquired black wing feathers, otherwise retains some bluish coloration. SUB-ADULT Y: like non-breeding Y, except for brown wing feathers with paler margins, coverts forming epaulet have narrow orange-brown margins (no red, white or buff). Bill brown. In captivity, Y took 3 years to acquire ad. plumage; Y at 2 years may show full gonadal development, but in field also only acquires breeding plumage in 3rd year (Verheyen 1953, Traylor 1965). ADULT X: like sub-ad. Y, but coloured edging to feathers on epaulet very indistinct. SIZE (82 ad. YY, 37 sub-ad. YY, 28 XX): wing, ad. Y 122–161 (138), sub-ad. Y 95–116 (108), X 87–97 (908); tail, ad. Y (breeding, n ¼ 82) 319–499 (411), ad. Y (non-breeding, n ¼ 27) 80–116 (950), sub-ad. Y 64–102 (779), X 54–75 (623); bill, ad. Y 174–207 (190), sub-ad. Y 173–197 (187), X 166–184 (175); tarsus, ad. Y 269–306 (286), sub-ad. Y 264–306 (285), X 231–256 (245). In all populations, Y wing-length and apparently also tail-length increase with age, perhaps over several years (Verheyen 1956, Traylor 1965, Craig 1989). WEIGHT: ad. Y (South Africa n ¼ 29) 33–46 (416); sub-ad. Y 37, 469; 1 juv. Y 35; ad. X (n ¼ 32) 25–39 (320). IMMATURE: like ad. X, but broad buffy edges to feathers on upperparts. NESTLING: not described.

Voice. Tape-recorded (58, 72, 86, 88, 99, 104, B, F). Typical song a rapid series of repeated notes on one pitch, ranging in tone from dry rattle to liquid ringing trill, with or without a few introductory notes; other songs slower, ‘tsi-tew-tewtew-tew-tew . . . ’ or ‘cha-pss-chay-chay-chay-chay’; also rendered ‘twi-twi-twi-twi-zizizizi’ (Maclean 1993). Makes low paper-tearing sound, which may be the ‘weezle weezle’ described as territorial song. Displays of captive Y punctuated by loud swizzling, blowing and twittering sounds; commonest call was ‘chippit-chippit’ in bursts, also ‘cheerit’ in response to other birds or people approaching; plaintive mewing sound during some displays (Brown et al. 1979). Contact call (both sexes) a sharp ‘chip’, ‘jit’ or ‘zik’, repeated and sometimes accelerated into series, becoming more intense as alarm, ‘tseek-tseek’ or ‘tscheep-tscheep’. Sub-adult Y makes quiet husky ‘chikker-chikker’. General Habits. Inhabits grasslands, often where grass short; not necessarily in marshy areas. Highlands: at 1800– 2800 m in Kenya and up to 2750 m in southern Africa; but also locally on coastal grasslands and in other lowland areas in central Africa. Feeds largely on ground; walks and runs. Y leaves territory to feed. Hawks flying termites and butterflies. Forms flocks, in Kenya with Jackson’s Widowbird E. Jacksoni and Red-billed Quelea Quelea quelea; in South Africa with Southern Red Bishop E. orix, Redcollared Widowbird E. ardens, Red-billed Quelea and Pintailed Whydah Vidua macroura. Roosts communally in marshes even when breeding (sometimes with Fan-tailed Widowbird E. axillaris) or in clumps of dense, high Eleusine jaegeri grass which may hold >1000 birds of several species; one roost in South Africa held c. 1 million Barn Swallows Hirundo rustica, several thousand Red-billed Quelea, and several hundred other ploceids including

247

248

PLOCEIDAE

>200 Long-tailed Widowbirds. YY going to roost in late afternoon fly high and fast; claims that Y with long tail is unable to fly in heavy rain or wind appear to be unfounded. Flight of Y undulating, in breeding and non-breeding season. On ground, long-tailed bird runs with short rapid steps, but hops if surface is uneven. Sub-adult YY not tolerated on territory, but they form groups of their own and carry out similar displays, even flying towards other birds with slow, measured wing-beats and the short tail depressed below level of body. In perched display, subad. Y tenses body, half opens wings and fans tail white swaying from side to side for several minutes, even when no other birds present. Moult of primaries follows typical passerine pattern, but elongated tail feathers seem to be shed simultaneously. Anting noted in captive birds (Poulsen 1956). Food. Seeds of Setaria flabellate and Triticum, also Paspalum dilatatum, P. distichum, Pennisetum clandestinum, Themeda triandra, achenes of Senecio juniperinus. In Free State, South Africa, greatest variety of seed types taken in spring and winter. Eats scarab beetles, feeds chicks with larvae, cicadas, aphids, beetles and spiders. In captivity takes mealworms, termites, grasshoppers, caterpillars, moths, spiders. Breeding Habits. Polygynous, Y with up to 5 XX; territorial, Y defends area of 05–30 ha, which may contain up to 5 nests. Y has favoured perches, often no more than 45–75 cm above ground: fence posts, termite mounds or small, scrubby Acacia trees. From perch, Y advertises presence with ‘tsik’ call, to which YY on adjoining territories reply. Any conspecific Y entering territory is driven out, escorted in fast flight with chaser’s tail streaming out behind (A). However, does not interact with Y Fantailed Widowbird where territories overlap. When group of brown-plumaged birds approaches, Y sets off in slow flight (B) with keeled tail (‘Cascade-tail display’: Skead 1995), at times hovering (C), calling a rolling, husky ‘wheezlewheezle’. If X lands in territory, Y may alight nearby and bob up and down with crown and nape feathers erected. Experimental manipulation of Y tail-length suggests that X may select longest-tailed Y as her mate (Andersson 1982); X evidently visits several territories in succession. Captive Y displays (to other species in aviary) with feathers on nape and crown erected, producing hooded or cowled appearance; on perch, struts with head held low, wings extended downwards and sometimes flapped slowly. Aerial display flights follow figure-of-eight course with tail trailing; bird also performs slow flights with keeled tail (Brown et al. 1979). Captive bird attacked Y Red-collared Widowbird and other red-plumaged birds in same aviary. Before copulating, Y stretches body upwards, sways rapidly from side to side in front of X; during coitus he spreads wings, enclosing her (Norris 1968). NEST: woven of fine grass including growing green material; lined with flowering grass heads, which may project to form short porch over side entrance. Sited close to ground in tuft of grass 38–60 cm tall, with blades of grass bent over and linked together above nest to form a bower; height above ground (n ¼ 42, Kenya) 22–53 (36) cm. Y may make a simple nest ring; X constructs nest unaided, within

Y’s territory. Captive Y once built a typical nest. Y gives alarm calls when predators approach, but does not approach nest. X continues to add lining of fine grass leaves and roots during incubation, until nest is densely padded. Nests may be as little as 1 m apart; clumping appears to be result of patchy habitat. EGGS: 1–4 (av. 21, n ¼ 109, Kenya). Greenish white or blue-green, heavily speckled with grey and olive-brown. Some with much larger flecking, or mesh of brown markings so close that little ground colour visible. SIZE: (n ¼ 29, South

Euplectes jacksoni Africa) 201–240  141–158 (219  159), (n ¼ 31, Kenya) av. 234  160. LAYING DATES: Zaı ¨re, Dec–Apr; Kenya, Apr–May (Brown and Britton 1980), Nov–Jan (Hake and Andersson 1992); Angola, Feb (breeding plumage Dec); Zambia, (breeding condition Feb); South Africa, Oct–Mar. INCUBATION: by X only, period 12–14 days. In aviary, X approached nest by well-defined path with grass drawn over it to form a tunnel. DEVELOPMENT AND CARE OF YOUNG: only X feeds young: 16 feeds per h per nestling, mainly grass seeds but also insects (c. 25% by volume). Eyes open after 5–6 days; first quills appear after 4–5 days, feathers opening out at 10–12 days. All droppings removed from nest. Nestling period 17

249

days, 17–20 days in captivity. Chicks can fly short distances at 15 days; leave nest before they can fly strongly, and remain concealed in grass. Captive young fed by X for 14 days after leaving nest: In captive brood, YY markedly larger than XX on fledging. BREEDING SUCCESS/SURVIVAL: of 36 nests in Kenya, 22 destroyed and success rate calculated at 16%; people, raptors, rodents, driver ants and flooding accounted for some losses. Y in Scho ¨nbrunn Zoo survived 17 years 11 months, developed at least partial breeding plumage every year, but flightless in final 2 years (Schifter 1970). Key References Andersson, M. (1982), Craig, A. J. F. K. (1989), Hake, M. and Andersson, M. (1992).

Euplectes jacksoni (Sharpe). Jackson’s Widowbird. Euplecte de Jackson.

Plate 15

Drepanoplectes jacksoni Sharpe, 1891. Ibis, 1891, p. 246; Kikuyu, Kenya.

(Opp. p. 218)

Range and Status. Endemic resident, restricted to highland grassland in Kenya and N Tanzania. Kenya, from Eldoret and Nandi east to Laikipia and Mt Kenya, south to Aberdares, Nairobi Nat. Park, Ngong Hills, Mau, Mara Game Res., Loita and Nguruman Hills; NE Tanzania, in Crater Highlands and Loliondo. Locally common, but considered near threatened, as habitat is being increasingly fragmented by agricultural development. Description. ADULT Y (breeding): plumage entirely black except for wings; tail feathers long and curved. Feathers of crown and nape have squared ends, forming a distinctive cape. Wing dark brown, feathers edged with buff, except for black tertials; epaulet orange-buff, greater coverts with orange-buff edging; underwing orange-buff. Upper mandible pale steel-blue, darker near nostrils, lower mandible dark with pale bluish tip; eyes dark brown; legs black. ADULT Y (non-breeding): forehead, crown, nape, mantle, rump and uppertail-coverts dark brown with buffy margins; tail dark brown with pale edges, the feathers distinctly pointed; broad buff superciliary stripe, lores and cheeks buff-brown; chin and throat whitish, breast buff with narrow, dark central streaks on feathers, belly whitish with dark streaks on flanks, thighs and undertail-coverts buff; epaulet orange-brown. Bill bluish, darker than in breeding state, especially upper mandible; eyes dark brown; legs brown. SUB-ADULT Y: like non-breeding Y, but bill completely brown, upper mandible darker, lower mandible paler. ADULT X: like sub-ad. Y, but smaller. SIZE (56 ad. YY, 14 sub-ad. YY, 21 XX): wing, Y 82–94 (899), sub-ad. Y 80–89 (848), X 74– 80 (776); tail, Y (breeding, n ¼ 26) 190–234 (212), Y (nonbreeding, n ¼ 17) 52–75 (613), sub-ad. Y 51–66 (556), X 46–56 (520); bill, Y 173–195 (183), sub-ad. Y 170–192 (180), X 161– 180 (170); tarsus, Y 276–312 (295), sub-ad. Y 274–310 (295), X 247–283 (264). WEIGHT: Y (n ¼ 212) av. 427; (n ¼ 13) 400– 485 (423); X (n ¼ 80) av. 297, (n ¼ 21) 290–420 (362). IMMATURE: like ad. X, but smaller. NESTLING: on hatching, pink-brown with sparse grey tufts of down toward back of head, at base of wings, and in dorsal and lumbar regions (van Someren 1956).

Field Characters. Length, breeding Y 28–30 cm, X 14 cm. Breeding Y black with pale bill; differs from sympatric Long-tailed Widowbird E. progne in shorter decurved tail of broad feathers, tawny epaulet. Non-breeding birds best

Euplectes jacksoni

identified by orange underwing (dark in Long-tailed and Hartlaub’s Marsh E. hartlaubi Widowbirds); Y larger than any Euplectes except Long-tailed Widowbird, retains black wing with tawny shoulder and feathers edges; X pale buff to orange-buff below, with heavy bill. Voice. Tape-recorded (B, C, McVIC, NOR). Y gives wheezing sizzling song during display, with rattle probably from wings. Song from perch ‘sisi-si glip-glip-glip-glip’; contact call rapidly repeated ‘tu’ (Moreau and Sclater 1938). Sharp ‘chip’ or ‘jit’ from perched bird, recorded by M. E. W. North, accelerating and getting louder as it flew off, may have been alarm call. In flight, soft ‘tu’. X

Euplectes jacksoni Africa) 201–240  141–158 (219  159), (n ¼ 31, Kenya) av. 234  160. LAYING DATES: Zaı ¨re, Dec–Apr; Kenya, Apr–May (Brown and Britton 1980), Nov–Jan (Hake and Andersson 1992); Angola, Feb (breeding plumage Dec); Zambia, (breeding condition Feb); South Africa, Oct–Mar. INCUBATION: by X only, period 12–14 days. In aviary, X approached nest by well-defined path with grass drawn over it to form a tunnel. DEVELOPMENT AND CARE OF YOUNG: only X feeds young: 16 feeds per h per nestling, mainly grass seeds but also insects (c. 25% by volume). Eyes open after 5–6 days; first quills appear after 4–5 days, feathers opening out at 10–12 days. All droppings removed from nest. Nestling period 17

249

days, 17–20 days in captivity. Chicks can fly short distances at 15 days; leave nest before they can fly strongly, and remain concealed in grass. Captive young fed by X for 14 days after leaving nest: In captive brood, YY markedly larger than XX on fledging. BREEDING SUCCESS/SURVIVAL: of 36 nests in Kenya, 22 destroyed and success rate calculated at 16%; people, raptors, rodents, driver ants and flooding accounted for some losses. Y in Scho ¨nbrunn Zoo survived 17 years 11 months, developed at least partial breeding plumage every year, but flightless in final 2 years (Schifter 1970). Key References Andersson, M. (1982), Craig, A. J. F. K. (1989), Hake, M. and Andersson, M. (1992).

Euplectes jacksoni (Sharpe). Jackson’s Widowbird. Euplecte de Jackson.

Plate 15

Drepanoplectes jacksoni Sharpe, 1891. Ibis, 1891, p. 246; Kikuyu, Kenya.

(Opp. p. 218)

Range and Status. Endemic resident, restricted to highland grassland in Kenya and N Tanzania. Kenya, from Eldoret and Nandi east to Laikipia and Mt Kenya, south to Aberdares, Nairobi Nat. Park, Ngong Hills, Mau, Mara Game Res., Loita and Nguruman Hills; NE Tanzania, in Crater Highlands and Loliondo. Locally common, but considered near threatened, as habitat is being increasingly fragmented by agricultural development. Description. ADULT Y (breeding): plumage entirely black except for wings; tail feathers long and curved. Feathers of crown and nape have squared ends, forming a distinctive cape. Wing dark brown, feathers edged with buff, except for black tertials; epaulet orange-buff, greater coverts with orange-buff edging; underwing orange-buff. Upper mandible pale steel-blue, darker near nostrils, lower mandible dark with pale bluish tip; eyes dark brown; legs black. ADULT Y (non-breeding): forehead, crown, nape, mantle, rump and uppertail-coverts dark brown with buffy margins; tail dark brown with pale edges, the feathers distinctly pointed; broad buff superciliary stripe, lores and cheeks buff-brown; chin and throat whitish, breast buff with narrow, dark central streaks on feathers, belly whitish with dark streaks on flanks, thighs and undertail-coverts buff; epaulet orange-brown. Bill bluish, darker than in breeding state, especially upper mandible; eyes dark brown; legs brown. SUB-ADULT Y: like non-breeding Y, but bill completely brown, upper mandible darker, lower mandible paler. ADULT X: like sub-ad. Y, but smaller. SIZE (56 ad. YY, 14 sub-ad. YY, 21 XX): wing, Y 82–94 (899), sub-ad. Y 80–89 (848), X 74– 80 (776); tail, Y (breeding, n ¼ 26) 190–234 (212), Y (nonbreeding, n ¼ 17) 52–75 (613), sub-ad. Y 51–66 (556), X 46–56 (520); bill, Y 173–195 (183), sub-ad. Y 170–192 (180), X 161– 180 (170); tarsus, Y 276–312 (295), sub-ad. Y 274–310 (295), X 247–283 (264). WEIGHT: Y (n ¼ 212) av. 427; (n ¼ 13) 400– 485 (423); X (n ¼ 80) av. 297, (n ¼ 21) 290–420 (362). IMMATURE: like ad. X, but smaller. NESTLING: on hatching, pink-brown with sparse grey tufts of down toward back of head, at base of wings, and in dorsal and lumbar regions (van Someren 1956).

Field Characters. Length, breeding Y 28–30 cm, X 14 cm. Breeding Y black with pale bill; differs from sympatric Long-tailed Widowbird E. progne in shorter decurved tail of broad feathers, tawny epaulet. Non-breeding birds best

Euplectes jacksoni

identified by orange underwing (dark in Long-tailed and Hartlaub’s Marsh E. hartlaubi Widowbirds); Y larger than any Euplectes except Long-tailed Widowbird, retains black wing with tawny shoulder and feathers edges; X pale buff to orange-buff below, with heavy bill. Voice. Tape-recorded (B, C, McVIC, NOR). Y gives wheezing sizzling song during display, with rattle probably from wings. Song from perch ‘sisi-si glip-glip-glip-glip’; contact call rapidly repeated ‘tu’ (Moreau and Sclater 1938). Sharp ‘chip’ or ‘jit’ from perched bird, recorded by M. E. W. North, accelerating and getting louder as it flew off, may have been alarm call. In flight, soft ‘tu’. X

250

PLOCEIDAE

approaching nest gave low ‘chip chio chio’, longer ‘cheiiio’ on leaving (van Someren 1956). General Habits. Inhabits open grassland at 1500–3000 m, primarily coarse Eleusine jaegeri or Themeda triandra; always gregarious, and forages in mixed flocks even when YY are in breeding plumage. Large flocks may damage crops in rural smallholdings. Often sympatric with Long-tailed Widowbird. Anting recorded in captive birds (Poulsen 1958). Breeding areas traditional, and same sites may be used for a number of years. Sub-ad. Y displays at any vacant dancing ring, but displays are incomplete, and not focused on central tuft as is the case in ad. Y. Perched Y nips off grass stems near it, even when not on dancing grounds. Although lengthy tail seems a handicap, Y is quite agile in flight (Loveridge 1922); on ground, runs with rapid short steps, hopping if the surface is uneven. Food. Grass seeds, particularly Themeda triandra, also Panicum. Eats termite alates. Breeding Habits. Very well known from dancing display of Y. Polygynous and highly territorial. Y makes a dancing display site, a lek, which X visits for mating only. Y in display flight arches tail when flying over his territory, which may include 2–3 dancing rings. Dancing ring is a circle of flattened grass c. 60 cm in diam., grass nipped off and trodden down by Y, surrounding a central tuft. Path formed by activity of Y may be down to bare earth. Central tuft is shaped to a dome by Y nipping off most of the tall grass blades; typically 20 cm tall, by 10 cm wide; ovalshaped, and on each side of its longest axis Y makes a shallow cup-shaped recess, 1 usually deeper than the other. Recesses are formed by Y taking short runs at tuft with lowered head; he then presses into them and smooths them out with breast. Collar erected in aggressive display; intruding Y with folded collar and tail will be ignored. Y attends dancing ring from mid-morning to late

afternoon, then leaves for feeding areas. He never perches on central tuft but sings from perch in tall grass next to ring (A). When dancing, Y stands on ring facing the tuft and jumps energetically to various heights from a few cm to almost 1 m. When jumping, his head is thrown back, neck feathers ruffled, tail plumes arched upwards and forwards to almost touch back of head, except for outer pair of feathers which hang downwards; his feet beat rapidly and half-open wings are quivered (B, C). Soft tinkling call accompanies jumps. After 5–6 jumps, Y pauses to rest. X flying over or watching from grass may then approach him. If X lands on ring, Y positions himself so that central tuft is between him and her. In ground display Y quivers vigorously, head back, hackles ruffled, body plumage fluffed and tail vertically spread and fluttering (D); accompanied by a tinkling call. After 50 s, Y may hop around tuft to approach X, then retreat; in final stiff approach he gives a distinctive gurgling call, used at no other stage: a prelude to copulation. Only 1 X is courted at a time; Y becomes agitated and his display is disrupted if a 2nd X arrives; both XX are then displaced in aggressive display. Y continues to display at dancing ring lacking central tuft, and X may land there briefly, but copulation does not occur at such modified rings (Andersson 1991). Dancing affected by gusty winds, and display is sporadic on windy days. Mating success of YY varies greatly; 22 did not mate, 6 mated once, 2 three times, 1 four times, 1 five times. Attraction of X depends on display intensity and Y’s time spent at lek, white tail length is apparently main

ESTRILDIDAE

factor in X’s choice of mate. Y tail length may correlate with body condition (Andersson 1989). Copulation normally occurs after X has spent >1 min at the ring; X solicits with slightly drooped, quivering wings. NEST: domed ball of woven grass with side entrance, with living grass bent down over it to form a bower. Lined with grass seedheads, often Panicum. Placed within 10 cm of ground in any grass tuft 50 cm tall; nests sometimes clustered, with >20 in a small area. Site not related to Y territory and is outside lek area, but within 300 m of it. Built by X only.

EGGS: 2–4; oval pale blue, greyish or greenish ground with dense, fine grey and brown flecks and squiggles. SIZE: 21–22  14–15 (van Someren 1956); av. (n ¼ 8) 225–154 (Serle 1943a). LAYING DATES: Kenya, Jan, Apr–June, Aug, Sept, Nov (Nairobi, displays Mar–Aug, dependent on rains; Naro Moru, lek activity peaks in Dec–Jan). INCUBATION: by X only, period 12–13 days; in captivity, 12 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only, on regurgitated grass seeds. Nest visited once per h, average stay 6 min, with c. 10 regurgitations. Faecal sacs removed by X and dropped in flight. Fledgling period 17 days; young initially hop through grass and herbs; roost with X. In captivity, apparently fed only on seeds; fledge at 16 days. Y does not attend nest, but alarm call on disturbance causes X to leave nest inconspicuously. BREEDING SUCCESS/SURVIVAL: nests in pastures may be trampled by cattle; X leaves nest without any defensive or distractive display. X deserts readily in early stages of incubation (van Someren 1956). Y recaptured 5 years after ringing, on new display ground 12 km from original site.

Key References Andersson, S. (1982, 1989, 1991, 1992, 1993), van Someren, V. D. (1946).

Family ESTRILDIDAE: firefinches, waxbills, grassfinches, munias, mannikins and others

Small, strongly patterned and coloured Old World finch-like granivores, mainly tropical, mostly gregarious; some in forest, most in bushy savanna. Most species with pink, red or carmine rumps and often faces, tails and underparts. Bill stout and conical, massive and notched in Spermophaga, small in Nigrita, slender in Parmoptila; generally bright red or silvery. Wings round-tipped; outermost primary (P10) reduced. Tail often attenuated with long, pointed central rectrices; otherwise medium, slightly rounded. Legs and feet black, grey or brown, never particularly strong. Most species perch readily and nest in woody and herbaceous vegetation, and feed in grass heads and on ground. Ortygospiza and Paludipasser are exclusively terrestrial. Eat ripe seeds, mainly small; large and hard in a few species; sometimes unripe; also insects in breeding season. Nigrita and Parmoptila insectivorous, latter eating ants. Drink by dipping bill into water then tilting head up and swallowing, also by sucking, with bill kept in water. Sucking is employed, particularly by Estrilda and Uraeginthus spp., when only very shallow water is available, or when bird drinks from rain droplets on leaves (I. Hinze, pers. comm.). Flight often whirring. Not migratory, but many species nomadic. Popular cagebirds; breeding biology studied better in captivity than in the wild. Complex, ritualized courtship, nestbuilding, nestling and parental behaviours. Considerable variety of vocalizations. Courting YY often carry piece of nesting material in bill and displays often involve tail movements. Many species sing during courtship; song not loud, sometimes inaudible only 1 m away. Copulation nearly always initiated by X. Nest domed, with side entrance and often a tunnel; untidy; made of grass stems and leaves, never woven. YY usually collect and carry nest material, XX usually sit in nest and build it; otherwise parents share more or less equally in duties. A few species nest in abandoned nests of weavers (Ploceidae). Several build roosting nest, and (Estrilda) ‘cock’s nest’. Nestlings with strongly patterned palate, tongue, floor of mouth, and papillae (tubercles) at corner of mouth (illustrated on p. 252); mouth brightly coloured in several species. Papillae brightly and reflectively coloured in some species, black and white in others. Mouth markings made even more conspicuous by lateral waving movements of tongue and entire head. Many nestlings solicit food by keeping head low in nest and twisting neck to one side so that wide-open mouth points upward; young fed by regurgitation. Many African species brood-parasitized by whydahs and indigobirds, Viduidae.

251

ESTRILDIDAE

factor in X’s choice of mate. Y tail length may correlate with body condition (Andersson 1989). Copulation normally occurs after X has spent >1 min at the ring; X solicits with slightly drooped, quivering wings. NEST: domed ball of woven grass with side entrance, with living grass bent down over it to form a bower. Lined with grass seedheads, often Panicum. Placed within 10 cm of ground in any grass tuft 50 cm tall; nests sometimes clustered, with >20 in a small area. Site not related to Y territory and is outside lek area, but within 300 m of it. Built by X only.

EGGS: 2–4; oval pale blue, greyish or greenish ground with dense, fine grey and brown flecks and squiggles. SIZE: 21–22  14–15 (van Someren 1956); av. (n ¼ 8) 225–154 (Serle 1943a). LAYING DATES: Kenya, Jan, Apr–June, Aug, Sept, Nov (Nairobi, displays Mar–Aug, dependent on rains; Naro Moru, lek activity peaks in Dec–Jan). INCUBATION: by X only, period 12–13 days; in captivity, 12 days. DEVELOPMENT AND CARE OF YOUNG: fed by X only, on regurgitated grass seeds. Nest visited once per h, average stay 6 min, with c. 10 regurgitations. Faecal sacs removed by X and dropped in flight. Fledgling period 17 days; young initially hop through grass and herbs; roost with X. In captivity, apparently fed only on seeds; fledge at 16 days. Y does not attend nest, but alarm call on disturbance causes X to leave nest inconspicuously. BREEDING SUCCESS/SURVIVAL: nests in pastures may be trampled by cattle; X leaves nest without any defensive or distractive display. X deserts readily in early stages of incubation (van Someren 1956). Y recaptured 5 years after ringing, on new display ground 12 km from original site.

Key References Andersson, S. (1982, 1989, 1991, 1992, 1993), van Someren, V. D. (1946).

Family ESTRILDIDAE: firefinches, waxbills, grassfinches, munias, mannikins and others

Small, strongly patterned and coloured Old World finch-like granivores, mainly tropical, mostly gregarious; some in forest, most in bushy savanna. Most species with pink, red or carmine rumps and often faces, tails and underparts. Bill stout and conical, massive and notched in Spermophaga, small in Nigrita, slender in Parmoptila; generally bright red or silvery. Wings round-tipped; outermost primary (P10) reduced. Tail often attenuated with long, pointed central rectrices; otherwise medium, slightly rounded. Legs and feet black, grey or brown, never particularly strong. Most species perch readily and nest in woody and herbaceous vegetation, and feed in grass heads and on ground. Ortygospiza and Paludipasser are exclusively terrestrial. Eat ripe seeds, mainly small; large and hard in a few species; sometimes unripe; also insects in breeding season. Nigrita and Parmoptila insectivorous, latter eating ants. Drink by dipping bill into water then tilting head up and swallowing, also by sucking, with bill kept in water. Sucking is employed, particularly by Estrilda and Uraeginthus spp., when only very shallow water is available, or when bird drinks from rain droplets on leaves (I. Hinze, pers. comm.). Flight often whirring. Not migratory, but many species nomadic. Popular cagebirds; breeding biology studied better in captivity than in the wild. Complex, ritualized courtship, nestbuilding, nestling and parental behaviours. Considerable variety of vocalizations. Courting YY often carry piece of nesting material in bill and displays often involve tail movements. Many species sing during courtship; song not loud, sometimes inaudible only 1 m away. Copulation nearly always initiated by X. Nest domed, with side entrance and often a tunnel; untidy; made of grass stems and leaves, never woven. YY usually collect and carry nest material, XX usually sit in nest and build it; otherwise parents share more or less equally in duties. A few species nest in abandoned nests of weavers (Ploceidae). Several build roosting nest, and (Estrilda) ‘cock’s nest’. Nestlings with strongly patterned palate, tongue, floor of mouth, and papillae (tubercles) at corner of mouth (illustrated on p. 252); mouth brightly coloured in several species. Papillae brightly and reflectively coloured in some species, black and white in others. Mouth markings made even more conspicuous by lateral waving movements of tongue and entire head. Many nestlings solicit food by keeping head low in nest and twisting neck to one side so that wide-open mouth points upward; young fed by regurgitation. Many African species brood-parasitized by whydahs and indigobirds, Viduidae.

251

252

ESTRILDIDAE

Nigrita canicapillus

Mandingoa nitidula

Estrilda astrild

Pyrenestes ostrinus

Uraeginthus angolensis

Uraeginthus bengalus

Granatina granatina

Hypargos niveoguttatus

Clytospiza monteiri

Lagonosticta rufopicta

Lagonosticta larvata

Lagonosticta rara

Amadina fasciata

Ortygospiza fuscocrissa

Spermestes cucullatus

Estrilda nonnula

Spermophaga haematina

Uraeginthus cyanocephalus Granatina ianthinogasterr

Pytilia phoenicoptera

Pytilia melba

Lagonosticta rubricata Lagonosticta rhodopareia

Spermestes bicolor

Spermestes fringilloides

Mouth markings and tubercles of nestlings and fledglings of some estrildids and viduids.

Sporaeginthus subflavus

Vidua obtusa

Vidua chalybeata

Vidua larvaticola

Nigrita luteifrons

253

Quite closely related with Viduidae, far more distantly with weavers Ploceidae and finches Fringillidae. Within Estrildidae, species relationships have been controversial, but recent molecular studies by M. Sorenson and R. B. Payne, (pers. comm.) suggest a phylogenetic arrangement as follows: Paludipasser locustella is only distantly related with all other estrildids, which fall into 5 clusters. Cluster (a), entirely African: Nigrita, Parmoptila; Nesocharis, Coccopygia; Mandingoa, Cryptospiza; Estrilda. Cluster (b), entirely African: Spermophaga, Pyrenestes; Uraeginthus, Granatina; Euschistospiza, Hypargos; Clytospiza, Pytilia, Lagonosticta. Cluster (c), mainly African: Amadina, Ortygospiza, Amandava. Cluster (d), 12 genera of Australasian grassfinches. Cluster (e), Oriental, Australasian and African: Odontospiza and Spermestes (Africa) with Heteromunia pectoralis (Australia); Euodice (Africa, India); Lonchura (Orient, Australasia) and Padda (Orient); and Lemuresthes nana (Madagascar). Sequences within each cluster can be seen as roughly evolutionary, from ‘ancestral’ to derived forms. We treat African genera in this sequence, except that we place Paludipasser after Ortygospiza. From the Asiatic Amandava we separate the single African species into Sporaeginthus. 137 species in 27 genera recognized by Goodwin (1982) and 140 in 27 genera by Sibley and Monroe (1990). We recognize 76 species in 25 genera in Africa; 21 genera endemic, 1 near-endemic (Estrilda), 1 shared with S Asia (Euodice) and 2 introduced from Asia (Amandava, Padda).

Genus Nigrita Strickland

Bill short and slender, with well-rounded culmen, usually somewhat flattened and expanded around nostrils; wings more pointed than rounded; tail black, fan-shaped, average length; plumage black and grey in 2 spp., grey and chestnut in 1, brown and white with black cap in 1; underparts uniform, no distinction between breast and belly. White wing spots in 1 species, N. canicapillus; these are single spots at tip of feather, as in Asian Amandava amandava, not paired as in twinspots. Palate and tongue marks estrildine, perhaps closest to Parmoptila, Cryptospiza and Pyrenestes (Goodwin 1982). See also Family diagnosis Occupies middle and upper levels in forest, often in canopy, not in low growth like most estrildines. Eats small insects and fruits. Bulky estrildine nest with side entrance. Endemic, 4 species. N. canicapillus and N. luteifrons are similarly plumaged but widely sympatric; unspotted W African race N. c. emiliae and white-backed Kungwe race N. c. candidus are divergent, perhaps incipient species (Hall and Moreau 1970). N. fusconotus, with more slender bill and more strongly graduated tail, was placed by Wolters (1966) in subgenus Percnopis.

Nigrita luteifrons Verreaux and Verreaux. Pale-fronted Negrofinch. Nigrette a ` front jaune

Plate 16

Nigrita luteifrons J. and E. Verreaux, 1851. Rev. Mag. Zool. (Paris), ser. 2, 3, p. 420; Gabon.

(Opp. p. 219)

Range and Status. Endemic resident. Sierra Leone (locality? G. D. Field in Gatter 1997); Liberia (once, Zlehtown, but probably rare and overlooked: Gatter 1997); Ivory Coast (Taı¨ Nat. Park and areas outside it: Gartshore et al. 1995); Ghana (Amedzofe, 0
520 N, 020
E); Togo, once at Misaho ¨he; Nigeria in forest zone, north to Ibadan, Ife and Okpo, uncommon, but sometimes outnumbers more abundant Grey-headed Negrofinch Nigrita canicapillus (Lagos: Gee and Heigham 1977); forest zone of S Cameroon from Ave´le´, Yaounde´ (seen 9 months of the year: Quantrill and Quantrill 1998) and Lobe´ke´ Faunal Res. north to Korup Nat. Park (few records), Limbe (not uncommon), Mt Cameroon (50–1300 m), Mt Kupe´ (uncommon, 750– 1000 m) and Bamenda, and in E to c. 5
300 N; Mbini, lowlands, and on Mt Alen at 500–950 m; Bioko; Gabon, widespread but not common; in NE even more infrequent than Grey-headed Negrofinch, in Lope´ Reserve confined to

borders of gallery forest; widespread in Congo (Nouabale´Ndoki and Odzala Nat. Parks, Le´fini, Mouilou Basin, Mayombe), outnumbering Grey-headed Negrofinch in Odzala (Dowsett and Dowsett-Lemaire 1997); Central African Republic around Bangui and in Lobaye Pre´fecture; not yet recorded in Dzanga Reserves in SW (Green and Carroll 1991) but almost certain to occur there since present across the borders in Cameroon and Congo; widespread but uncommon in Zaı¨re, from Ubangi, Uele and Semliki south to Mayombe, Kwango and Kasai (Luluabourg), and in E to Lubena west of L. Edward at 1000 m, Irangi Forest (Kizungu 2001) and Itombwe (to 1480 m); extreme W Uganda in Bwamba; uncommon Angola (Cabinda, and N Cuanza Norte at Canzele and Quiculungo). Formerly thought to be confined to Lower Guinea forests (e.g. Louette 1981) but recent discoveries in Upper

Nigrita luteifrons

253

Quite closely related with Viduidae, far more distantly with weavers Ploceidae and finches Fringillidae. Within Estrildidae, species relationships have been controversial, but recent molecular studies by M. Sorenson and R. B. Payne, (pers. comm.) suggest a phylogenetic arrangement as follows: Paludipasser locustella is only distantly related with all other estrildids, which fall into 5 clusters. Cluster (a), entirely African: Nigrita, Parmoptila; Nesocharis, Coccopygia; Mandingoa, Cryptospiza; Estrilda. Cluster (b), entirely African: Spermophaga, Pyrenestes; Uraeginthus, Granatina; Euschistospiza, Hypargos; Clytospiza, Pytilia, Lagonosticta. Cluster (c), mainly African: Amadina, Ortygospiza, Amandava. Cluster (d), 12 genera of Australasian grassfinches. Cluster (e), Oriental, Australasian and African: Odontospiza and Spermestes (Africa) with Heteromunia pectoralis (Australia); Euodice (Africa, India); Lonchura (Orient, Australasia) and Padda (Orient); and Lemuresthes nana (Madagascar). Sequences within each cluster can be seen as roughly evolutionary, from ‘ancestral’ to derived forms. We treat African genera in this sequence, except that we place Paludipasser after Ortygospiza. From the Asiatic Amandava we separate the single African species into Sporaeginthus. 137 species in 27 genera recognized by Goodwin (1982) and 140 in 27 genera by Sibley and Monroe (1990). We recognize 76 species in 25 genera in Africa; 21 genera endemic, 1 near-endemic (Estrilda), 1 shared with S Asia (Euodice) and 2 introduced from Asia (Amandava, Padda).

Genus Nigrita Strickland

Bill short and slender, with well-rounded culmen, usually somewhat flattened and expanded around nostrils; wings more pointed than rounded; tail black, fan-shaped, average length; plumage black and grey in 2 spp., grey and chestnut in 1, brown and white with black cap in 1; underparts uniform, no distinction between breast and belly. White wing spots in 1 species, N. canicapillus; these are single spots at tip of feather, as in Asian Amandava amandava, not paired as in twinspots. Palate and tongue marks estrildine, perhaps closest to Parmoptila, Cryptospiza and Pyrenestes (Goodwin 1982). See also Family diagnosis Occupies middle and upper levels in forest, often in canopy, not in low growth like most estrildines. Eats small insects and fruits. Bulky estrildine nest with side entrance. Endemic, 4 species. N. canicapillus and N. luteifrons are similarly plumaged but widely sympatric; unspotted W African race N. c. emiliae and white-backed Kungwe race N. c. candidus are divergent, perhaps incipient species (Hall and Moreau 1970). N. fusconotus, with more slender bill and more strongly graduated tail, was placed by Wolters (1966) in subgenus Percnopis.

Nigrita luteifrons Verreaux and Verreaux. Pale-fronted Negrofinch. Nigrette a ` front jaune

Plate 16

Nigrita luteifrons J. and E. Verreaux, 1851. Rev. Mag. Zool. (Paris), ser. 2, 3, p. 420; Gabon.

(Opp. p. 219)

Range and Status. Endemic resident. Sierra Leone (locality? G. D. Field in Gatter 1997); Liberia (once, Zlehtown, but probably rare and overlooked: Gatter 1997); Ivory Coast (Taı¨ Nat. Park and areas outside it: Gartshore et al. 1995); Ghana (Amedzofe, 0
520 N, 020
E); Togo, once at Misaho ¨he; Nigeria in forest zone, north to Ibadan, Ife and Okpo, uncommon, but sometimes outnumbers more abundant Grey-headed Negrofinch Nigrita canicapillus (Lagos: Gee and Heigham 1977); forest zone of S Cameroon from Ave´le´, Yaounde´ (seen 9 months of the year: Quantrill and Quantrill 1998) and Lobe´ke´ Faunal Res. north to Korup Nat. Park (few records), Limbe (not uncommon), Mt Cameroon (50–1300 m), Mt Kupe´ (uncommon, 750– 1000 m) and Bamenda, and in E to c. 5
300 N; Mbini, lowlands, and on Mt Alen at 500–950 m; Bioko; Gabon, widespread but not common; in NE even more infrequent than Grey-headed Negrofinch, in Lope´ Reserve confined to

borders of gallery forest; widespread in Congo (Nouabale´Ndoki and Odzala Nat. Parks, Le´fini, Mouilou Basin, Mayombe), outnumbering Grey-headed Negrofinch in Odzala (Dowsett and Dowsett-Lemaire 1997); Central African Republic around Bangui and in Lobaye Pre´fecture; not yet recorded in Dzanga Reserves in SW (Green and Carroll 1991) but almost certain to occur there since present across the borders in Cameroon and Congo; widespread but uncommon in Zaı¨re, from Ubangi, Uele and Semliki south to Mayombe, Kwango and Kasai (Luluabourg), and in E to Lubena west of L. Edward at 1000 m, Irangi Forest (Kizungu 2001) and Itombwe (to 1480 m); extreme W Uganda in Bwamba; uncommon Angola (Cabinda, and N Cuanza Norte at Canzele and Quiculungo). Formerly thought to be confined to Lower Guinea forests (e.g. Louette 1981) but recent discoveries in Upper

Nigrita luteifrons

253

Quite closely related with Viduidae, far more distantly with weavers Ploceidae and finches Fringillidae. Within Estrildidae, species relationships have been controversial, but recent molecular studies by M. Sorenson and R. B. Payne, (pers. comm.) suggest a phylogenetic arrangement as follows: Paludipasser locustella is only distantly related with all other estrildids, which fall into 5 clusters. Cluster (a), entirely African: Nigrita, Parmoptila; Nesocharis, Coccopygia; Mandingoa, Cryptospiza; Estrilda. Cluster (b), entirely African: Spermophaga, Pyrenestes; Uraeginthus, Granatina; Euschistospiza, Hypargos; Clytospiza, Pytilia, Lagonosticta. Cluster (c), mainly African: Amadina, Ortygospiza, Amandava. Cluster (d), 12 genera of Australasian grassfinches. Cluster (e), Oriental, Australasian and African: Odontospiza and Spermestes (Africa) with Heteromunia pectoralis (Australia); Euodice (Africa, India); Lonchura (Orient, Australasia) and Padda (Orient); and Lemuresthes nana (Madagascar). Sequences within each cluster can be seen as roughly evolutionary, from ‘ancestral’ to derived forms. We treat African genera in this sequence, except that we place Paludipasser after Ortygospiza. From the Asiatic Amandava we separate the single African species into Sporaeginthus. 137 species in 27 genera recognized by Goodwin (1982) and 140 in 27 genera by Sibley and Monroe (1990). We recognize 76 species in 25 genera in Africa; 21 genera endemic, 1 near-endemic (Estrilda), 1 shared with S Asia (Euodice) and 2 introduced from Asia (Amandava, Padda).

Genus Nigrita Strickland

Bill short and slender, with well-rounded culmen, usually somewhat flattened and expanded around nostrils; wings more pointed than rounded; tail black, fan-shaped, average length; plumage black and grey in 2 spp., grey and chestnut in 1, brown and white with black cap in 1; underparts uniform, no distinction between breast and belly. White wing spots in 1 species, N. canicapillus; these are single spots at tip of feather, as in Asian Amandava amandava, not paired as in twinspots. Palate and tongue marks estrildine, perhaps closest to Parmoptila, Cryptospiza and Pyrenestes (Goodwin 1982). See also Family diagnosis Occupies middle and upper levels in forest, often in canopy, not in low growth like most estrildines. Eats small insects and fruits. Bulky estrildine nest with side entrance. Endemic, 4 species. N. canicapillus and N. luteifrons are similarly plumaged but widely sympatric; unspotted W African race N. c. emiliae and white-backed Kungwe race N. c. candidus are divergent, perhaps incipient species (Hall and Moreau 1970). N. fusconotus, with more slender bill and more strongly graduated tail, was placed by Wolters (1966) in subgenus Percnopis.

Nigrita luteifrons Verreaux and Verreaux. Pale-fronted Negrofinch. Nigrette a ` front jaune

Plate 16

Nigrita luteifrons J. and E. Verreaux, 1851. Rev. Mag. Zool. (Paris), ser. 2, 3, p. 420; Gabon.

(Opp. p. 219)

Range and Status. Endemic resident. Sierra Leone (locality? G. D. Field in Gatter 1997); Liberia (once, Zlehtown, but probably rare and overlooked: Gatter 1997); Ivory Coast (Taı¨ Nat. Park and areas outside it: Gartshore et al. 1995); Ghana (Amedzofe, 0
520 N, 020
E); Togo, once at Misaho ¨he; Nigeria in forest zone, north to Ibadan, Ife and Okpo, uncommon, but sometimes outnumbers more abundant Grey-headed Negrofinch Nigrita canicapillus (Lagos: Gee and Heigham 1977); forest zone of S Cameroon from Ave´le´, Yaounde´ (seen 9 months of the year: Quantrill and Quantrill 1998) and Lobe´ke´ Faunal Res. north to Korup Nat. Park (few records), Limbe (not uncommon), Mt Cameroon (50–1300 m), Mt Kupe´ (uncommon, 750– 1000 m) and Bamenda, and in E to c. 5
300 N; Mbini, lowlands, and on Mt Alen at 500–950 m; Bioko; Gabon, widespread but not common; in NE even more infrequent than Grey-headed Negrofinch, in Lope´ Reserve confined to

borders of gallery forest; widespread in Congo (Nouabale´Ndoki and Odzala Nat. Parks, Le´fini, Mouilou Basin, Mayombe), outnumbering Grey-headed Negrofinch in Odzala (Dowsett and Dowsett-Lemaire 1997); Central African Republic around Bangui and in Lobaye Pre´fecture; not yet recorded in Dzanga Reserves in SW (Green and Carroll 1991) but almost certain to occur there since present across the borders in Cameroon and Congo; widespread but uncommon in Zaı¨re, from Ubangi, Uele and Semliki south to Mayombe, Kwango and Kasai (Luluabourg), and in E to Lubena west of L. Edward at 1000 m, Irangi Forest (Kizungu 2001) and Itombwe (to 1480 m); extreme W Uganda in Bwamba; uncommon Angola (Cabinda, and N Cuanza Norte at Canzele and Quiculungo). Formerly thought to be confined to Lower Guinea forests (e.g. Louette 1981) but recent discoveries in Upper

254

ESTRILDIDAE

Nigrita luteifrons

contrasting black face patch; juvenile uniform grey-brown. Often first detected by song.

Voice. Tape-recorded (104, B, CHA, ERA). Whistled song soft and musical, 6 notes descending the scale, last note long and drawn-out, ‘whee-hee-hee-hay-hay-hyooooo’. Call side to be a faint musical whistling ‘choo’ (Goodwin 1982).

?

?

Guinea suggest it has been overlooked, probably due to confusion with Grey-headed Negrofinch. Description. N. l. luteifrons Alexander: Nigeria to Angola and Zaı¨re. ADULT Y: forehead buffy white, merging into grey crown; sides of crown paler grey; rest of upperparts and upper sides of neck grey like centre of crown, but rump usually paler. Tail black, upperside slightly glossy. Lores and above eye to ear-coverts, cheeks and lower sides of neck black, sharply demarcated from sides of crown, and continuous with wholly black underparts. Flight feathers blackish grey; rest of upperwing dark grey, slightly glossy. Axillaries and underwing-coverts greyish white. Bill black; eyes reddish; legs pale brown or flesh. ADULT X: forehead, crown, upperparts, wings and tail like ad. Y. Lores to upper cheeks and above and behind eye black, forming small face patch; ear-coverts, lower cheeks and lower sides of neck grey; entire underparts ashy grey. Bill and legs as in ad. Y but eyes yellowish or creamy white. SIZE (10 YY, 10 XX): wing, Y 55–61 (587), X 57–59 (579); tail, Y 33–40 (378), X 34–39 (374); bill, Y 11–12 (112), X 11–125 (114). WEIGHT: 1 Y 13. IMMATURE: juvenile like ad. X, but more uniformly grey, lacking black face patch and white forehead. NESTLING: margin of gape black with 4 white tubercles, 1 just above angle of gape, 2 above and 1 below it; spots on palate and tongue like those of Estrilda (Bates 1911). N. l. alexanderi Ogilvie-Grant: Bioko. Forecrown as well as forehead buffy white; whitish sides to crown broader and better defined than in nominate race; grey upperparts generally paler. Slightly larger with heavier bill: Y, wing (n ¼ 8) 58–64 (613), bill (n ¼ 9) 115–13 (121). WEIGHT: 2 XX 12, 145. TAXONOMIC NOTE: racial status of Upper Guinea birds not yet determined.

Field Characters. Length 115 cm. Like Grey-headed Negrofinch but smaller, with short stubby bill. Y has pale buffy forehead, plain wing without white spots; X very different, with pale grey body, buffy forehead and

General Habits. Inhabits lowland primary, secondary and logged forest, gallery forest, thickets at forest edge and clearings; also cocoa and oil palm plantations, coffee forest, cultivated land with significant tree cover, trees in villages, gardens; in Gabon mainly in secondary forest, only entering primary forest in upper levels of canopy and emergents, and then rarely >300 m from forest edge; frequents habitats more open and lower than Grey-headed Negrofinch, old plantations and orchards, tall bushes in clearings, openings in forest made by storms, borders of streams in cultivated land, groves of trees near villages; occurs with Grey-headed Negrofinch at abrupt transitions between primary and secondary forest, at edges of clearings and beside roads and paths, preferring more open areas at heights of 2–18 m, while Grey-headed higher up at forest edge (Brosset and Erard 1986); associates with Greyheaded Negrofinch at fruiting trees. For list of dominant trees and plants in Irangi Forest, Zaı¨re, see under Greyheaded Negrofinch. Singly, in pairs or family groups, often with flocks of insectivores, gleaning large branches and working through foliage; visits flowers, often together with sunbirds; foraging actions tit-like; attends swarms of ants and termites, catching them in air like flycatcher.

Food. Mainly insects in Gabon (Brosset and Erard 1986), especially scale insects (Cameroon: Bates 1911); fruit, including Octhocosmus africanus and Ficus pseudomangifera, husks of oil palm fruits; some seeds.

Breeding Habits. Apparently territorial (Gabon). NEST: 2 described: (1) made of dry green moss and a few grass stems, lined with seeding grass panicles, diam. 130, width of side entrance 35, placed 35 m high in bushy top of garden hedge (Cameroon: Serle 1954); (2) oval, of fine grass, with entrance high on side, in bush c. 3 m above ground (Rand et al. 1959); another in Gabon (not described) in top of mango tree in village. EGGS: 4 (1 clutch); white. SIZE: c. 17  115. LAYING DATES: Ivory Coast (juv. Oct); Cameroon, May– June; Bioko, Oct; Gabon, Mar–Apr; Congo, May, July; Zaı¨re (Itombwe) Mar, June, Nov. DEVELOPMENT AND CARE OF YOUNG: young fed by both parents, which sang when they returned to nest, and young immediately started begging (Rand et al. 1959).

Key References

Clement, P. et al. (1993), Goodwin, D. (1982).

Nigrita canicapillus

255

Nigrita canicapillus (Strickland). Grey-headed Negrofinch. Nigrette a ` calotte grise.

Plate 16

Aethiops canicapillus Strickland, 1841. Proc. Zool. Soc. London, p. 30; Fernando Po.

(Opp. p. 219)

Range and Status. Endemic resident. Guinea (Macenta Pre´f.), common, and along border with Sierra Leone and Liberia (Mt Nimba), north in SE to Beyla; Sierra Leone (Botanic Reserve of Fourah Bay College, Mt Aureol (08
280 N, 13
140 W) and Gola Forest); Liberia, common from coast to N mts; Mali, once, south of Kangaba, Mar (Lamarche 1993); Ivory Coast, widespread north to Taı¨ Nat. Park, Mt Nimba, Odienne, Korhogo and Comoe´ Nat. Park; common in Yapo and Bossematie´ forests, also in Taı¨ Nat. Park, where seen on 30% of day trips (Gartshore et al. 1995); Ghana, common in forest zone from Kakum Forest, Cape Coast and Aburi Botanical Gardens (Moyer 1996) north to c. 7
N; Togo, common and widespread from coast north to Aledjo; Benin, frequent in Noyau Central; Nigeria, common throughout forest zone, north to Ibadan, Ife, Akure, Nsukka, Enugu and Oban Hills; S Cameroon lowlands, including Lobe´ke´ Faunal Res. (Dowsett-Lemaire and Dowsett 2000), north to c. 6
N, including Korup Nat. Park (common and widespread); common at Yaounde´, where recorded every month, and seen on 55 out of 70 trips (Quantrill and Quantrill 1998); also at lower levels in highlands: Rumpi Hills (1000–1250 m), Mt Cameroon (500–1500 m), Mt Kupe (350–1600 m), Mt Nlonako (110– 1250 m), north to Ribau; frequent on Mt Kupe´, most numerous in cultivated areas (Bowden 2001). Bioko, up to 2000 m; widespread in Mbini, on Mont Alen from 325 to 1200 m (Dowsett-Lemaire and Dowsett 1999); widespread in Gabon; common and widespread in Congo (Nouabale´Ndoki and Odzala Nat. Parks, Le´fini Reserve, Kouilou basin, Mayombe); SW Central African Republic from Dzanga-Ndoki Nat. Park and Dzanga-Sangha Rainforest Reserve to Bangui area (Bangui, Ndele, Botambi, Ndoko), and in SE around Baroua; Zaı¨re, throughout forest zone, from Mayombe to Semliki and Kivu (including Idjwi I.), south to Kwango, Kasai and Katanga (Haut-Lomami: Kayembe Makulu); common around edges of forest, less numerous in forest interior; infrequent in N Ubangi (Dejaifve 1994), very common in Itombwe; up to 1500 m in Ruwenzoris and Mugunga (Albert Nat. Park), 1600 m in mts east of Rutshuru, 1850 m in mts west of L. Edward (Bitakongo), 2000 m in Itombwe; at relatively low levels (700–950 m) in Irangi Forest near Kahuzi-Biega Nat. Park (Kizungu 2001). N Angola south in W to Cuanza Sul (Gabela), and in N Lunda Norte (Dundo, Camaiala R.); Rwanda (Nyungwe Forest, mainly below 2100 m, locally to 2400 m); Sudan (Bengengai, Imatong Mts, Lotti and Talanga forests, to 2400 m, fairly common); common and widespread in forests of W and S Uganda, mainly below 1500 m, from Impenetrable (Bwindi) (1200–2400 m), Malabigambo and Namalala north through Kalinzu, Kasyoha-Kitomi and Kibale to Bwamba, Itwara and Budongo, also to Masindi, Acholi and Sezibwa R. and Mt Elgon; Kenya, fairly common in highland forest, 1550– 3000 m, from Mt Elgon, Kapenguria, Cheranganis, Aberdares, Mt Kenya and Nyambenis to Kakamega, Nandi, Trans-Mara, Sotik, Kilgoris, Lolgorien, Mau and Nairobi,

Nigrita canicapillus

Mali: see Introduction.

also in S Kerio Valley in forest remnant above 2230 m (Wilson and Wilson 1994) and in Nguruman Hills; Tanzania in NW (Minziro Forest, Bukoba), W (KungweMahari Mt) and NE in Crater Highlands, Mbulu, Arusha Nat. Park and Mt Kilimanjaro (1900–2500 m). Density, 5–9 pairs per km2 in open secondary habitats, 3–5 in forest (Liberia); 5–6 pairs per km2 (Gabon). Description. N. c. canicapillus (Strickland): Bioko; S Nigeria to Gabon and the Lower and Middle Congo, intergrading with schistaceus in NE Zaı¨re from about Lisala to Semliki. ADULT Y: forehead and forecrown to lores, around eye, cheeks, ear-coverts, lower sides of neck and entire underparts black; top of crown to mantle, back and scapulars grey, sharply demarcated from black areas by white line; outermost scapulars broadly tipped white. Rump whitish, uppertail-coverts dark grey, tail black. Flight feathers blackish brown; primary coverts and alula blackish; tertials, greater coverts, median and lesser coverts blackish tipped with white spot. Axillaries and underwing-coverts white; inner borders of undersides of flight feathers greyish white. Bill black; eyes red; legs dark brown or blackish. Sexes alike. SIZE (10 YY, 10 XX, Cameroon): wing, Y 67–72 (701), X 68–73 (698); tail, Y 45– 51 (489), X 45–51 (482); bill, Y 14–155 (146), X 135–15 (144); tarsus, Y 165–18 (171), X 155–175 (166). Birds of Bioko slightly larger: (6 YY, 3 XX) wing, Y 71–73 (72), X 71–72 (717); bill, Y 15–16 (153), X 15–155 (151). WEIGHT: Cameroon, Y (n ¼ 4) 17–19 (182), X (n ¼ 4) 17–21 (192), 1 juv. Y 16, also 5 YY and 1 X 15–21 (176); Bioko (9 YY, 2 XX) 17–225 (211); IMMATURE: juvenile differs from ad. in having upperparts dark brownish grey, paler grey on rump; underparts dark brownish grey, paler on belly; wings browner, spots at tips of tertials, greater coverts and median coverts smaller, tinged buff. NESTLING: natal down greyish white; iris light grey; gape with 2 white tubercles at base of each mandible, palate with 5 black

256

ESTRILDIDAE

spots, black crescent on lower mandible (C. H. Fry, pers. comm.; fig. p. 252). N. c. emiliae Sharpe: Guinea and Sierra Leone to Togo. Differs from nominate race in having rump grey, border of hindcrown and nape pale grey rather than whitish; wing spots small, greyish white, confined to median and lesser coverts. Smaller: (10 YY) wing 62–68 (645), bill 12–135 (129). WEIGHT: Liberia, Y (n ¼ 7) 122–178 (152), X (n ¼ 7) 14–167 (154). N. c. schistaceus Sharpe: S Sudan to Uganda, Kenya west of Rift and NW Tanzania (Bukoba); intergrades with nominate race in Ruwenzoris and Kivu. Like nominate race, with whitish rump, but wing spots confined to median and lesser coverts; slightly darker grey above. WEIGHT: Uganda, Y (n ¼ 4) 17–23, X (n ¼ 6) 16–195, unsexed (n ¼ 29) 16–21 (avs?); Kenya, Y (n ¼ 3) 155–18 (165), 2 XX 165, 180, unsexed (n ¼ 3) 17–205 (186). N. c. diabolicus (Reichenow and Neumann): central Kenya east of Rift and N Tanzania (Crater Highlands to Kilimanjaro). Like schistaceus but slightly darker above, and rump grey like back; greater coverts and tertials (but not outer scapulars) tipped white. N. c. candidus Moreau: Kungwe Mt, Tanzania. Top of crown to upper mantle entirely whitish; rump whitish; greater coverts and tertials tipped white. N. c. angolensis Bannerman: NW Angola to S Zaı¨re (east to Kasai and Manyema). Like nominate race, but crown to back darker grey; rump greyish white, usually with darker grey barring. Slightly larger; wing, Y (n ¼ 3) 71–73 (723), X (n ¼ 3) 70–73 (717). TAXONOMIC NOTE: N. c. candidus and N. c. emiliae have been considered incipient species or megasubspecies.

Field Characters. Length 125–15 cm. A dark forest bird, uniform black below; grey crown and back separated from black forehead and face by white line; white spots on wingcoverts, white rump conspicuous in flight; eye red (turning yellow after death). Juvenile uniform dark grey with pale tips to wing-coverts and tertials, hint of a pale line from crown down sides of neck, eye pale grey. Race candidus (Kungwe-Mahari Mt) much whiter on head and rump; race emiliae (Guinea to Togo) lacks white line between crown and face. Distinguished from smaller Pale-fronted Negrofinch N. luteifrons by broad black forehead, white spots on wings and pale rump. Hard to see in forest where it remains hidden among canopy foliage; song often the best clue to its presence. Voice. Tape-recorded (104, B, C, F, ERA, HOR, KEI, LEM, PAY). Territorial song, usually given from top of tall tree, sweet, ringing, musical drawn-out notes, last ones lower; middle note may be highest, ‘hooo-heee-hooo’; notes may be 2-syllable, ‘huyoo-heeyee-hoop’ or 3-syllable, ‘heeyeeyee-heeyeeyee-hyoo-hyoo’; aggressive song, heard only east of Benin, often given near to the ground (Chappuis 2000), ascending whistles that start much lower than territorial song, ‘haw . . . hoo . . . her . . . hay . . . hee’, sometimes with introductory bubble and watch-winding call, ‘bubububub . . . drrrrrr . . . haw . . . her . . . hee’. Variable, perhaps regionally; in Nyungwe (Rwanda), a double whistle given 3 times, ‘fifu, fifu, fifu’, lasting c. 15 s (Dowsett-Lemaire 1990). Call said to be a plaintive ‘tsway’, metallic, like rusty wire spring (Bannerman 1949). General Habits. Inhabits primary and secondary forest, clearings, edges and gallery forest, and various habitats outside true forest: farmbush, cocoa and coffee plantations,

forest plantations, logged forest (Gola Forest, Sierra Leone, where more common outside forest: Allport et al. 1989); all treed habitats in Liberia, up to 1600 m on Mt Nimba; oil palm and cocoa plantations (Ghana); deserted farmland, wooded thickets and gardens (Nigeria); farms with or without tree cover, gardens (Cameroon); plantations, moss forest and montane heathland (Bioko); old orchards, old plantations of oil palm Elaeis, stream borders, clumps of bushes in savanna (Gabon); thickets, gardens, roadside scrub (Congo); old fields with tall isolated trees (Itombwe); coffee forest (Angola). Primary forest at Irangi, Zaire, characterized by canopy trees Gilbertiodendron dewevrei, Julbernardia sereti, Cynometra alexandri and Piptadeniastrum africanum, with Puelia ciliata and Afromomum spp. dominant in the herb layer, and shrub stratum characterized by Sapium ellipticum, Scaphopetalum thonneri and Thomandersia laurifolia; secondary forest dominated by trees Uapaca guineensis and Musanga cecropioides, by shrubs and lianas Macaranga spinosa, Albizia gummifera, Harungana madagascariensis, Alchornea cordifolia and A. floribunda, with understorey plants including Costus afer, Sporobolus sp., Panicum sp., and Setaria sp. (Kizungu 2001). Occurs at all levels, but in primary forest most often in canopy, from 25 m up to tops of tallest emergents; in clearings descends below 20 m to forage in liana tangles, and sometimes occurs in undergrowth with band of insectivores (Gabon); in Rwanda forages in thickets in small openings in forest, and around liana columns up to mid level; in Liberia keeps higher than Chestnut-breasted Negrofinch N. bicolor, although in Angola forages in oil palms in company with it and White-breasted Negrofinch N. fusconotus, sometimes on or near ground (Dean 2000). Pairs or family parties in mixed species flocks, especially with sunbirds; joins upper level guilds (Sierra Leone); searches for insects through clumps of foliage and along branches, including upper branches of dead trees; hops from branch to branch; gleans thin twigs 2–4 singing YY per km2, both in mature and old secondary forest; in Gabon, varies in abundance from place to place and year to year; averages 3–4 pairs per km2 in secondary forest regularly altered by

Nigrita fusconotus

?

Mali: see Introduction.

man, but in some years density may double or triple (Makokou: Brosset and Erard 1986). Description. N. f. fusconotus Fraser: Bioko and SE Nigeria to Angola, Zaı¨re and Kenya. ADULT Y: top of head to lores, below eye, ear-coverts, upper sides of neck and hindneck glossy violetblack. Mantle, scapulars, back and rump earth brown; uppertailcoverts glossy violet-black. Tail black, upperside glossy, tinged violet. Chin, cheeks and lower side of neck white, becoming offwhite on rest of underparts, flanks and sides of breast suffused with grey. Flight feathers, outer 2 tertials and primary coverts blackish brown; greater coverts, inner median coverts and innermost tertial paler grey-brown with slight oily green gloss; lesser coverts, outer median coverts and alula glossy violet-black. Axillaries and underwing-coverts whitish. Bill black; eyes brown or hazel to dark red; legs dark horn or dark slate-grey. Sexes alike. SIZE (10 YY, 8 XX): wing, Y 51–54 (526), X 47–53 (51): tail, Y 41–46 (442), X 36–44 (408); bill, Y 11–125 (117), X 105–12 (113); tarsus, Y 13–15 (137), X 135–15 (141). WEIGHT:

260

ESTRILDIDAE

Cameroon, 1 X 8, Bioko, unsexed (n ¼ 5) 85–11 (99); Uganda, Y (n ¼ 4) 8–9, unsexed (n ¼ 9) 8–10; Kenya, 1 X 7, 1 unsexed 95. IMMATURE: juvenile similar to ad., but top of head sooty brown and chin and cheeks to breast greyer; uppertail-coverts blackish with little gloss, wing-coverts less glossy. NESTLING: covered with white down, gape blackish with light coloured fleshy tubercles; black spots on palate (Chapin 1954). N. f. uropygialis Sharpe: Guinea to Ghana and SW Nigeria. Differs from nominate race in having rump pale buff, contrasting with rest of upperparts. WEIGHT: Liberia, Y (n ¼ 3) 8– 10 (89), X (n ¼ 1) 108.

Field Characters. Length 10–115 cm. A mainly treetop negrofinch which could be mistaken for a warbler as it forages among leaves. White below, with back cap, rump and tail, pale brown back; W race uropygialis has pale rump. Juv. similar but black areas dark brown, underparts greyer. Distinctive song best clue to its presence. Voice. Tape-recorded (104, B, C, CHA, ERA, HOR, KEI, LEM). Song of birds from Ivory Coast and Ghana (race uropygialis), slow, sweet whistles, long and slurred, ‘weeehyooo-hyooo’, ‘wayee-wayee-hyoo-heeeyoo’, sometimes with brief introductory notes, ‘tipit-wayeee-wooo’. Birds of nominate race (Gabon, Rwanda, Uganda, Kenya) have very different song, a high-pitched descending trill which decelerates, each note becoming longer and more widely spaced toward the end, ‘tsrrrrrrrrrr-ee-tee-tee-tee-tay-taytay-too-too’; notes at end of song may be double, ‘heedoheedo-heedo’. The great difference between songs of the 2 races suggests that they may be 2 species (Chappuis 2000), but the song described in Gabon (Lope´: Christy and Clarke 1994), a series of rising whistles, ‘sui-suit-suit-suitsuit-suit-suit’, accelerating into a trill and then slowing down, may bridge the difference. Further study and more tapes are needed. General Habits. Inhabits lowland and intermediate primary and secondary forest, especially edges and clearings and along tracks, logged forest and gallery forest; in Rwanda (Nyungwe) prefers crowns of Albizia trees (Dowsett-Lemaire 1990); also moss forest and montane heath (Bioko), savanna woodland (Benin), farmbush and cocoa plantations (Sierra Leone); around Makokou (Gabon) mainly in man-made secondary habitats: parklike cultivated land with tall trees around villages, old

overgrown plantations, various stages of regenerating forest where the canopy has not grown back, oil palm plantations (Brosset and Erard 1986). For description of Irangi Forest vegetation see Grey-headed Negrofinch. Mainly in middle and upper levels, typically in canopy (>30 m in Liberia); in clearings descends to lower levels chasing insects (Gabon: Lope´), and occurs in small trees and undergrowth, sometimes feeding on ground. Pairs or family parties, but 10 together on berries in cocoa plantation (Liberia) and numbers concentrate around clusters of ripe oil palm nuts (Gabon). Joins mixed species flocks in upper levels; works through foliage like Apalis warbler, pokes at bases of leaf stems, hanging acrobatically like a tit; forages in flowers and buds in tree canopy (Angola); feeds on pulp of fallen oil palm fruit. Sings from regular song post on topmost branch of tall forest tree, especially dead branch. Food. Small insects, including caterpillars and Heteroptera; fruits, including Macaranga assas, Ficus pseudomangifera and ‘tiny orange berries’ (Chapin 1954); oil, fruit pulp and oily fibrous husks of oil palm nuts; small seeds; some nectar. Breeding Habits. Y proclaims territory by singing from top of tall tree. NEST: round, untidy domed structure with side-top entrance, built of a variety of materials: fine fibres of dry plantain leaf stalks; thin twigs, dry grass leaves, flower stems and bits of torn bark; rough mass of brownish materials, including fine shreds of bark, dry grass, dead leaves and a little moss, not woven at all; and 1 entirely of fine bark fibre from a vine, looking like nest of tree-rat Thamnomys; placed 6–10 m above ground at base of palm frond, or towards end of thin leafy branch of small tree. EGGS: 3–6; elongate, without gloss; pure white. SIZE: (n ¼ 4) 13–145  103–11 (137  107); (n ¼ 6) 14–145  11. LAYING DATES: Liberia (independent young Jan, Mar– Apr); Ghana, Feb; Nigeria (nest-building Apr, breeding condition Jan); Cameroon (breeding condition June); Bioko, Nov; Gabon (nest-building Feb); Angola (breeding condition Mar, Aug–Sept); Zaı¨re, Ituri June–Sept, Itombwe Jan, Nov; E Africa: Uganda, July (breeding condition Apr–May, Nov–Dec), Kenya, Sept, Region B Apr, July–Aug. Key References

Clement, P. et al. (1993), Goodwin, D. (1982).

Genus Parmoptila Cassin

Tiny, rainforest-undergrowth, warbler-like birds; speckled brown, 2 species sexually dimorphic (YY with scarlet forehead), third only slightly so. Bills moderately broad-based but in profile fine and warbler-like; area around nostrils depressed, ridge of culmen slightly curved, central part of cutting edge of upper mandible arched; end of tongue narrowly cleft to about one-third of tongue’s length, tip and edges brush-like, thickly set with filaments on sides (Moreau et al. 1969); wings rounded, P1 very small; rather weak legs and feet; nestlings with black crescent below tongue and

260

ESTRILDIDAE

Cameroon, 1 X 8, Bioko, unsexed (n ¼ 5) 85–11 (99); Uganda, Y (n ¼ 4) 8–9, unsexed (n ¼ 9) 8–10; Kenya, 1 X 7, 1 unsexed 95. IMMATURE: juvenile similar to ad., but top of head sooty brown and chin and cheeks to breast greyer; uppertail-coverts blackish with little gloss, wing-coverts less glossy. NESTLING: covered with white down, gape blackish with light coloured fleshy tubercles; black spots on palate (Chapin 1954). N. f. uropygialis Sharpe: Guinea to Ghana and SW Nigeria. Differs from nominate race in having rump pale buff, contrasting with rest of upperparts. WEIGHT: Liberia, Y (n ¼ 3) 8– 10 (89), X (n ¼ 1) 108.

Field Characters. Length 10–115 cm. A mainly treetop negrofinch which could be mistaken for a warbler as it forages among leaves. White below, with back cap, rump and tail, pale brown back; W race uropygialis has pale rump. Juv. similar but black areas dark brown, underparts greyer. Distinctive song best clue to its presence. Voice. Tape-recorded (104, B, C, CHA, ERA, HOR, KEI, LEM). Song of birds from Ivory Coast and Ghana (race uropygialis), slow, sweet whistles, long and slurred, ‘weeehyooo-hyooo’, ‘wayee-wayee-hyoo-heeeyoo’, sometimes with brief introductory notes, ‘tipit-wayeee-wooo’. Birds of nominate race (Gabon, Rwanda, Uganda, Kenya) have very different song, a high-pitched descending trill which decelerates, each note becoming longer and more widely spaced toward the end, ‘tsrrrrrrrrrr-ee-tee-tee-tee-tay-taytay-too-too’; notes at end of song may be double, ‘heedoheedo-heedo’. The great difference between songs of the 2 races suggests that they may be 2 species (Chappuis 2000), but the song described in Gabon (Lope´: Christy and Clarke 1994), a series of rising whistles, ‘sui-suit-suit-suitsuit-suit-suit’, accelerating into a trill and then slowing down, may bridge the difference. Further study and more tapes are needed. General Habits. Inhabits lowland and intermediate primary and secondary forest, especially edges and clearings and along tracks, logged forest and gallery forest; in Rwanda (Nyungwe) prefers crowns of Albizia trees (Dowsett-Lemaire 1990); also moss forest and montane heath (Bioko), savanna woodland (Benin), farmbush and cocoa plantations (Sierra Leone); around Makokou (Gabon) mainly in man-made secondary habitats: parklike cultivated land with tall trees around villages, old

overgrown plantations, various stages of regenerating forest where the canopy has not grown back, oil palm plantations (Brosset and Erard 1986). For description of Irangi Forest vegetation see Grey-headed Negrofinch. Mainly in middle and upper levels, typically in canopy (>30 m in Liberia); in clearings descends to lower levels chasing insects (Gabon: Lope´), and occurs in small trees and undergrowth, sometimes feeding on ground. Pairs or family parties, but 10 together on berries in cocoa plantation (Liberia) and numbers concentrate around clusters of ripe oil palm nuts (Gabon). Joins mixed species flocks in upper levels; works through foliage like Apalis warbler, pokes at bases of leaf stems, hanging acrobatically like a tit; forages in flowers and buds in tree canopy (Angola); feeds on pulp of fallen oil palm fruit. Sings from regular song post on topmost branch of tall forest tree, especially dead branch. Food. Small insects, including caterpillars and Heteroptera; fruits, including Macaranga assas, Ficus pseudomangifera and ‘tiny orange berries’ (Chapin 1954); oil, fruit pulp and oily fibrous husks of oil palm nuts; small seeds; some nectar. Breeding Habits. Y proclaims territory by singing from top of tall tree. NEST: round, untidy domed structure with side-top entrance, built of a variety of materials: fine fibres of dry plantain leaf stalks; thin twigs, dry grass leaves, flower stems and bits of torn bark; rough mass of brownish materials, including fine shreds of bark, dry grass, dead leaves and a little moss, not woven at all; and 1 entirely of fine bark fibre from a vine, looking like nest of tree-rat Thamnomys; placed 6–10 m above ground at base of palm frond, or towards end of thin leafy branch of small tree. EGGS: 3–6; elongate, without gloss; pure white. SIZE: (n ¼ 4) 13–145  103–11 (137  107); (n ¼ 6) 14–145  11. LAYING DATES: Liberia (independent young Jan, Mar– Apr); Ghana, Feb; Nigeria (nest-building Apr, breeding condition Jan); Cameroon (breeding condition June); Bioko, Nov; Gabon (nest-building Feb); Angola (breeding condition Mar, Aug–Sept); Zaı¨re, Ituri June–Sept, Itombwe Jan, Nov; E Africa: Uganda, July (breeding condition Apr–May, Nov–Dec), Kenya, Sept, Region B Apr, July–Aug. Key References

Clement, P. et al. (1993), Goodwin, D. (1982).

Genus Parmoptila Cassin

Tiny, rainforest-undergrowth, warbler-like birds; speckled brown, 2 species sexually dimorphic (YY with scarlet forehead), third only slightly so. Bills moderately broad-based but in profile fine and warbler-like; area around nostrils depressed, ridge of culmen slightly curved, central part of cutting edge of upper mandible arched; end of tongue narrowly cleft to about one-third of tongue’s length, tip and edges brush-like, thickly set with filaments on sides (Moreau et al. 1969); wings rounded, P1 very small; rather weak legs and feet; nestlings with black crescent below tongue and

Parmoptila woodhousei

261

black-spotted palate. Untidy dome nest. Pure white eggs. Eat small insects, mainly ants (although brush-tipped tongues in passerine birds are associated with pollen- and nectar-feeding). Endemic; a superspecies with 3 species; P. rubrifrons and P. jamesoni (similar YY) separated by P. woodhousei; woodhousei and jamesoni (similar XX) meet.

Parmoptila woodhousei superspecies

1 2

3

2,3 1 P. rubrifrons 2 P. woodhousei 3 P. jamesoni

2

Parmoptila woodhousei Cassin. Woodhouse’s Antpecker. Parmoptile a ` gorge rousse.

Plate 16

Parmoptila Woodhousei Cassin, 1859. Proc. Acad. Nat. Sci. Philadelphia, 11, p. 40; Camma River, Gabon.

(Opp. p. 219)

Forms a superspecies with P. rubrifrons and P. jamesoni. Range and Status. Endemic resident, Nigeria to SW Zaı¨re and N Angola. Nigeria, rare, records from Lagos (once, a breeding pair, Gee and Heigham 1977), Okomu Forest Res. (near Benin; once) and, close to Cameroon border, Calabar, Oban West, Ikom, Bashu and Boshi-Okwango Forest Reserves (6
120 –6
330 N). Cameroon, widespread but infrequent in forest zone, c. 20 localities from Ikom and Korup to Lobe´ke´; frequent to common in Korup Nat. Park; rare in Kumba area; uncommon on Mt Kupe´ at 700–1050 m. Central African Republic, between Nola and Mbaiki (Sangha/Lobaye Pre´fs) and in Lobaye Pre´f. (Carroll 1988); Botambi; one, Ouossi R. west of Baroua (5
200 N, 24
200 E). Mbini, up to 1200 m on Mt Alen. Gabon, uncommon, Camma R., Gamba area (5 localities: Sargeant 1993), widespread and quite common in Ivindo Basin and Res. de la Lope´. Congo, fairly widespread in Odzala Nat. Park, local in Dzanga-Ndoki and Nouabale´-Ndoki Nat. Parks (Ndoki, Mbe´li, Bomassa, Mombongo), uncommon in Mayombe (2 localities near Dimonika), frequent in Kouilou Basin. Zaı¨re, uncommon; known from Bikoro (L. Tumba) and lower Ubangi valley, Basongo and Lusanga. Angola,

Parmoptila woodhousei

Parmoptila woodhousei

261

black-spotted palate. Untidy dome nest. Pure white eggs. Eat small insects, mainly ants (although brush-tipped tongues in passerine birds are associated with pollen- and nectar-feeding). Endemic; a superspecies with 3 species; P. rubrifrons and P. jamesoni (similar YY) separated by P. woodhousei; woodhousei and jamesoni (similar XX) meet.

Parmoptila woodhousei superspecies

1 2

3

2,3 1 P. rubrifrons 2 P. woodhousei 3 P. jamesoni

2

Parmoptila woodhousei Cassin. Woodhouse’s Antpecker. Parmoptile a ` gorge rousse.

Plate 16

Parmoptila Woodhousei Cassin, 1859. Proc. Acad. Nat. Sci. Philadelphia, 11, p. 40; Camma River, Gabon.

(Opp. p. 219)

Forms a superspecies with P. rubrifrons and P. jamesoni. Range and Status. Endemic resident, Nigeria to SW Zaı¨re and N Angola. Nigeria, rare, records from Lagos (once, a breeding pair, Gee and Heigham 1977), Okomu Forest Res. (near Benin; once) and, close to Cameroon border, Calabar, Oban West, Ikom, Bashu and Boshi-Okwango Forest Reserves (6
120 –6
330 N). Cameroon, widespread but infrequent in forest zone, c. 20 localities from Ikom and Korup to Lobe´ke´; frequent to common in Korup Nat. Park; rare in Kumba area; uncommon on Mt Kupe´ at 700–1050 m. Central African Republic, between Nola and Mbaiki (Sangha/Lobaye Pre´fs) and in Lobaye Pre´f. (Carroll 1988); Botambi; one, Ouossi R. west of Baroua (5
200 N, 24
200 E). Mbini, up to 1200 m on Mt Alen. Gabon, uncommon, Camma R., Gamba area (5 localities: Sargeant 1993), widespread and quite common in Ivindo Basin and Res. de la Lope´. Congo, fairly widespread in Odzala Nat. Park, local in Dzanga-Ndoki and Nouabale´-Ndoki Nat. Parks (Ndoki, Mbe´li, Bomassa, Mombongo), uncommon in Mayombe (2 localities near Dimonika), frequent in Kouilou Basin. Zaı¨re, uncommon; known from Bikoro (L. Tumba) and lower Ubangi valley, Basongo and Lusanga. Angola,

Parmoptila woodhousei

262

ESTRILDIDAE

rare: Uı´ge (Uı´ge town), N Cuanza Norte (Golungo Alto, Quiculungo), N Lunda Norte (Dundo). Density of 4–5 pairs per km2 on M’Passa plateau and 7–8 pairs per km2 on islands in Ivindo R., Gabon. Description. P. w. woodhousei Cassin: SE Nigeria to SW Central African Republic, Congo and W Zaı¨re. ADULT Y: forehead to hindneck dark brown, tinged rufous, feathers of forehead tipped crimson, those of crown tipped cinnamon-buff and with fine pale shaft streaks; mantle to uppertail-coverts rich brown with faint buff feather-tips, mantle feathers with pale buff shaft streaks. Tail feathers dark brown. Lores, cheeks and ear-coverts rufous-brown, merging with top of head and with more strongly rufous chin and upper throat. Lower throat, sides of neck, and breast to undertailcoverts whitish, washed buffy brown, especially on breast, and densely spotted dark olive-grey; each feather with two or more pairs of dark lateral spots. Remiges dark brown, narrowly edged buffy brown; upperwing-coverts and tertials warm brown, alula and greater coverts fringed paler cinnamon-brown, median and lesser coverts with small cinnamon-buff tips. Axillaries and underwing-coverts white, tinged brownish. Bill black; eyes reddish brown; legs pale brown or whitish brown. ADULT X: like ad. Y but forehead like crown, feathers lacking crimson tips. SIZE: (10 YY, 10 XX): wing, Y 48–53 (505), X 49–53 (505); tail, Y 37–41 (386), X 35–41 (382); bill, Y 11–125 (119), X 11–125 (118); tarsus, Y 14–15 (146), X 14–155 (148). WEIGHT: 1 Y 95. IMMATURE: juvenile differs from ad. in being more uniform brown above, with few pale shaft streaks; pale brown below, mottled darker and barred dark brown on flanks and undertailcoverts. NESTLING: 4 small white lobes at each corner of mouth (A). P. w. ansorgei Hartert: NW Angola. Slightly paler than nominate race. WEIGHT: 1 Y 9, 1 X 95.

Field Characters. Length 11 cm. A warbler-like bird of forest undergrowth, where it appears dull and drab in the half-light except for the rufous-orange face and throat (Christy and Clarke 1994); underparts spotted, Y with touch of red on forehead. Juv. has uniform rich brown underparts, with a little barring on flanks and undertail-coverts. Can only be confused with Jameson’s Antpecker P. jamesoni, which it meets in central Zaı¨re; distinguished from Y by spotted underparts, lack of prominent red patch on forehead, from X by spotted (not barred) underparts and lack of spots on throat. Juv. very similar to juv. Redfronted Antpecker P. rubrifrons but lighter and browner above, less rufous below. Voice. Tape-recorded (104, B, CHA, LEM). Call (song?) a long, shrill, high-pitched ‘tseeep’ or down-slurred ‘tseeeoo’, probably the same as the ‘tsi-up’ of Christy and Clarke (1994). General Habits. Inhabits primary lowland forest and old secondary formations; often in thick Afromomum and parasol-tree understorey growth near large clearings, and particularly in places where thick curtains of lianas reach down to ground (Gabon, Brosset and Erard 1986); disturbed parts of lowland rainforest with dense low herb layer (Mt Kupe´, Cameroon); swamp forest, at all levels, and all forest types in Nouabale´-Ndoki Nat. Park, Congo; but in secondary forest rarely moves out of ground layer or lower strata, keeping between 3 and 12 m above ground;

gallery forest and coffee forest (Angola). Rather silent; gives ‘tsi-up’ call only uncommonly; Y sings near nest. Occurs singly or often in pairs, or in family parties. Regularly joins mixed-species foraging flocks of insectivores. Shy, wary, and usually difficult to observe; but often stays for long time in a place where can be easy to watch (Christy and Clarke 1994). Forages amongst leaves much like a warbler, by inspecting them methodically and moving on rapidly; searches forks and crevices between twigs and around flakes of dry bark, particularly of trees of family Marantaceae; inspects clusters of dry leaves; opens green leaves stuck to each other with caterpillar silk; plunders papier-mache´-like nests of Tetramorium ants stuck below leaves. In captivity eats up to 30 ants per min, and survives only as long as live ants provided and ambient temperature kept at 24
C (Mitsch 1973). Captive birds not at all sociable with their own or other estrildid spp., but not aggressive; roost solitarily on branches. Sedentary. Food. Mainly arboreal ants; also caterpillars, small orthopterans and fruits of Xylopia aethiopica. In captivity in Europe, preferred large ants Formica rufa to small Lasius spp., and ate aphids, small moths, beetles and spiders (but refused houseflies and mealworms: Mitsch 1973). Breeding Habits. Solitary nester. NEST: large, bulky, domed mass of leaves, grass, bits of bark and vegetation debris, decorated loosely outside with bits of green moss, lined with fine fibres (once from banana leaves). Entrance 5 cm wide, at one side, one with ‘a sort of portico covering’ (Bates 1909). 20 cm high and up to 30 cm wide. Placed on forked horizontal twig. One c. 45 m up in isolated tree in small clearing; one in a small Martretia tree 25 m above water in flooded forest; one inside lower part of large cluster of dead leaves 3 m up in leafy tree. Built by Y and X; Y appears to build most of inside and X much of outside of nest; Y gives material to X and sings nearby when she finishes incorporating it into nest fabric. EGGS: 3–4, usually 4. Pure white. SIZE: (n ¼ 3) 130– 145  100–105. LAYING DATES: Nigeria, Lagos, (X in breeding condition, Mar), Calabar, (dependent young, May); Gabon, all year (nest-building Jan, Apr; nestlings Mar; young recently out of nest Oct, Feb; active gonads July, Dec); Angola, (moderately enlarged gonads Apr).

Parmoptila rubrifrons DEVELOPMENT AND CARE OF YOUNG:

young fed by both

parents.

263

Key References Bates, G. L. (1909), Brosset, A. and Erard, C. (1986), Christy, P. and Clarke, W. (1994), Dowsett-Lemaire, F. and Dowsett, R. J. (1991), Mitsch, H. (1973).

Parmoptila rubrifrons (Sharpe and Ussher). Red-fronted Antpecker. Parmoptile a` front rouge.

Plate 16

Pholidornis rubrifrons Sharpe and Ussher, 1872. Ibis p. 182; Denkera, Ghana.

(Opp. p. 219)

Forms a superspecies with P. woodhousei and P. jamesoni.

Parmoptila rubrifrons

Range and Status. Endemic resident, western Africa. Mali, known only from 2 YY taken at Misse´ni, June. Sierra Leone, Gola Forest, 3 records. Liberia, locally not uncommon, in Lofa County, Yekepa, Ye´ale´, Nimba, Sapo Nat. Park, Glaro Forest, and many sites in Grebo and Zwedru Forests. Guinea, Ziama Massif, Macenta, 2 records. Ivory Coast, Taı¨ Nat. Park, uncommon (4 localities) and Yapo Forest (scarce, Balchin 1990). Ghana, rare, known from Kaizaraku (Denkera, south of Kumasi), forests in Ashanti, Atewa, Subri River and Kakum Forest Reserves. Description. ADULT Y: broad scarlet band across forehead and forecrown, down to eye; hindcrown to hindneck dark greyish olive with pale grey scaly feather-fringes; rest of upperparts dark greyish olive-brown with faintly marked paler grey feather fringes. Tail dark grey-brown. Sides of head and neck dark greyish olive, spotted buffish white, spots fine and dense on lores, cheeks and ear-coverts, larger and less dense on superciliary band below hindcrown and nape and on side of neck. Chin pale buffy, merging with deep rufous-chestnut of rest of underparts. Flight feathers dark grey-brown, edged paler brown; upperwing-coverts and tertials greyish olive-brown. Axillaries and underwingcoverts rufous. Bill blackish, eyes chestnut-brown or dark red; legs pale yellowish brown. ADULT X: forehead and crown to hindneck dark olive-brown, fringed pale cinnamon-buff; rest of upperparts, sides of head, upperwing-coverts and tertials dark olive-brown. Flight feathers and tail as in ad. Y. Entire underparts greyish white, spotted blackish brown, more heavily from chin to breast, each feather with at least 2 pairs of lateral spots, flank feathers and undertail-coverts with several complete blackish bars. Bare parts as in ad. Y. SIZE: (10 YY, 10 XX): wing, Y 48–53 (502), X 49–53 (503); tail, Y 36–44 (404), X 39–43 (403); bill, Y 11–125 (118), X 11–125 (119); tarsus, Y 14–155 (150), X 145–16 (151). WEIGHT: Liberia, Y (n ¼ 19) 8–105 (92), X (n ¼ 13) 8–105 (98), imm. Y (n ¼ 3) 73, 75, 93; Ivory Coast (Taı¨), unsexed (n ¼ 3) 10–11 (av. given as 10). IMMATURE: juvenile more uniform above than ad.; underparts pale buff, mottled dark brown. Immature Y like ad. X, but underparts unspotted rufous-buff.

Field Characters. Length 10–11 cm. Y told by bright red forehead, spotted face and rufous-chestnut face and underparts, X by dark olive-brown upperparts and face and heavily spotted underparts. Juv. similar but more uniform above, underparts buffy, mottled dark brown. Voice. Not tape-recorded. Only described vocalization ‘whseeet, rather vigorous for such a small bird’ (Demey and Fishpool 1994); seems very like that of Woodhouse’s Antpecker.

Mali: see Introduction.

General Habits. Occurs in mature and logged forests, usually not far from water or at edge of high-forest swamp (Gatter 1997). Keeps within 10 m (Liberia) or 5 m (Ivory Coast) of ground. Gleans leaves and thin twigs, often in company of other species. Feeds at single ants’ nest for up to a minute; never seen at flowers (Gatter 1997). Y uttering weak ‘pee-you’ call opened bill surprisingly wide (Balchin 1990). Food. Mainly red ants; other small insects including beetles; some small seeds. Breeding Habits. Solitary nester. NEST: huge untidy mass. 3 nests, Liberia, 17  25, 25  30 and 25  40 cm. Sited 25–35 m above swampy or inundated ground, in small understorey tree. One nest built by Y alone, taking c. 8 days. LAYING DATES: Liberia, Mt Nimba, (oviduct egg, Apr; enlarged gonads Sept and Jan–Apr), Zwedru (3 nests, Oct, Nov); Ivory Coast, (juvs Dec, Jan, June). Nothing further known. Key References

Gatter, W. (1997), Woodcock, M. (2003).

Parmoptila rubrifrons DEVELOPMENT AND CARE OF YOUNG:

young fed by both

parents.

263

Key References Bates, G. L. (1909), Brosset, A. and Erard, C. (1986), Christy, P. and Clarke, W. (1994), Dowsett-Lemaire, F. and Dowsett, R. J. (1991), Mitsch, H. (1973).

Parmoptila rubrifrons (Sharpe and Ussher). Red-fronted Antpecker. Parmoptile a` front rouge.

Plate 16

Pholidornis rubrifrons Sharpe and Ussher, 1872. Ibis p. 182; Denkera, Ghana.

(Opp. p. 219)

Forms a superspecies with P. woodhousei and P. jamesoni.

Parmoptila rubrifrons

Range and Status. Endemic resident, western Africa. Mali, known only from 2 YY taken at Misse´ni, June. Sierra Leone, Gola Forest, 3 records. Liberia, locally not uncommon, in Lofa County, Yekepa, Ye´ale´, Nimba, Sapo Nat. Park, Glaro Forest, and many sites in Grebo and Zwedru Forests. Guinea, Ziama Massif, Macenta, 2 records. Ivory Coast, Taı¨ Nat. Park, uncommon (4 localities) and Yapo Forest (scarce, Balchin 1990). Ghana, rare, known from Kaizaraku (Denkera, south of Kumasi), forests in Ashanti, Atewa, Subri River and Kakum Forest Reserves. Description. ADULT Y: broad scarlet band across forehead and forecrown, down to eye; hindcrown to hindneck dark greyish olive with pale grey scaly feather-fringes; rest of upperparts dark greyish olive-brown with faintly marked paler grey feather fringes. Tail dark grey-brown. Sides of head and neck dark greyish olive, spotted buffish white, spots fine and dense on lores, cheeks and ear-coverts, larger and less dense on superciliary band below hindcrown and nape and on side of neck. Chin pale buffy, merging with deep rufous-chestnut of rest of underparts. Flight feathers dark grey-brown, edged paler brown; upperwing-coverts and tertials greyish olive-brown. Axillaries and underwingcoverts rufous. Bill blackish, eyes chestnut-brown or dark red; legs pale yellowish brown. ADULT X: forehead and crown to hindneck dark olive-brown, fringed pale cinnamon-buff; rest of upperparts, sides of head, upperwing-coverts and tertials dark olive-brown. Flight feathers and tail as in ad. Y. Entire underparts greyish white, spotted blackish brown, more heavily from chin to breast, each feather with at least 2 pairs of lateral spots, flank feathers and undertail-coverts with several complete blackish bars. Bare parts as in ad. Y. SIZE: (10 YY, 10 XX): wing, Y 48–53 (502), X 49–53 (503); tail, Y 36–44 (404), X 39–43 (403); bill, Y 11–125 (118), X 11–125 (119); tarsus, Y 14–155 (150), X 145–16 (151). WEIGHT: Liberia, Y (n ¼ 19) 8–105 (92), X (n ¼ 13) 8–105 (98), imm. Y (n ¼ 3) 73, 75, 93; Ivory Coast (Taı¨), unsexed (n ¼ 3) 10–11 (av. given as 10). IMMATURE: juvenile more uniform above than ad.; underparts pale buff, mottled dark brown. Immature Y like ad. X, but underparts unspotted rufous-buff.

Field Characters. Length 10–11 cm. Y told by bright red forehead, spotted face and rufous-chestnut face and underparts, X by dark olive-brown upperparts and face and heavily spotted underparts. Juv. similar but more uniform above, underparts buffy, mottled dark brown. Voice. Not tape-recorded. Only described vocalization ‘whseeet, rather vigorous for such a small bird’ (Demey and Fishpool 1994); seems very like that of Woodhouse’s Antpecker.

Mali: see Introduction.

General Habits. Occurs in mature and logged forests, usually not far from water or at edge of high-forest swamp (Gatter 1997). Keeps within 10 m (Liberia) or 5 m (Ivory Coast) of ground. Gleans leaves and thin twigs, often in company of other species. Feeds at single ants’ nest for up to a minute; never seen at flowers (Gatter 1997). Y uttering weak ‘pee-you’ call opened bill surprisingly wide (Balchin 1990). Food. Mainly red ants; other small insects including beetles; some small seeds. Breeding Habits. Solitary nester. NEST: huge untidy mass. 3 nests, Liberia, 17  25, 25  30 and 25  40 cm. Sited 25–35 m above swampy or inundated ground, in small understorey tree. One nest built by Y alone, taking c. 8 days. LAYING DATES: Liberia, Mt Nimba, (oviduct egg, Apr; enlarged gonads Sept and Jan–Apr), Zwedru (3 nests, Oct, Nov); Ivory Coast, (juvs Dec, Jan, June). Nothing further known. Key References

Gatter, W. (1997), Woodcock, M. (2003).

264

ESTRILDIDAE

Plate 16

Parmoptila jamesoni (Shelley). Jameson’s Antpecker. Parmoptile de Jameson.

(Opp. p. 219)

Pholidornis jamesoni Shelley, 1890. Ibis p. 163, pl. 5, fig. 1; Yambuya, Aruwimi River, Congo (i.e. Zaı¨re). Forms a superspecies with P. rubrifrons and P. woodhousei.

Parmoptila jamesoni

Range and Status. Endemic resident. In Zaı¨re, frequent to common, known from 11 localities in Itombwe and at least 30 elsewhere as mapped, mainly in Ituri Forest; common in Itombwe, at 880–1600 m. Uganda, uncommon in forests in W: Budongo, Bwamba, Kibale, Kalinzu and BwindiImpenetrable. Tanzania, Minziro Forest (N. Baker and M. Woodcock, pers. comm.). Description. ADULT Y: forehead and forecrown down to eye scarlet; hindcrown to upper ear-coverts, upper side of neck, mantle, scapulars and uppertail-coverts olive-grey, with faint scaling due to lighter grey feather tips. Tail olive-grey. Broad stripe through lores dark olive-grey. Entire underparts, including chin, throat, cheeks, lower ear-coverts and lower side of neck, orange-rufous. Flight feathers dark grey-brown, edged pale brown; upperwing-coverts and tertials olive-grey. Axillaries and underwing-coverts pale rufous. Bill blackish; eyes dark red or brownish red; legs yellowish or yellowish buff. ADULT X: upperparts greyish olive-brown, feathers of forehead and forecrown with small rufous tips. Sides of forehead to above and just behind eye rufous, forming a broad superciliary stripe; below it a dusky grey stripe through lores and behind eye. Chin, throat, cheeks, lower ear-coverts and lower sides of neck rufous. Sides of breast olive-brown, spotted pale olive-buff. Rest of breast to flanks and undertail-coverts pale grey-buff with dark greyish brown barring, broader and bolder on breast and flanks, each feather with 2 or more complete bars; upper breast feathers with small rufous tips. Tertials and upperwing-coverts olive-brown. Tail, flight feathers and bare parts as ad. Y. SIZE: (1Y, 1X): wing, Y 53, X 55; tail, Y 36, X 37; bill, Y 115, X 115; tarsus, Y 165, X 17. WEIGHT: 1 Y 95. IMMATURE: juvenile has upperparts plain olive-brown; underparts buff with faint dusky olive barring, most pronounced from lower breast to flanks and undertail-coverts; cheeks, sides of neck and throat, and upper breast suffused olive-brown. NESTLING: newly fledged young had iris dark brown; skin of gape black with 3 small yellowish lobes at each side; interior of mouth pale yellow with 5 black spots on palate and black crescent beneath tongue (Chapin 1954).

Field Characters. Length 10–11 cm. Needs to be distinguished from Woodhouse’s Antpecker P. woodhousei where they meet in Zaı¨re; Y readily told by bright red forehead and rufous face and underparts; X like X P. woodhousei (rufous superciliary stripe, cheeks and throat) but upperparts greyer, sides of breast olive and remaining underparts buffish with dark barring. Juv. similar to X but plainer above, cheeks and throat olive, rest of underparts buffy, faintly barred. Voice. Not known. General Habits. Inhabits lower storey of moist primary and logged forest, usually quite near water or at border of forest swamps; forest plantations; gallery forest; scrub along forest edges. In pairs or family parties. Regularly in mixed-species foraging flocks. Forages silently, working along branches and examining green foliage and hanging

bunches of dead brown leaves; pokes head into flowers like white-eye Zosterops; feeds at single ants’ nest for up to 1 min at a time. Pair spent some time poking into piece of peeled-off bark, presumably feeding on ants, working so energetically that it attracted attention of a Black-billed Weaver Ploceus melanogaster which chased them away (Uganda: S. Keith, pers. comm.). Food. Insects, including small beetles, ants, larvae, pupae and tiny cocoons; small seeds. Of 9 stomachs, Zaı¨re, only ants (ads, larvae, pupae) in 5 and only various small insects in other 4.

Breeding Habits. NEST: not described. LAYING DATES: Zaı ¨re, Ituri, (breeding birds Feb, May, Sept), Itombwe, (breeds Oct–May; juvs Feb, May, June, Nov). Nothing further known.

Key References

Chapin, J. P. (1954), Woodcock, M. (2003).

Nesocharis shelleyi

265

Genus Nesocharis Alexander

Genus characterized by distinctive grey, black and olive-green plumage. Bill short, fine (N. shelleyi) or stout, with decurved culmen and sharp point, with suggestion of hook at end of upper mandible; wings very short and rounded; tail short and square (N. shelleyi) to medium length and graduated (N. capistrata). Lively and acrobatic when foraging; 2 largely insectivorous, 1 specializing in seeds of Compositae; 2 use abandoned nest of weaver (mainly) or sunbird, N. capistrata builds typical estrildine nest, ball shaped with side entrance. Endemic. 3 species, 2 (N. shelleyi and N. ansorgei) forming a superspecies. Placed by Delacour (1943) in Estrilda; Immelmann et al. (1975) put it closest to Coccopygia melanotis. Steiner (1960) placed it in his tribe Cryptospizae with Cryptospiza, Nigrita, Pyrenestes and Parmoptila; of these, Goodwin (1982) thought it closest to Cryptospiza; now thought to be closest to Coccopygia (see Family diagnosis).

Nesocharis shelleyi superspecies

1 2 1 N. shelleyi 2 N. ansorgei

Nesocharis shelleyi Alexander. Shelley’s Oliveback. Dos-vert a` te ˆte noire.

Plate 17

Nesocharis shelleyi Alexander, 1903. Bull. Br. Orn. Club 13, p. 48; Moka, Fernando Po.

(Opp. p. 266)

Forms a superspecies with N. ansorgei. Range and Status. Endemic resident. Widespread in mts of SE Nigeria (Gaskaka-Gumti Game Res., Mambilla Plateau, Leinde Fadali at 1650 m, Gotel Mts (Gangirwal at 2100 m, Chappal Waddi at 1900 m), Obudu Plateau); Cameroon north to Tchabal Mbabo, on Bamenda Plateau, Mt Manenguba at 1950–2000 m, Mt Oku at 2200–2400 m, Mt Kupe´ at 1550–2000 m, Banso Mts north of Kumbo, Mt Cameroon at 1200–2100 m; Bioko. Widespread and locally common in Cameroon highlands; easily overlooked, and abundance may be underestimated; probably not under any immediate threat

(Stuart 1986). Listed as having a globally restricted range (500 birds (Parker 1999). Description. E. m. melanotis (Temminck): SE Africa. ADULT Y: forehead to hindneck down to eye and upper side of neck grey; mantle, scapulars and upper back yellowish olive-green with faint darker barring; lower back, rump and uppertail-coverts orangered. Tail black. Lores sometimes grey, sometimes black. Cheeks and ear-coverts to chin and upper throat black; lower throat white, joining white partial collar on lower side of neck. Breast pale grey to buffy grey, darkening to olive-grey on flanks; centre of belly to undertail-coverts yellowish buff. Flight feathers greybrown, narrow outer edges of primaries and edges of secondaries olive-green; tertials, alula and upperwing-coverts olive-green, tertials and greater coverts tinged red. Underwing-coverts and axillaries buff. Bill black above, red below with pinkish base; eyes bright red; legs brownish black. ADULT X: differs from ad. Y in having chin and throat whitish; cheeks and ear-coverts light grey, paler than top of head; rarely any red wash on tertials and greater coverts. SIZE (10 YY, 8 XX): wing, Y 46–48 (472), X 45–48 (470); tail, Y 34–37 (355), X 33–36 (350); bill, Y 95–11 (103), X 95– 11 (102); tarsus, Y 13–145 (138), X 13–14 (137). WEIGHT: Y (n ¼ 12) 57–84 (75), X (n ¼ 8) 72–84 (80), unsexed (n ¼ 160) 50–88 (63). IMMATURE: juvenile similar to ad. X, but green upperparts duller and lack fine barring, rump and uppertail-coverts pale rusty orange; bill all blackish; eyes dark brown. NESTLING: palate unmarked; near corner of mouth, side of maxilla with single papilla, black with white border, and side of mandible with 2 white papillae or tubercles surrounded by black; dark band across tongue. E. m. bocagei (Shelley): Angola and Namibia. Differs from nominate race in having flanks olive-green, lower breast to undertail-coverts yellow, and underpart feathers distinctly barred; brighter green (less yellowish) above, but Y lacks red tinge on wings; barring on upperpart feathers stronger and present on tertials and upperwing-coverts. TAXONOMIC NOTE: treated as conspecific with C. quartinia by Hall and Moreau (1970), Goodwin (1982) and Dowsett and Dowsett-Lemaire (1993a), although it is not strictly the case that they are behaviourally and ecologically identical as stated by the last authors. Several breeding pairs of C. m. melanotis, C. m. bocagei and C. quartinia in captivity together are ‘without strife or stress . . . and . . . behave as if specifically distinct, showing sexual

Coccopygia melanotis

interest only in others of the same form’ (Goodwin 1982). It suggests not only that C. melanotis and C. quartinia are distinct species but also that melanotis and bocagei may be specifically distinct. We take a middle course and recognize neither 1 nor 3 but 2 allospecies.

Field Characters. Length 9–10 cm. A small waxbill of rank vegetation at forest edge, with short, square black tail, red rump and uppertail-coverts; distinguished by combination of grey head and breast, olive-green back and wings and buffy yellow belly, also by 2-tone bill (black upper and red lower mandible). Y has black face and throat and broad white collar below throat; X has white throat and halfcollar extending under grey ear-coverts. Immature like X but duller, throat grey-brown, rump rusty orange, bill all black. In Angolan race bocagei, black barring on upperparts and flanks is pronounced, belly bright yellow. X and juv. best distinguished by range from extremely similar Yellow-bellied Waxbill C. quartinia stuartirwini, see superspecies map. Voice. Tape-recorded (88, 99, 104, F, GREG, MOY, PAY, STJ). Contact call a thin ‘seee’ or down-slurred ‘seeeu’, given at irregular intervals; at high intensity an explosive ‘tswee!’ or repeated ‘tseek-tseek-tseek’ (Goodwin 1982); when several birds calling together, shorter ‘see’, also ‘tsidit’ and high ‘tsit’. Apparent song, 4-note ‘see-see-soo-see’, 3rd note lower, pure and sweet; another ‘song’ described as a single soft but penetrating ‘teeeeeeit!’ or ‘tuuuuueet!’ (Goodwin 1982). Alarm, sharp ‘teerrrr!’.

Coccopygia quartinia General Habits. Inhabits borders of montane evergreen and Drakensberg mistbelt forests, lowland and coastal forests, grassy clearings in moist woodland; bushy hillsides, gardens, farms, fallow land with dense cover, exotic plantations, thick riverine scrub; in fynbos, near wooded kloofs; brushy undergrowth and grassy tracks in coastal dune scrub. Appears to be loosely associated with alien grass Pennisetum setaceum on burnt, developed land in SW Cape Prov. (Martin and Pepler 2002). Occurs in pairs and flocks of up to c. 12. Forages in heads of grasses and on ground; active and agile. Quiet, unobtrusive, can be tame and confiding, especially in gardens around farmsteads and houses. Does not mix with other estrildids. Appears to be highly mobile though essentially resident; perhaps some altitudinal movements in KwaZulu-Natal Drakensberg (Harrison et al. 1997). Food. Seeds and small insects. Seeds of Rhynchelytrum repens, Digitaria stentiana and Panicum maximum; presumably eats seeds of Pennisetum setaceum (see above); in captivity eats seeds of P. laevifolium, Setaria sphacelata, Sporobolus fimbriatus, Chloris virgata, Digitaria sanguinalis, Sonchus, Taraxacum and Cichorium, and invertebrates including spiders, aphids, termites, mealworms and small moths (Brickell and Konigkramer 1997). Breeding Habits. Solitary nester; monogamous; territorial.

273

NEST: rather flimsy oval or pear-shaped ball made of broad-bladed dry grass, the lower-side entrance with short spout; thickly lined with grass inflorescences and feathers (of e.g. Speckled Pigeon Columba guinea). Inflorescences can project from entrance tube. 1 nest c. 150  100. Sited in fork up to 2 m above ground in bush, small tree including conifers and Euphorbia ingens, hedge or creeper; often in conspicuous place in gardens. EGGS: 3–9, usually 4–5 (18 clutches, av. 47); large clutches may be due to 2 XX laying in same nest (Brickell and Konigkramer 1997). White. SIZE: (n ¼ 34) 122–163  99–110 (137  104). LAYING DATES: Zimbabwe, (nest-building, July); South Africa: Transvaal, Nov–Feb; KwaZulu-Natal, Nov–Apr; Cape Prov., Oct–May (E Cape, Nov–Jan). INCUBATION: by Y and X. Period, in captivity: 12–13 days. DEVELOPMENT AND CARE OF YOUNG: fed by Y and X. Period: 19–22 days. Young out of nest follow X wherever she moves or settles, soliciting food with great persistence, hustling one another and pushing her from both sides (Vincent 1949). BREEDING SUCCESS/SURVIVAL: once parasitized by Pin-tailed Whydah Vidua macroura (Martin 1983).

Key References Brickell, N. (1985a), Clement, P. et al. (1993), Goodwin, D. (1982).

Coccopygia quartinia (Bonaparte). Yellow-bellied Waxbill. Astrild a ` ventre jaune.

Plate 18

E[strelda] quartinia Bonaparte, 1851? Consp. Avium, 1 (1850), p. 461; Abyssinia.

(Opp. p. 267)

Forms a superspecies with C. melanotis. Range and Status. Endemic resident, eastern Africa. Eritrea, frequent to common at 900–2450 m on E escarpment; commonest in high rainfall areas. Ethiopia, locally common to abundant in W and SE Highlands, Rift Valley and S Ethiopia, mainly between 900 and 3000 m; south to Gardulla, Burgi, Adola and Yabello. Sudan, common up to 2400 m in Imatong, Dongotona and Didinga Mts. E Zaı¨re, uncommon (Kivu Volcanoes) to frequent (W Rwenzori) and locally common (northwest of L. Edward), south to Itombwe (common at 1830–1910 m: Luiko, Minembwe, Nzombe), Mt Kabobo (frequent at 2420–2480 m) and Marungu Highlands (common, Pepa and Ketendwe areas). E Africa, at 900– 3000 m as mapped. Uganda, in W from Budongo, Bugoma and Mubende to Kigezi; in NE and E, Mts Lonyili, Morongole, Moroto and Elgon. Kenya, fairly common and widespread in W and central highlands; also on Mts Kulal, Nyiru and Marsabit, Ndoto and Mathews Ranges, and Nguruman, Chyulu and Taita Hills. Tanzania, in NW at Bukoba, in NE from Loliondo and Crater Highlands to Mt Hanang, Mts Kilimanjaro and Meru, Arusha Nat. Park, Pare, Usambara, Ukaguru and Uluguru ranges (single record near Dar-es-Salaam, perhaps an escapee), in S in Udzungwa, Mbeya, Kipengere, Njombe, Songea and

Coccopygia quartinia

Coccopygia quartinia General Habits. Inhabits borders of montane evergreen and Drakensberg mistbelt forests, lowland and coastal forests, grassy clearings in moist woodland; bushy hillsides, gardens, farms, fallow land with dense cover, exotic plantations, thick riverine scrub; in fynbos, near wooded kloofs; brushy undergrowth and grassy tracks in coastal dune scrub. Appears to be loosely associated with alien grass Pennisetum setaceum on burnt, developed land in SW Cape Prov. (Martin and Pepler 2002). Occurs in pairs and flocks of up to c. 12. Forages in heads of grasses and on ground; active and agile. Quiet, unobtrusive, can be tame and confiding, especially in gardens around farmsteads and houses. Does not mix with other estrildids. Appears to be highly mobile though essentially resident; perhaps some altitudinal movements in KwaZulu-Natal Drakensberg (Harrison et al. 1997). Food. Seeds and small insects. Seeds of Rhynchelytrum repens, Digitaria stentiana and Panicum maximum; presumably eats seeds of Pennisetum setaceum (see above); in captivity eats seeds of P. laevifolium, Setaria sphacelata, Sporobolus fimbriatus, Chloris virgata, Digitaria sanguinalis, Sonchus, Taraxacum and Cichorium, and invertebrates including spiders, aphids, termites, mealworms and small moths (Brickell and Konigkramer 1997). Breeding Habits. Solitary nester; monogamous; territorial.

273

NEST: rather flimsy oval or pear-shaped ball made of broad-bladed dry grass, the lower-side entrance with short spout; thickly lined with grass inflorescences and feathers (of e.g. Speckled Pigeon Columba guinea). Inflorescences can project from entrance tube. 1 nest c. 150  100. Sited in fork up to 2 m above ground in bush, small tree including conifers and Euphorbia ingens, hedge or creeper; often in conspicuous place in gardens. EGGS: 3–9, usually 4–5 (18 clutches, av. 47); large clutches may be due to 2 XX laying in same nest (Brickell and Konigkramer 1997). White. SIZE: (n ¼ 34) 122–163  99–110 (137  104). LAYING DATES: Zimbabwe, (nest-building, July); South Africa: Transvaal, Nov–Feb; KwaZulu-Natal, Nov–Apr; Cape Prov., Oct–May (E Cape, Nov–Jan). INCUBATION: by Y and X. Period, in captivity: 12–13 days. DEVELOPMENT AND CARE OF YOUNG: fed by Y and X. Period: 19–22 days. Young out of nest follow X wherever she moves or settles, soliciting food with great persistence, hustling one another and pushing her from both sides (Vincent 1949). BREEDING SUCCESS/SURVIVAL: once parasitized by Pin-tailed Whydah Vidua macroura (Martin 1983).

Key References Brickell, N. (1985a), Clement, P. et al. (1993), Goodwin, D. (1982).

Coccopygia quartinia (Bonaparte). Yellow-bellied Waxbill. Astrild a ` ventre jaune.

Plate 18

E[strelda] quartinia Bonaparte, 1851? Consp. Avium, 1 (1850), p. 461; Abyssinia.

(Opp. p. 267)

Forms a superspecies with C. melanotis. Range and Status. Endemic resident, eastern Africa. Eritrea, frequent to common at 900–2450 m on E escarpment; commonest in high rainfall areas. Ethiopia, locally common to abundant in W and SE Highlands, Rift Valley and S Ethiopia, mainly between 900 and 3000 m; south to Gardulla, Burgi, Adola and Yabello. Sudan, common up to 2400 m in Imatong, Dongotona and Didinga Mts. E Zaı¨re, uncommon (Kivu Volcanoes) to frequent (W Rwenzori) and locally common (northwest of L. Edward), south to Itombwe (common at 1830–1910 m: Luiko, Minembwe, Nzombe), Mt Kabobo (frequent at 2420–2480 m) and Marungu Highlands (common, Pepa and Ketendwe areas). E Africa, at 900– 3000 m as mapped. Uganda, in W from Budongo, Bugoma and Mubende to Kigezi; in NE and E, Mts Lonyili, Morongole, Moroto and Elgon. Kenya, fairly common and widespread in W and central highlands; also on Mts Kulal, Nyiru and Marsabit, Ndoto and Mathews Ranges, and Nguruman, Chyulu and Taita Hills. Tanzania, in NW at Bukoba, in NE from Loliondo and Crater Highlands to Mt Hanang, Mts Kilimanjaro and Meru, Arusha Nat. Park, Pare, Usambara, Ukaguru and Uluguru ranges (single record near Dar-es-Salaam, perhaps an escapee), in S in Udzungwa, Mbeya, Kipengere, Njombe, Songea and

Coccopygia quartinia

274

ESTRILDIDAE

Matengo Highlands (common in Matengo: Mtindi, Lipumba, Kindimba, Litembo, Ugano, Mahuka), and in W on Mahari Mt and Ufipa Plateau. Zambia, over 1700 m on Nyika Plateau; Mukutu and Mafinga Mts; Muchinga ˆ i, Escarpment, Danger Hill and Musense (Serenje). Malaw widespread above 1375 m; up to 2165 m on Mt Mulanje (where occurs down to 885 m) and up to 2440 m on Nyika Plateau; occurs in Dzalanyama Mts, Lilongwe. Zimbabwe, up to 2200 m in Nyanga Highlands; Chimanimani Mts, Chipinga Uplands around Mt Selinda (single record near Harare, perhaps an escapee). Mozambique, Manica Platform, Mts Furancungo, Gorongoza and Namuli (Ryan et al. 1999). Description. E. q. kilimensis (Sharpe) (includes ‘nyanzae’): SE Sudan, E Zaı¨re and Uganda to Kenya and central Tanzania. ADULT Y: forehead to hindneck and upper sides of neck rather dark grey; mantle, scapulars and upper back dark olive-green with fine dusky barring; lower back to uppertail-coverts orangered. Tail black. Lores dark grey; cheeks and ear-coverts grey, lighter than top of head. Chin and throat to lower sides of neck light grey; breast grey, merging with olive-green flanks; belly and undertail-coverts deep olive-yellow, often tinged orange. Flight feathers grey-brown, narrow outer edges of primaries and edges of secondaries dark olive-green; tertials, alula and upperwingcoverts dark olive-green. Underwing-coverts and axillaries olive-yellow. Bill black above, red below; eyes crimson or redbrown; legs grey-brown. ADULT X like ad. Y, but underparts average rather paler and lack orange tinge. SIZE (10 YY, 10 XX): wing, Y 45–48 (461), X 44–47 (456); tail, Y 34–37 (352), X 33– 37 (347); bill, Y 95–10 (98), X 9–10 (96); tarsus, Y 14–15 (146), X 135–15 (144). WEIGHT: SE Sudan, unsexed (n ¼ 10) 6–7 (65); Kenya, unsexed, (n ¼ 15) 6–8 (69). IMMATURE: juvenile like ad. X but lacks barring on upperparts; rump rusty orange; bill all-black. NESTLING: pink, with slight tufts of down on head and dorsum; gape with small tubercles; palate and tongue with pattern of small black spots (van Someren 1956). E. q. quartinia Bonaparte: Eritrea and Ethiopia. Grey cap paler than in kilimensis, upperparts lighter and greener; breast paler grey; yellow belly and undertail-coverts lighter and cleaner, with no orange tinge in Y. E. q. stuartirwini Clancey: S and E Tanzania to E Zimbabwe. Grey cap slightly paler than in kilimensis, upperparts greener, breast slightly paler grey. WEIGHT: S Tanzania, unsexed, (n ¼ 25) 55–75 (67); Zimbabwe, Y (n ¼ 2) 57, 62, X (n ¼ 2) 70, 72.

Field Characters. Length 9–10 cm. A small, short-tailed bird of highland undergrowth. Sexes similar, with grey head, olive-green back and wings, red rump and black tail; belly colour variable, bright yellow to pale yellow, buff or orange; bill red and black. Immature has black bill, orange rump. Fawn-breasted Waxbill Estrilda paludicola of damp grassland has all-red bill, rich brown back and wings. In Zimbabwe comes close to range of Swee Waxbill C. melanotis but they do not overlap. Voice. Tape-recorded (99, PAY). Calls, sibilant ‘pssssl’, ‘ssss’ and ‘psseet’, also a lower ‘tyoo’. Song, 5-syllable ‘pew, sit-tu, silly’ and variations; quiet, inserted among calls almost like an aside. Alarm a sharp ‘teerrrr’, like Swee Waxbill.

General Habits. Inhabits wet and sometimes dry upland grass, low bushes around lower borders of mountain forest, moist thickets, open savannas, fallow fields and old farmland; large gardens; grassy forest clearings (Itombwe Highlands, Zaı¨re) and bracken-briar (Marungu Highlands, ˆ i, Zimbabwe, Mozambique) and Zaı¨re; Zambia, Malaw coarse mountain vegetation, edges of evergreen forest, and regenerating growth in felled pine plantations (Zimbabwe). Occurs in pairs or flocks of 8–20, usually 10–15 birds; often with other estrildids. Forages on ground but mainly by perching on grass heads, pecking seeds out; in Kenya concentrates in areas thickly overgrown with sticky-grass Setaria (van Someren 1956). When flushed from ground, flock flies up into nearest bush or small tree, then drops down to ground again when danger passes. 3 birds once seen at old nest of weaver Ploceus baglafecht reichenowi, ‘which they seemed to have relined with grass tops and probably used as a roost’ (Chapin 1954). Resident, but in Zimbabwe reduction in reporting rate in upland habitats in May–Oct and occurrences down to 350–400 m in Haroni-Rusitu area in Sept suggest movement to lower altitudes (Harrison et al. 1997). Food. Very small grass seeds, including those of Setaria. Breeding Habits. Solitary nester. NEST: flimsy but quite bulky long-oval ball made of rather coarse, interlaced grass stems, with entrance at side just above supporting branch; quite thickly lined with white plant down, fine grass heads and sometimes with a few feathers. Entrance has short spout or porch, made of flowering grass heads. Placed in fork within thick foliage near top of shrub or small tree, 12–5 m above ground; often in clump of mistletoe in tall tree; sometimes near ˆ i, nests only in paper-wasps’ nest or in thorn tree; in Malaw secondary growth outside forest (Dowsett-Lemaire 1983). Built by both sexes, taking 7–10 days. Double-brooded (in long rains in Kenya). ˆ i) or nearer to 5 (Kenya). Pure EGGS: 3–6, av. 4 (Malaw white; very thin shelled. SIZE: (n ¼ 2) 135  110, 14  10. LAYING DATES: Eritrea, (building Sept, nestlings Nov); Ethiopia, Mar, June–July, Aug–Nov; Sudan, Oct–Nov, E Zaı¨re, Mt Avu, (Aug), Itombwe, July; Uganda, Bwamba, Apr, July–Aug; E Africa, Region D, Dec–Jan, May–July ˆ i, Feb–May. (half of 14 clutches in June); Malaw INCUBATION: period 12–13 days. DEVELOPMENT AND CARE OF YOUNG: young fed halfdigested mass of tiny grass seeds, by regurgitation in small portions to each chick in turn; feeding whole brood takes up to 5 mins; feeds up to 1 h apart; both parents often come to nest together, 1 remaining outside until the other has finished. Period: 14–16 days. Parents and young sometimes return to nest for a few nights to sleep. Begging call of fledglings wheezy and not unmusical, ‘ss-ss-see’, loud for size of bird (Sclater and Moreau 1932–1933). Young out of nest stay with parents until next nesting, when they join up with young from other nests (van Someren 1956). Key Reference van Someren, V. G. L. (1956).

Mandingoa nitidula

275

Genus Mandingoa Hartert

Bill sharply pointed, culmen almost straight, nostrils hidden by feathers; wings short and pointed, outer primary minute; tail rounded, shorter than outstretched legs. Plumage mostly green except for black belly feathers with paired white spots at tip, which have placed it among the twinspots. Endemic, 1 species. Often placed in Hypargos, but tail shorter and wing more pointed, and Wolters (in Hall and Moreau 1970) has shown that they differ in various structural characters; green plumage not shared by any other twinspots. Red face patch suggestive of Red-faced Crimsonwing Cryptospiza reichenovii, but Cryptospiza is completely unspotted; general colour and pattern suggests a possible relationship with Pytilia (Goodwin 1982). Now thought to be closest to Cryptospiza, in estrildid Cluster (a) (R. B. Payne, pers. comm.; see Family diagnosis).

Mandingoa nitidula (Hartlaub). Green Twinspot. Se ´ne´gali vert.

Plate 20

Estrelda nitidula Hartlaub, 1865. In Gurney, Ibis, 1865, p. 269; Natal.

(Opp. p. 283)

Range and Status. Endemic resident, widespread in Afrotropics; scarce to locally abundant. Guinea, in SW (Kounounkan) and along borders of Sierra Leone and Liberia, rare Macenta; W and SE Sierra Leone, including Gola Forest; Liberia, locally common in N highlands, rare on coast and at Zwedru; Ivory Coast, frequent in SW in wettest parts of forest zone, including Nimba, Taı¨ and Danane, also present in E in Bossematie´ Forest and in Comoe´ Nat. Park; Ghana in forest zone north to Bia Nat. Park and Mampong; Togo, 1 record (Misaho ¨he), presumed rare resident (Cheke and Walsh 1996); 3 records Nigeria, Ibadan, Oban Hills, and recently at Boje-Enyi, Cross River State (Bull. Afr. Bird Club 2, 126). Forest zone of Cameroon north to Tchabal Mbabo, west to Korup Nat. Park, east to Lobe´ke´, but scarce or absent in extreme S; known from Yaounde´ but not seen by Quantrill and Quantrill (1998), and not recorded for Mbini or adjacent N Gabon; Bioko; Central African Republic in SW and SE (Baroua), north to Sangba; Gabon, locally abundant around Makokou, and listed for Doume´, Ogooue district (Bannerman 1949), doubtless more widespread; Congo, known from Nouabale´-Ndoki Nat. Park (common), Odzala Nat. Park (rare) and in S at Sibiti and Dimonika; Zaı¨re, widespread in N from N Ubangui (frequent: Dejaifve 1994), Uele and Ituri, and in E south to Itombwe (very common); known from widely scattered localities elsewhere, lower Congo in W, Upemba Nat. Park and Kasaji in S, doubtless more widespread than shown on map; uncommon in Angola from Uı´ge and Luhanda (Malanje) south to Quitondo and Gabela in Cuanza Sul; E Rwanda (valleys of Akagera, Kibaya, Kagogo and Nyaruvomo rivers, around lakes Mhanga and Mugesera, and near Kigali: vande Weghe 1981b); not reported from Nyungwe Forest; Burundi (Musigati, Kigwena, Bururi Forest); common extreme S Sudan (Bengengai Forest, Imatong and Dongotona Mts, up to 2400 m); Ethiopia in W Highlands (upper reaches of Omo R.), Rift Valley, including Nechisar Nat. Park (frequent), and recently at Wendo Genet (Farnsworth et al. 2000), and in S at Alghe and Arero, generally uncommon; widespread W and S Uganda from Budongo,

Mandingoa nitidula

?

Bugoma, Bwamba (abundant), Maramagambo, KasyohaKitomi, Impenetrable (Bwindi) and Sango Bay forests east to Entebbe, Kifu and Mabira, also in N (Kidepo Valley Nat. Park) and extreme NW (Otze Forest White Rhino Sanctuary: Forbes-Watson 1994); Kenya along coast from Tana R. to Vanga, inland on Shimba, Taita and Ngulia Hills, central highlands (N Nairobi, Kiambu, Aberdares, Meru Forest, and Kikuyu Escarpment Forest at 2350 m (Taylor and Taylor 1988); status uncertain: periodic reports from Mt Elgon, Kerio Valley, Kakamega, North Nandi Forest, Mara Game Res. and Lolgorien may in part represent wandering or displaced birds (Zimmerman et al. 1996); formerly on Mt Marsabit. Tanzania, widespread in coastal forests, inland to Eastern Arc mts, North Pares

Mandingoa nitidula

275

Genus Mandingoa Hartert

Bill sharply pointed, culmen almost straight, nostrils hidden by feathers; wings short and pointed, outer primary minute; tail rounded, shorter than outstretched legs. Plumage mostly green except for black belly feathers with paired white spots at tip, which have placed it among the twinspots. Endemic, 1 species. Often placed in Hypargos, but tail shorter and wing more pointed, and Wolters (in Hall and Moreau 1970) has shown that they differ in various structural characters; green plumage not shared by any other twinspots. Red face patch suggestive of Red-faced Crimsonwing Cryptospiza reichenovii, but Cryptospiza is completely unspotted; general colour and pattern suggests a possible relationship with Pytilia (Goodwin 1982). Now thought to be closest to Cryptospiza, in estrildid Cluster (a) (R. B. Payne, pers. comm.; see Family diagnosis).

Mandingoa nitidula (Hartlaub). Green Twinspot. Se ´ne´gali vert.

Plate 20

Estrelda nitidula Hartlaub, 1865. In Gurney, Ibis, 1865, p. 269; Natal.

(Opp. p. 283)

Range and Status. Endemic resident, widespread in Afrotropics; scarce to locally abundant. Guinea, in SW (Kounounkan) and along borders of Sierra Leone and Liberia, rare Macenta; W and SE Sierra Leone, including Gola Forest; Liberia, locally common in N highlands, rare on coast and at Zwedru; Ivory Coast, frequent in SW in wettest parts of forest zone, including Nimba, Taı¨ and Danane, also present in E in Bossematie´ Forest and in Comoe´ Nat. Park; Ghana in forest zone north to Bia Nat. Park and Mampong; Togo, 1 record (Misaho ¨he), presumed rare resident (Cheke and Walsh 1996); 3 records Nigeria, Ibadan, Oban Hills, and recently at Boje-Enyi, Cross River State (Bull. Afr. Bird Club 2, 126). Forest zone of Cameroon north to Tchabal Mbabo, west to Korup Nat. Park, east to Lobe´ke´, but scarce or absent in extreme S; known from Yaounde´ but not seen by Quantrill and Quantrill (1998), and not recorded for Mbini or adjacent N Gabon; Bioko; Central African Republic in SW and SE (Baroua), north to Sangba; Gabon, locally abundant around Makokou, and listed for Doume´, Ogooue district (Bannerman 1949), doubtless more widespread; Congo, known from Nouabale´-Ndoki Nat. Park (common), Odzala Nat. Park (rare) and in S at Sibiti and Dimonika; Zaı¨re, widespread in N from N Ubangui (frequent: Dejaifve 1994), Uele and Ituri, and in E south to Itombwe (very common); known from widely scattered localities elsewhere, lower Congo in W, Upemba Nat. Park and Kasaji in S, doubtless more widespread than shown on map; uncommon in Angola from Uı´ge and Luhanda (Malanje) south to Quitondo and Gabela in Cuanza Sul; E Rwanda (valleys of Akagera, Kibaya, Kagogo and Nyaruvomo rivers, around lakes Mhanga and Mugesera, and near Kigali: vande Weghe 1981b); not reported from Nyungwe Forest; Burundi (Musigati, Kigwena, Bururi Forest); common extreme S Sudan (Bengengai Forest, Imatong and Dongotona Mts, up to 2400 m); Ethiopia in W Highlands (upper reaches of Omo R.), Rift Valley, including Nechisar Nat. Park (frequent), and recently at Wendo Genet (Farnsworth et al. 2000), and in S at Alghe and Arero, generally uncommon; widespread W and S Uganda from Budongo,

Mandingoa nitidula

?

Bugoma, Bwamba (abundant), Maramagambo, KasyohaKitomi, Impenetrable (Bwindi) and Sango Bay forests east to Entebbe, Kifu and Mabira, also in N (Kidepo Valley Nat. Park) and extreme NW (Otze Forest White Rhino Sanctuary: Forbes-Watson 1994); Kenya along coast from Tana R. to Vanga, inland on Shimba, Taita and Ngulia Hills, central highlands (N Nairobi, Kiambu, Aberdares, Meru Forest, and Kikuyu Escarpment Forest at 2350 m (Taylor and Taylor 1988); status uncertain: periodic reports from Mt Elgon, Kerio Valley, Kakamega, North Nandi Forest, Mara Game Res. and Lolgorien may in part represent wandering or displaced birds (Zimmerman et al. 1996); formerly on Mt Marsabit. Tanzania, widespread in coastal forests, inland to Eastern Arc mts, North Pares

276

ESTRILDIDAE

(1500–1750 m), W and E Usambaras, Ngurus (1500 m), Ukagurus (1500–1850 m), Ulugurus (300–1500 m), Udzungwas (750–1000 m), and to Arusha District, including Mt Meru; scattered localities in W (Bukoba District including Minziro Forest), Ukerewe I., Gombe and Mahali Nat. Parks; recently found in Serengeti on Grumeti R. (Bull. Afr. Bird Club 8, 66). Zanzibar and Pemba; Zambia as ˆ i at 500– mapped, apparently rare; widespread Malaw 1500 m, up to 2120 m on Nyika Plateau; E Zimbabwe up to 1200 m, south to Mt Selinda; occurs in Manica e Sofala and other parts of central Mozambique, formerly along coast of Sul do Save but has declined there due to destruction of coastal forest and capture for cage bird trade, population now 2600 m) and Mt Kenya through Nandi, Mau and Trans-Mara Forests to Nguruman, Namanga and Chyulu Hills; said to be formerly in Nairobi (Zimmerman et al. 1996) but still there, since observed in Nairobi Arboretum in May 2000 (Kenya Birds 9, 2002, 36); N Tanzania in Loliondo area, Longido, Mt Meru and Kilimanjaro. Density in primary forest 1 bird per ha at Kijabe, Kenya (Carlson 1986), 17 birds per 100 ha at Chilomo, Ethiopia (Tadesse et al. 2001). Description. C. s. kilimensis Sclater (includes ‘chyuluensis’): S Sudan to E Uganda (Elgon), W and central Kenya (north to Mt Kenya), SE Kenya (Chyulus) and N Tanzania; intergrades with nominate race in N Kenya. ADULT Y: forehead to upper mantle dark greyish olive; lower mantle and scapulars to uppertailcoverts dark crimson; tail blackish. Lores, around eye, cheeks, ear-coverts and sides of neck greyish olive; chin and throat light olive-buff, merging into olive-grey underparts, darkest on belly and undertail-coverts; lower flanks dark crimson. Remiges, alula and primary coverts dark grey-brown, outer webs of tertials dark crimson; inner greater coverts broadly fringed crimson, outer feathers mainly olive with narrow crimson outer edge; median and lesser coverts dark greyish olive, some inner medians tipped crimson. Axillaries and underwing-coverts greyish white; inner borders of undersides of flight feathers pale grey. Bill black; eyes brown, rim of eyelids red or pink; legs brown. ADULT X: like ad. Y, but with less crimson on wing coverts and little on flanks; lacks pink eye-rim. SIZE (10 YY, 10 XX): wing, 56–61 (579), X 56–58 (570); tail, Y 40–44 (428), X 40–44 (419); bill, Y 12–13 (126), X 115–13 (124); tarsus, Y 17–185 (179), X 165–185 (175). WEIGHT: Sudan, Y (n ¼ 25) 11–145 (122), X (n ¼ 28) 11–15 (128); W Kenya, Y (n ¼ 8) 123–15 (135), X (n ¼ 5) 112–142 (129), unsexed (n ¼ 19) 114–157 (128). IMMATURE: juvenile like ad. X, but red less intense, more cinnamon-rufous; confined to back, rump, uppertail-coverts, tips of mantle and scapular feathers and outer webs of tertials; only a tinge on flanks. NESTLING: skin and natal down pale; 4 tubercles near corner of mouth, 1 on elongate pad on base of upper mandible, the other 3 at gape; inside mouth like Red-faced Crimsonwing C. reichenovii, mediolateral palate spots elongate, tongue with 2 spots (R. B. Payne, pers. comm.). C. s. salvadorii Reichenow: Ethiopia to N Kenya, intergrading with kilimensis at Mt Uraguess. Slightly darker above than

Cryptospiza salvadorii

kilimensis; darker olive-grey below with contrasting pale buffish chin and throat. C. s. ruwenzori Sclater: Albertine Rift mts. Head and neck greyer; underparts generally greyer and paler.

Field Characters. Length 10–11 cm. Dull olive with dark crimson back, rump and wings; distinguished from other crimsonwings by lack of red on head, except that Y has conspicuous red eye-ring; X similar but with paler lores and chin but no eye-ring, and lacks the definitive pale face patch of X Red-faced Crimsonwing. Juv. slightly greyer than juv. Red-faced Crimsonwing, but perhaps not safely separable in the field. Voice. Tape-recorded (B, McVIC). Contact call from birds foraging in undergrowth, an irregular, very soft, barely audible ‘tsip’; when flushed, louder ‘chip-chip’; in bolder contact calls the ‘tsip’ is followed by a rapid, thin metallic trill lasting c. 1 s, ‘tsip . . . tissssssssss’. During courtship display, ‘tsig-tsig’ or ‘tseeg-tseeg’, possibly a variant of contact call; song (captivity), soft, melodious but almost plaintive ‘dee-goo-goo-day-dee’ (R. Burkard in Goodwin 1982). General Habits. Inhabits undergrowth of montane forest, both in interior and around clearings and tree-falls, along tracks and at edges; on Mt Kilimanjaro, extremely common on W slopes (1700–2500 m), less common on S slopes (1600–2900 m), probably indicating preference for dry forest or less intact forest habitats (Cordeiro 1994);

Cryptospiza reichenovii habitat on Mt Morongole dry montane forest; in Ethiopia, olive-Podocarpus-juniper forest, possibly also lowland subtropical humid forest (Urban and Brown 1971). Occurs in bamboo, in secondary growth (e.g. bracken briar, Sericostachys) at forest edges and along streams, sometimes in riparian thickets in savanna. Altitudinal range 1500–3000 m. Singly, in pairs or small groups of up to 6; at Mau Narok, Kenya, occurs after Aug in flocks of up to 50, feeding on tall grasses at edge of cultivation (Sessions 1966). Forages on low grasses or fallen seeds, on ground and just above it, often along forest tracks; likes lower sprays of grass seeds which have bent over onto leaves on forest floor. Shy and secretive; when disturbed slips away into mid-level foliage. Wanders to lower elevations as grass seeds ripen.

Breeding Habits. Courtship and greeting displays as in Red-faced Crimsonwing (Goodwin 1982). NEST: ball or oval, with side opening with or without 5 cm long tunnel, of grass, twigs and tendrils, covered with moss, ext. diam. 150–180; 2–4 m above ground in forest tree, top of sapling, or liana cluster. EGGS: 3–5; white with minute grey spots. SIZE: (n ¼ 1) 164  125. LAYING DATES: Zaı ¨re, June; Sudan, Oct–Jan; Ethiopia, Aug; E Africa: Kenya, Dec (juvs May–July), Tanzania, Feb, Oct, Region A, Oct–Dec, Region D, Jan–Feb, Aug, Nov; all records in dry months following rains.

Food. Seeds of grasses, including Setaria, and of Impatiens; probably also some insects.

Key References (1996).

281

Goodwin, D. (1982), Zimmerman D. A. et al.

Cryptospiza reichenovii (Hartlaub). Red-faced Crimsonwing. Se´ne ´gali de Reichenow.

Plate 17

Pytelia reichenovii Hartlaub, 1874. Ibis, p. 166; Bondongo, Cameroon.

(Opp. p. 266)

Range and Status. Endemic resident, scarce to locally common or abundant. Extreme SE Nigeria (Obudu Plateau, and at >1350 m on Leinde Fadali); Cameroon, widespread in highlands from Mt Oku, where fairly common at 2100–2850 m, to Mt Manenguba (2100 m), Mt Kupe´ (850–1950 m), Rumpi Hills Forest Res. at 1000– 1550 m) (up to 1700 m on Dikume Balue), and Mt Cameroon (900–2000 m); Bioko; Angola around Gabela. Mountains of Albertine Rift up to 2500 m, from S Lendu Plateau (Hall and Moreau 1970) and Ruwenzoris (Zaı¨re/ Uganda) and nearby Kibale Forest, south to Impenetrable (Bwindi) Forest (Uganda), Kivu Volcanos and Itombwe (Zaı¨re), Volcanos, Nyungwe and Cyamudongo (Rwanda), Rwegura, valleys of Lua and Ruhwa rivers, Teza (Burundi) and Mahali Mt (Tanzania). Widespread and locally abundant in highlands of E Tanzania at 900–2520 m (down to 300 m in E Usambaras), from Crater Highlands, Mt Meru, Kilimanjaro (1750 m), and North Pares (1500– 1900 m) to Mt Rungwe, Njombe and Songea; in Zambia ˆ i, mainly above only on Nyika Plateau; widespread Malaw 1360 m, sometimes down to 900 m; Mozambique on Chiperone, Namuli and Gorongoza Mts and highlands along Zimbabwe border; common in E highlands of Zimbabwe at 1400–2000 m from Inyanga and Vumba to Melsetter and Chimanimani Mts; may even be able to extend its range locally where there is sufficient undergrowth (Irwin 1981). Description. C. r. reichenovii (Hartlaub): Bioko, Cameroon, Angola. ADULT Y: forehead to upper mantle dark olive; lower mantle and scapulars to uppertail-coverts dark crimson, tail blackish. Broad patch through lores and around and behind eye crimson; lower cheeks, ear-coverts and sides of neck dark olive; chin pale olive-buff, merging into dark olive underparts, more blackish on belly and undertail-coverts; lower flanks dark crimson; undertail-coverts often tinged crimson. Remiges, primary coverts and alula blackish brown, outer edge of innermost

Cryptospiza reichenowii

secondary often crimson, outer webs of tertials dark crimson; inner greater coverts broadly edged and tipped crimson, outer feathers mainly dark olive with crimson confined to narrow outer edge; median coverts dark olive, inner feathers broadly tipped crimson; lesser coverts dark olive. Axillaries and underwingcoverts olive-grey; inner borders of undersides of flight feathers pale grey. Bill black; eyes dark brown; legs dark brown or greybrown, soles of feet yellowish. ADULT X: similar to ad. Y but lacks red face patch; lores and around eye pale olive-buff, separated from olive-buff chin by dark olive band across lower cheek. Crimson edges of greater and median coverts narrower. Partial

Cryptospiza reichenovii habitat on Mt Morongole dry montane forest; in Ethiopia, olive-Podocarpus-juniper forest, possibly also lowland subtropical humid forest (Urban and Brown 1971). Occurs in bamboo, in secondary growth (e.g. bracken briar, Sericostachys) at forest edges and along streams, sometimes in riparian thickets in savanna. Altitudinal range 1500–3000 m. Singly, in pairs or small groups of up to 6; at Mau Narok, Kenya, occurs after Aug in flocks of up to 50, feeding on tall grasses at edge of cultivation (Sessions 1966). Forages on low grasses or fallen seeds, on ground and just above it, often along forest tracks; likes lower sprays of grass seeds which have bent over onto leaves on forest floor. Shy and secretive; when disturbed slips away into mid-level foliage. Wanders to lower elevations as grass seeds ripen.

Breeding Habits. Courtship and greeting displays as in Red-faced Crimsonwing (Goodwin 1982). NEST: ball or oval, with side opening with or without 5 cm long tunnel, of grass, twigs and tendrils, covered with moss, ext. diam. 150–180; 2–4 m above ground in forest tree, top of sapling, or liana cluster. EGGS: 3–5; white with minute grey spots. SIZE: (n ¼ 1) 164  125. LAYING DATES: Zaı ¨re, June; Sudan, Oct–Jan; Ethiopia, Aug; E Africa: Kenya, Dec (juvs May–July), Tanzania, Feb, Oct, Region A, Oct–Dec, Region D, Jan–Feb, Aug, Nov; all records in dry months following rains.

Food. Seeds of grasses, including Setaria, and of Impatiens; probably also some insects.

Key References (1996).

281

Goodwin, D. (1982), Zimmerman D. A. et al.

Cryptospiza reichenovii (Hartlaub). Red-faced Crimsonwing. Se´ne ´gali de Reichenow.

Plate 17

Pytelia reichenovii Hartlaub, 1874. Ibis, p. 166; Bondongo, Cameroon.

(Opp. p. 266)

Range and Status. Endemic resident, scarce to locally common or abundant. Extreme SE Nigeria (Obudu Plateau, and at >1350 m on Leinde Fadali); Cameroon, widespread in highlands from Mt Oku, where fairly common at 2100–2850 m, to Mt Manenguba (2100 m), Mt Kupe´ (850–1950 m), Rumpi Hills Forest Res. at 1000– 1550 m) (up to 1700 m on Dikume Balue), and Mt Cameroon (900–2000 m); Bioko; Angola around Gabela. Mountains of Albertine Rift up to 2500 m, from S Lendu Plateau (Hall and Moreau 1970) and Ruwenzoris (Zaı¨re/ Uganda) and nearby Kibale Forest, south to Impenetrable (Bwindi) Forest (Uganda), Kivu Volcanos and Itombwe (Zaı¨re), Volcanos, Nyungwe and Cyamudongo (Rwanda), Rwegura, valleys of Lua and Ruhwa rivers, Teza (Burundi) and Mahali Mt (Tanzania). Widespread and locally abundant in highlands of E Tanzania at 900–2520 m (down to 300 m in E Usambaras), from Crater Highlands, Mt Meru, Kilimanjaro (1750 m), and North Pares (1500– 1900 m) to Mt Rungwe, Njombe and Songea; in Zambia ˆ i, mainly above only on Nyika Plateau; widespread Malaw 1360 m, sometimes down to 900 m; Mozambique on Chiperone, Namuli and Gorongoza Mts and highlands along Zimbabwe border; common in E highlands of Zimbabwe at 1400–2000 m from Inyanga and Vumba to Melsetter and Chimanimani Mts; may even be able to extend its range locally where there is sufficient undergrowth (Irwin 1981). Description. C. r. reichenovii (Hartlaub): Bioko, Cameroon, Angola. ADULT Y: forehead to upper mantle dark olive; lower mantle and scapulars to uppertail-coverts dark crimson, tail blackish. Broad patch through lores and around and behind eye crimson; lower cheeks, ear-coverts and sides of neck dark olive; chin pale olive-buff, merging into dark olive underparts, more blackish on belly and undertail-coverts; lower flanks dark crimson; undertail-coverts often tinged crimson. Remiges, primary coverts and alula blackish brown, outer edge of innermost

Cryptospiza reichenowii

secondary often crimson, outer webs of tertials dark crimson; inner greater coverts broadly edged and tipped crimson, outer feathers mainly dark olive with crimson confined to narrow outer edge; median coverts dark olive, inner feathers broadly tipped crimson; lesser coverts dark olive. Axillaries and underwingcoverts olive-grey; inner borders of undersides of flight feathers pale grey. Bill black; eyes dark brown; legs dark brown or greybrown, soles of feet yellowish. ADULT X: similar to ad. Y but lacks red face patch; lores and around eye pale olive-buff, separated from olive-buff chin by dark olive band across lower cheek. Crimson edges of greater and median coverts narrower. Partial

284

ESTRILDIDAE

albinos and other aberrantly plumaged birds known (Sclater and Moreau 1932–1933). SIZE (10 YY, 5 XX, Cameroon): wing, Y 52– 55 (539), X 52–57 (538); tail, Y 35–41 (383), X 36–41 (394); bill, Y 125–135 (130), X 125–14 (130); tarsus, Y 17–19 (181), X 17– 19 (181). WEIGHT: Cameroon, Y (n ¼ 9) 10–15 (127), YX (n ¼ 12) 11–13 (116). IMMATURE: juvenile similar to ad. X but lacks pale patch around eye; underparts duller, more greyish; crimson less bright, confined to tertial edges, tips of scapulars and lower mantle feathers, and back, rump and uppertail-coverts; little on upperwingcoverts. NESTLING: at c. 7 days, skin pinkish slate, silver-grey pin feathers on wings, tail and triangle at base of neck, white fluff above eyes, bill and feet blue-grey, eye dark brown; gape pale yellow with 4 tiny tubercles on each side outlined in black, palate pale yellow with 3 black crescents in a line and below and between them 2 short vertical black strokes, tongue with 2 vertical black marks side by side (W. M. Moreau in Sclater and Moreau 1933). C. r. ocularis Sharpe: mts of Albertine Rift. Like nominate race, but on average slightly paler olive below. WEIGHT: Rwanda, Y (n ¼ 6) 112–13 (123), X (n ¼ 3) 117–137 (126). C. r. australis Shelley (including ‘homogenes’): Tanzania to Mozambique and Zimbabwe. Distinctly paler and greyer below than nominate race. WEIGHT: N Tanzania, Y (n ¼ 20) 106–157 (128), X (n ¼ 20) 95–148 (128); S Tanzania Y (n ¼ 23) 115–17 (130), X (n ¼ 14) 105–145 (128); southern Africa, Y (n ¼ 152) 115–162 (135), X (n ¼ 125) 112–174 (133). Max. diurnal weight change 15 g (118% of sample mean weight) (Dowsett 1983).

Field Characters. Length 10–11 cm. Olive-brown with dark crimson upperparts, wings and flanks. Distinguished from similar Abyssinian Crimsonwing C. salvadorii, which it overlaps on Kilimanjaro and Mt Meru, by face patches (red in Y, pale in X); immature without any indication of eye mask difficult to separate from Abyssinian Crimsonwing but less grey (Stevenson and Fanshawe 2002). Tail black (Lesser Seedcracker Pyrenestes minor has brown back, red tail). Keeps to shade, where red colour difficult to see. Voice. Tape-recorded (99, 104, B, C, F, GRI, GUT, SVEN). Main call high, thin, metallic ‘sit-sit’, often repeated rapidly in trills of different lengths; also gives louder, lower ‘sweeee’ or ‘tzeet’. Tiny song audible for only c. 10 m, 4 long-drawn soft, sweet notes descending in major thirds, each followed by a staccato chirp, first note quavering and plaintive, last so low as to be inaudible to some human ears (Sclater and Moreau 1932–1933). Another song contains a long note with curious nasal complaining quality, preceded by tikking notes, ‘tikatik-wraaaaangh’, with or without final ‘tu’. Low conversational chuckling notes in captivity, ‘chup’, ‘tslip’, ‘tuk-a-tuk’. Sings during courtship display and when perched near mate or fledged young. General Habits. Inhabits dense undergrowth and shrubbery, including tree ferns, in mature montane forest, both closed and open, especially along roads, around clearings and at forest edge; within forest often along streams; on ˆ i, favours thick carpets of AcanthaNyika Plateau, Malaw ceae or other subshrubs and herbs near streams (DowsettLemaire 1983); at forest edge, thickets of bracken-briar, tall heath Philippia, bracken under Leucosidea bushes; also

plantations and moss forest (Bioko), open areas with herbaceous vegetation (Cameroon, Mt Kupe´: Bowden 2001), old cultivation with Bidens pilosus and other weeds, millet and maize patches and scrub around farms (Zaı¨re), overgrown gardens (Mozambique), plantations of tea and of exotic conifers (Zimbabwe); at Amani, Tanzania, only exceptionally outside forest shade, to a maximum distance of 90 m (Sclater and Moreau 1932–1933). Pairs and parties of up to 10; sometimes with Western Bluebill Spermophaga haematina, Black-crowned Waxbill Estrilda nonnula and Black-and-white Mannikin Spermestes bicolor. Silent, shy; forages on ground or just above it, in undergrowth or on grass beside roads or in streambeds. In pine plantations extracts seeds from cones in trees up to 15 m or more above ground (Irwin 1981). Drinks and bathes frequently. In captivity very active for first 2 and last 2 h of daylight, moving incessantly; also in captivity, birds ‘anted’ with millipedes but not with ants (P. Kunkel in Goodwin 1982). Occurrence at sea level in Cameroon suggests some birds move to lower levels in non-breeding season. Altitudinal migration down to lowland forest during dry/cold season recorded for 4 mountain ranges in Tanzania: Pares, Ukagurus/Ngurus, Rubehos and Udzungwas (Burgess and Mlingwa 2000). Some local dispersal outside breeding season (up to 3 km by ringed bird) on Nyika Plateau, ˆ i (Dowsett-Lemaire 1983). Malaw Food. Insects; seeds, especially of grasses, including Setaria chevaleri and S. longiseta and Panicum spp. including P. adenophorum and P. maximum, Streblochaete longiarista, Acalypha psilostachya, Hagenia abyssinica, Hypoestes verticillaris, Urera hypsilodendron; small green bean-like seeds of Impatiens spp., pine cone seeds. e.g. Pinus patula, fragments of pounded maize left by villagers; young birds also take other parts of plants (Eisentraut 1963). Seen feeding among flowers of Leucosidea (Ginn et al. 1989). In captivity, besides seeds, insect food, e.g. ant pupae, whiteworms, gnat larvae and mealworms, also spiders. ˆ i, Nyika: Breeding Habits. Territory size 08–4 ha (Malaw Dowsett-Lemaire 1983). Every year birds nest in same area, even in same tree, and 3- or 4-year-old nests are often present near current nest (Zimbabwe: Ginn et al. 1989). Courtship display like that of Estrilda spp.; display performed by both sexes, with or without nest material in bill, Y singing, X silent; displaying bird raises feathers of breast, flanks and lower belly and cocks tail, turns toward partner and thrusts head and body upwards by stretching and bending legs; head is held stiffly up, not thrown back; display may end in copulation but often does not. In another display, Y perches next to X and sings to her with head and body held stiffly up; X sometimes responds by adopting similar stance and quivering tail. Y also sings like this to flying but still dependent young, which gather round him and listen attentively (Goodwin 1982). In greeting display stands upright with breast and belly feathers partly erected, tail cocked, head turned slightly towards partner with feathers arranged to make it appear triangular; birds then hop sideways toward each other.

Estrilda NEST: large untidy ball or oval, with wide side entrance pointing slightly upwards, of grass blades, plant stems, mosses, dead leaves and leaf skeletons and fine dry grass, giving it appearance of old nest (Fuggles-Couchman 1986), lined with feathery seed heads of small grass Panicum, threads of Marasmius, or feathers; (n ¼ 1) ext. diam. 96–120, int. diam. 48, entrance tunnel 48 long, another ext. diam. 144; placed 3–6 m above ground at edge of forest or clearing, in tree fern (e.g. Cyathea usambarensis) or forest sapling (e.g. Cylicomorpha parviflora), once in creepers overhanging riverside pool, once in cluster of dead vines hanging c. 8 m above ground; sites in Zimbabwe include cocoa tree Erythroxylum emarginatum, tangle of creepers, Hypericum scrub beside stream, and (often) thorny shrub Maytenus heterophylla under overhanging branch (nest record cards, National Museum, Bulawayo); built by X with material brought by Y (Rwanda), in captivity mainly by Y. EGGS: 3–5, white. SIZE: (n ¼ 5) 167–171  118–122 (169  l20). LAYING DATES: Nigeria (breeding condition Dec); Cameroon, Oct–Dec (juvs Feb); Angola (juvs May, Sept);

Zaı¨re, Apr–June, Aug, Oct (juv. Mar); Rwanda (nestbuilding Jan); E Africa: Tanzania, ‘practically every month of the year’ (Amani: Moreau 1936), (elsewhere nestbuilding Feb, brood patch July–Aug), Region B Mar; ˆ i, Mar–July, Sept (juvs Aug–Oct); Mozambique Malaw Oct–Mar; Zimbabwe Sept–Mar. INCUBATION: by both sexes. DEVELOPMENT AND CARE OF YOUNG: young brooded and fed by both parents; fledge at 21 days, fed by Y for another 10–12 days; beg by stretching necks straight towards parent; for first few nights after fledging, young called back to nest to roost by parents, but not all come, some roosting outside. BREEDING SUCCESS/SURVIVAL: ringing returns on Nyika ˆ i, show minimum age of oldest bird 45 Plateau, Malaw years, one territory held at least 4 seasons (Dowsett 1985); ad. Y recaptured after 5 years 140 days (Zimbabwe: Harwin et al. 1994). Key References Goodwin D. (1982), Maclean, G. L. (1993), Sclater, W. L. and Moreau, R. E. (1932–1933).

Genus Estrilda Swainson

Small estrildines, sexes alike or nearly so, many spp. with finely barred brownish plumage, red or carmine rump and flanks; bill robust, black or waxy red; wings short and rounded; tail graduated, rectrices slender, not noticeably short. Gregarious, active, adept at extracting seeds from grass heads; live amongst grasses, often forage on ground; a few spp. somewhat arboreal. Nest sited on ground in grass tuft or (3 spp.) a few m up in thorn tree; round, made almost entirely of grass stems and inflorescences, with side entrance at end of short passage, protected and concealed by ring of protruding, converging grass stems, usually with remarkable false nest (so-called cock’s nest) built on top. Cock’s nest or false nest is like real nest but usually less substantial, with wide, obvious entrance. Function has been controversial, but is now generally agreed to distract predators from real nest. Parent can only enter real nest tunnel by crouching mouse-like, and once in, pliant material around tunnel entrance closes. It may be hard for predator to enter to take incubating bird or eggs, but there is no other escape and the survival stratagem is to deflect predator (compare false entrance of penduline tits’ nests, Remizidae, Vol. VI, p. 106). Bits of earth and excrement placed on false nest and feathers and eggshells around and inside its entrance, together with fussy behaviour around and inside false nest of bird arriving to relieve mate incubating in real nest, serve to distract and mislead predator, as does pungent smell of dead nestling that can be wiped on outside of nest and ‘built into’ wall of false nest (in captive E. troglodytes real nest is kept scrupulously clean, but in wild E. astrild it is not) (I. Hinze, pers. comm.). Estrilda is sister clade to Mandingoa/Cryptospiza. Taxonomically controversial, because each of the 5 distinct superspecies or species-groups has populations separated distantly, sedentary, not strongly differentiated in plumage characters, but in some cases with small but seemingly meaningful differences (voice, habitat, structure, facial features) that have led to marked tendency to upgrading from subspecies to megasubspecies to full species (either allospecies or sibling species or independent species). Favourite cagebirds, whose biology is often better known from aviary than from wild studies. Near endemic; c. 16 species (E. rufibarba Arabian, remainder African) in 5 alliances: (i) E. caerulescens/E. perreini/E. thomensis superspecies (pale grey, unbanded; bill black or part carmine); (ii) E. paludicola/E. poliopareia superspecies and E. melpoda (brown above, pale below; bill red; E. melpoda with large orange face patch; E. poliopareia with faint banding and pale eye); under the phylogenetic species concept, E. paludicola might become 5 allospecies, of which ochrogaster is the most distinct (though less so than E. poliopareia); (iii) E. troglodytes/E. rufibarba/E. astrild/E. nigriloris superspecies (first 2 blacktailed and in arid habitats, last 2 brown-tailed and in mesic habitats; pale brown and pink, banded; eyestripe red, or black in nigriloris; bill red, or blackish in rufibarba) and E. rhodopyga (bill blackish); (iv) E. atricapilla/E. kandti superspecies and E. nonnula (banded; black-capped; bill black-and-red; mutual relations of much interest); and (v) E. charmosyna/E. erythronotos superspecies (banded; black-faced; bill bluish). Genetic structure of 12 taxa tested by R. B. Payne indicates rather different alliances: group (a1) paludicola, troglodytes, melpoda, rhodopyga; (a2) astrild, (a3) nonnula and atricapilla very closely allied;

285

Estrilda NEST: large untidy ball or oval, with wide side entrance pointing slightly upwards, of grass blades, plant stems, mosses, dead leaves and leaf skeletons and fine dry grass, giving it appearance of old nest (Fuggles-Couchman 1986), lined with feathery seed heads of small grass Panicum, threads of Marasmius, or feathers; (n ¼ 1) ext. diam. 96–120, int. diam. 48, entrance tunnel 48 long, another ext. diam. 144; placed 3–6 m above ground at edge of forest or clearing, in tree fern (e.g. Cyathea usambarensis) or forest sapling (e.g. Cylicomorpha parviflora), once in creepers overhanging riverside pool, once in cluster of dead vines hanging c. 8 m above ground; sites in Zimbabwe include cocoa tree Erythroxylum emarginatum, tangle of creepers, Hypericum scrub beside stream, and (often) thorny shrub Maytenus heterophylla under overhanging branch (nest record cards, National Museum, Bulawayo); built by X with material brought by Y (Rwanda), in captivity mainly by Y. EGGS: 3–5, white. SIZE: (n ¼ 5) 167–171  118–122 (169  l20). LAYING DATES: Nigeria (breeding condition Dec); Cameroon, Oct–Dec (juvs Feb); Angola (juvs May, Sept);

Zaı¨re, Apr–June, Aug, Oct (juv. Mar); Rwanda (nestbuilding Jan); E Africa: Tanzania, ‘practically every month of the year’ (Amani: Moreau 1936), (elsewhere nestbuilding Feb, brood patch July–Aug), Region B Mar; ˆ i, Mar–July, Sept (juvs Aug–Oct); Mozambique Malaw Oct–Mar; Zimbabwe Sept–Mar. INCUBATION: by both sexes. DEVELOPMENT AND CARE OF YOUNG: young brooded and fed by both parents; fledge at 21 days, fed by Y for another 10–12 days; beg by stretching necks straight towards parent; for first few nights after fledging, young called back to nest to roost by parents, but not all come, some roosting outside. BREEDING SUCCESS/SURVIVAL: ringing returns on Nyika ˆ i, show minimum age of oldest bird 45 Plateau, Malaw years, one territory held at least 4 seasons (Dowsett 1985); ad. Y recaptured after 5 years 140 days (Zimbabwe: Harwin et al. 1994). Key References Goodwin D. (1982), Maclean, G. L. (1993), Sclater, W. L. and Moreau, R. E. (1932–1933).

Genus Estrilda Swainson

Small estrildines, sexes alike or nearly so, many spp. with finely barred brownish plumage, red or carmine rump and flanks; bill robust, black or waxy red; wings short and rounded; tail graduated, rectrices slender, not noticeably short. Gregarious, active, adept at extracting seeds from grass heads; live amongst grasses, often forage on ground; a few spp. somewhat arboreal. Nest sited on ground in grass tuft or (3 spp.) a few m up in thorn tree; round, made almost entirely of grass stems and inflorescences, with side entrance at end of short passage, protected and concealed by ring of protruding, converging grass stems, usually with remarkable false nest (so-called cock’s nest) built on top. Cock’s nest or false nest is like real nest but usually less substantial, with wide, obvious entrance. Function has been controversial, but is now generally agreed to distract predators from real nest. Parent can only enter real nest tunnel by crouching mouse-like, and once in, pliant material around tunnel entrance closes. It may be hard for predator to enter to take incubating bird or eggs, but there is no other escape and the survival stratagem is to deflect predator (compare false entrance of penduline tits’ nests, Remizidae, Vol. VI, p. 106). Bits of earth and excrement placed on false nest and feathers and eggshells around and inside its entrance, together with fussy behaviour around and inside false nest of bird arriving to relieve mate incubating in real nest, serve to distract and mislead predator, as does pungent smell of dead nestling that can be wiped on outside of nest and ‘built into’ wall of false nest (in captive E. troglodytes real nest is kept scrupulously clean, but in wild E. astrild it is not) (I. Hinze, pers. comm.). Estrilda is sister clade to Mandingoa/Cryptospiza. Taxonomically controversial, because each of the 5 distinct superspecies or species-groups has populations separated distantly, sedentary, not strongly differentiated in plumage characters, but in some cases with small but seemingly meaningful differences (voice, habitat, structure, facial features) that have led to marked tendency to upgrading from subspecies to megasubspecies to full species (either allospecies or sibling species or independent species). Favourite cagebirds, whose biology is often better known from aviary than from wild studies. Near endemic; c. 16 species (E. rufibarba Arabian, remainder African) in 5 alliances: (i) E. caerulescens/E. perreini/E. thomensis superspecies (pale grey, unbanded; bill black or part carmine); (ii) E. paludicola/E. poliopareia superspecies and E. melpoda (brown above, pale below; bill red; E. melpoda with large orange face patch; E. poliopareia with faint banding and pale eye); under the phylogenetic species concept, E. paludicola might become 5 allospecies, of which ochrogaster is the most distinct (though less so than E. poliopareia); (iii) E. troglodytes/E. rufibarba/E. astrild/E. nigriloris superspecies (first 2 blacktailed and in arid habitats, last 2 brown-tailed and in mesic habitats; pale brown and pink, banded; eyestripe red, or black in nigriloris; bill red, or blackish in rufibarba) and E. rhodopyga (bill blackish); (iv) E. atricapilla/E. kandti superspecies and E. nonnula (banded; black-capped; bill black-and-red; mutual relations of much interest); and (v) E. charmosyna/E. erythronotos superspecies (banded; black-faced; bill bluish). Genetic structure of 12 taxa tested by R. B. Payne indicates rather different alliances: group (a1) paludicola, troglodytes, melpoda, rhodopyga; (a2) astrild, (a3) nonnula and atricapilla very closely allied;

285

286

ESTRILDIDAE

(b1) caerulescens, thomensis; (b2) perreini; (b3) erythronotos and delamerei conspecific. Alliance (i) formerly held to link with Lagonosticta, and its members were formerly placed in that genus; resembles Lagonosticta in some aspects of behaviour, form of bill, P9 not emarginate, and shortness of tail (Chapin 1954), but not at all closely related at molecular level (R. B. Payne pers. comm.).

Estrilda paludicola superspecies

Estrilda caerulescens superspecies

1

1

2

1 E. caerulescens 2 E. thomensis 3 E. perreini

3

2

1 E. poliopareia 2 E. paludicola

3

2 3

Estrilda astrild superspecies

Estrilda atricapilla superspecies

2 1

2 1

1

2

2

1,2

1 1,2

2 1 2 1

1 2

2 1 E. troglodytes 2 E. astrild 3 E. nigriloris

1 E. atricapilla 2 E. kandti

3

2

2

1

Estrilda caerulescens

287

Estrilda charmosyna superspecies

1

2 1 E. charmosyna 2 E. erythronotos

2

Estrilda caerulescens (Vieillot). Lavender Waxbill. Astrild queue-de-vinaigre.

Plate 18

Fringilla caerulescens Vieillot, 1817. Nouv. Dict. Hist. Nat., nouv. ´ed., 12, p. 176; Zone Torride, Senegal.

(Opp. p. 267)

Forms a superspecies with E. perreini and E. thomensis. Range and Status. Endemic resident, W Africa. Introduced to Hawaii. Senegal, near Saint-Louis, Lingue`re, Tambacounda, Dakar and Thie`s in NW; otherwise barely north of 14
N, but south of it frequent or uncommon but widespread, east to Niokolo-Koba Nat. Park. Gambia, locally common in most areas. Mali, records from 65 localities, southwest of 15
N and 3
W, mainly southwest of 13
N and 8
W; commonest in Mandingo Mts (Lamarche 1993). Guinea-Bissau, widespread: Sa˜o Dominges, Bula, Mansoa, Farim, Contabane, Dandum (Rodwell 1996). Guinea, Koundara and Gaoual areas (Morel and Morel 1988), possibly Conakry (Richards 1982). Liberia, rare, Mt Nimba, 1967–1968. Ivory Coast, frequent north of 9
N, e.g. Comoe´ Nat. Park; a few, Abidjan, 1987 (escapes?: Demey and Fishpool 1991). Burkina Faso, rare, Yatenga area; frequent around Ouagadougou; only at 11
170 N, 3
390 W (Holyoak and Seddon 1989). Niger; frequent in ‘W’ Nat. Park and north to 13
N (Korogoungou, Bengou, Boboye). Ghana, widespread and not uncommon in N (Bongo Hills, Sisali [Sissili] valley, Tumu, Gambaga), south to Mole. Togo, uncommon in N, south to Aledjo and Pe´wa (9
150 N). Benin, frequent in Be´te´rou area; uncommon in N, except around Kani and in Arli and Pendjari Nat. Park (across border with Burkina Faso), where common. Nigeria, locally not uncommon, Kainji Lake Nat. Park to Kaiama, north to Zaria, Kano and Maiduguri areas (Gaya, Tamburawa;

Estrilda caerulescens

?

? ?

Estrilda caerulescens

287

Estrilda charmosyna superspecies

1

2 1 E. charmosyna 2 E. erythronotos

2

Estrilda caerulescens (Vieillot). Lavender Waxbill. Astrild queue-de-vinaigre.

Plate 18

Fringilla caerulescens Vieillot, 1817. Nouv. Dict. Hist. Nat., nouv. ´ed., 12, p. 176; Zone Torride, Senegal.

(Opp. p. 267)

Forms a superspecies with E. perreini and E. thomensis. Range and Status. Endemic resident, W Africa. Introduced to Hawaii. Senegal, near Saint-Louis, Lingue`re, Tambacounda, Dakar and Thie`s in NW; otherwise barely north of 14
N, but south of it frequent or uncommon but widespread, east to Niokolo-Koba Nat. Park. Gambia, locally common in most areas. Mali, records from 65 localities, southwest of 15
N and 3
W, mainly southwest of 13
N and 8
W; commonest in Mandingo Mts (Lamarche 1993). Guinea-Bissau, widespread: Sa˜o Dominges, Bula, Mansoa, Farim, Contabane, Dandum (Rodwell 1996). Guinea, Koundara and Gaoual areas (Morel and Morel 1988), possibly Conakry (Richards 1982). Liberia, rare, Mt Nimba, 1967–1968. Ivory Coast, frequent north of 9
N, e.g. Comoe´ Nat. Park; a few, Abidjan, 1987 (escapes?: Demey and Fishpool 1991). Burkina Faso, rare, Yatenga area; frequent around Ouagadougou; only at 11
170 N, 3
390 W (Holyoak and Seddon 1989). Niger; frequent in ‘W’ Nat. Park and north to 13
N (Korogoungou, Bengou, Boboye). Ghana, widespread and not uncommon in N (Bongo Hills, Sisali [Sissili] valley, Tumu, Gambaga), south to Mole. Togo, uncommon in N, south to Aledjo and Pe´wa (9
150 N). Benin, frequent in Be´te´rou area; uncommon in N, except around Kani and in Arli and Pendjari Nat. Park (across border with Burkina Faso), where common. Nigeria, locally not uncommon, Kainji Lake Nat. Park to Kaiama, north to Zaria, Kano and Maiduguri areas (Gaya, Tamburawa;

Estrilda caerulescens

?

? ?

288

ESTRILDIDAE

L. Alo, Ngadda R.), south to Nasarawa, Jos and Yankari Nat. Park. Cameroon, uncommon, from about 12
N to Benue plain. Chad, uncommon: records at Ndjamena, Massenya and Niellim. Central African Republic, uncommon, Manovo-Gounda-Saint Floris and Bamingui-Bangoran Nat. Parks. Description. ADULT Y: forehead to hindneck, mantle, scapulars and upper back grey with slight bluish tinge; lower back, rump and uppertail-coverts crimson. Tail blackish, upperside of T1 and of outer webs of T2–T5 dark crimson. Black streak through lores linked narrowly below eye to short black post-ocular streak. Cheeks and ear-coverts to chin and throat greyish white, merging into grey sides of neck and grading through light grey breast to sooty or blackish grey on lower flanks, vent and thighs; rearmost flank feathers with small white spots near tip; undertail-coverts dull crimson. Flight feathers grey-brown, edges of primaries and outer webs of secondaries grey; rest of upperwing grey. Underwing-coverts and axillaries greyish white. Bill reddish to pinkish grey with tip, culmen, cutting edges and lower part of lower mandible blackish; eyes dark brown; legs dark olive, blackish brown or black. ADULT X: like ad. Y but dark grey lower belly and flanks less blackish. SIZE (9 YY, 6 XX): wing, Y 50–54 (520), X 49–54 (512); tail, Y 37–41 (388), X 37–39 (383); bill, Y 11–125 (118), X 115–125 (117); tarsus, Y 14–155 (147), X 14– 15 (147). WEIGHT: N Nigeria, unsexed (n ¼ 7) 79–92 (85), N Ghana, unsexed (n ¼ 15) 67–91 (80). IMMATURE: juvenile paler than ad., lacks black eye-streak and white spots on flanks; crimson rump duller; less crimson on tail and undertail-coverts. NESTLING: mouth pattern like that of Lagonosticta senegala; gape tubercles bluish white (Goodwin 1982).

Field Characters. Length 10 cm. Uniform light grey with crimson rump, tail and undertail-coverts; lower belly blackish, black line through lores to just behind eye; bill dark brown with pinkish base. Immature paler, reds duller, undertail black and red, loral line faint. Cannot be confused with any bird in its range; does not overlap Grey or Cinderella Waxbills E. perreini and E. thomensis. Voice. Tape-recorded (104, B, BRU, GUT, PAY). Contact call, thin, high-pitched ‘seew-seee’ or ‘see-see-squee-see’. High-intensity contact call of X, ‘tseeeht-tseet’, answered by Y with his ‘song phrase’, a whistled ‘see-you’, second note lower and more melodious (Goodwin 1982). Also gives a short, hard ‘dit-tsit’. Captive bird gave sweet, down-slurred ‘tseeoo’ (song?). Alarm longer and more intense than call, ‘tseeay’ or an explosive ‘squeep’. Song a thin and squeaky series of ‘see-see-see-swee’ notes (Clement et al. 1993). Nest call a soft ‘tseeteek tseeteek teek teek’ (Goodwin 1982). General Habits. Inhabits grassland with scattered bushes and trees; thickets and their edges in woodland, often stony or rocky (Mali); rank patches in more open country; margins of cultivation; places with short grass including road verges; dense scrub in wet areas with Raphia thickets, also mature woodland and dense growth around bases of inselbergs (Nigeria). In pairs or flocks of 6–8, or 20 or more. Forages on ground, often with Uraeginthus bengalus and E. troglodytes, but rather more arboreal than them; said to forage in trees and to feed on pollen at tree flowers, like a sunbird (Thonnerieux et al. 1989). On ground, holds grass stem

under foot and extracts seeds from inflorescence with bill. Active, alert and inquisitive; but notably secretive when nesting. In bush, once hung by one foot whilst investigating a leaf held in the other (Goodwin 1982). Flies usually low down; flight quite fast but does not look strong. Unlike most congeners, does not wag or switch tail from side to side when intending to creep or fly (Kunkel 1959). Roosts in old nests of its own or of other species; in captivity, if roosting in branches, sometimes hangs head downwards (Immelmann et al. 1964). Food. Seeds of grass and small fruits; some insects. In captivity, readily drinks artificial nectar. Breeding Habits. Solitary nester; evidently monogamous. Courtship display: Y, with piece of nest material held by one end in bill, forehead feathers sleeked so that head looks triangular, tail held at an angle, bobs up and down; X responds first by giving greeting display, then by quivering tail. In Greeting (or Nodding) Display, 2 birds perch half facing each other, heads ‘triangular’, tails angled, rump and flank feathers smoothly erected; one bird adopts upright posture and ‘freezes’ for a moment with bill inclined, then nods or bows and again freezes; other bird does the same but usually out of synchrony, so that its head is down when other bird’s head is up; Y sings, X gives contact calls (Goodwin 1982). Newly paired mates perform Greeting Display often, but when bond is established they seldom do. Greeting Display can be between 2 YY unknown to each other, when display may end aggressively with chasing and even violent fighting. NEST: 5 in Gambia untidy, woven domes, 20 cm diam., with downward-pointing entrance spout at one side c. 50 long and 35 in diam., built in forks in citrus trees and garden shrubs (Moore 1983); 1 in Senegal bulky, globular, with long entrance tunnel at top; made of grass; hidden in green creepers well off ground (Barlow et al. 1997). In captivity, builds spherical nest, diam. (n ¼ 5) c. 200, with downward-sloping entrance tunnel c. 50 long and 35 in diam. (once c. 150 long), or sometimes uses old Ploceus weaver nests with tubular entrances; sited in fork in shrub or Citrus tree (Brickell and Konigkramer 1997). Uses feet to hold down piece of material when working it into fabric of nest with bill. Often places feathers, bits of wood, large bird droppings and in particular dead nestlings on top of nest (in captivity. One aviary nest had well-concealed entrance below a grassy overhang on which the birds placed small pieces of cuttlefish bone and soil (Hinze 2000). EGGS: 4–6. White. LAYING DATES: Gambia, Aug–Sept; Burkina Faso, (courtship, Sept); Nigeria, (nest-building, Oct; dependent young, Dec). INCUBATION: by both sexes. Period: 11–12 days (usually 12, in captivity). DEVELOPMENT AND CARE OF YOUNG: for 1st week, young fed entirely on invertebrates; thereafter, insects and seeds (Hinze 2000). Nestlings call loudly whenever parent enters nest with food. Period: 16–19 days. Young out of nest fed by parents for a further 14 days, and do not return to nest to roost.

Estrilda perreini BREEDING SUCCESS/SURVIVAL: 5 nests in one year and 3 in next all either destroyed by heavy rains or abandoned (Moore 1983).

289

Key References Brickell, N. and Konigkramer, T. (1997), Clement, P. et al. (1993), Goodwin, D. (1982), Hinze, I. (2000), Immelmann, K. et al. (1964), Kunkel, P. (1967), Moore, A. (1983).

Estrilda perreini (Vieillot). Grey Waxbill. Astrild a ` queue noire.

Plate 18

Fringilla perreini Vieillot, 1817. Nouv. Dict. Hist. Nat., nouv. ´ed., 12, p. 179; Malimbe, Portuguese Congo.

(Opp. p. 267)

Forms a superspecies with E. caerulescens and E. thomensis. Range and Status. Endemic resident, Gabon, only in Tchibanga area, where rare. Congo, only near Djambala and along lower Congo River. Zaı¨re, frequent around Boma near mouth of Congo R., extending upstream to Brazzaville, Kinshasa and Kwamouth; not uncommon in Katanga, north to Baraka; records from between Fizi and Mulala, NW shores of L. Tanganyika. Tanzania, near Kigoma; S Ufipa Plateau; Iringa Highlands, Marengo Highlands, near Jumbe, twice in Mikumi Nat. Park. Angola, Cabinda, Uı´ge, S Cunza Norte, Malanje, N Bie´, Cuanza Sul, Huambo and along escarpment to NW Huı´la; also in SW Lunda Sul, N Lunda Norte and N Moxico; probably much more widespread than mapped. Zambia, uncommon and local; Northern Prov. from near Muyombe and Old Fife, along Muchinga Escarpment from Muzyatama and Kolala to Danger Hill; also Lavushi Manda and north to Kasama and Mbala; in 26% of 300 ˆ i, only Zambia squares (R.J. Dowsett, pers. comm.). Malaw at Chiromo and Mlolo. Zimbabwe, very local, but can be common; up to 800 m in Honde and Nyamkwarara valleys, uncommon in Muneni valley, abundant up to 700 m in Lusitu-Haroni area; Buzi R. in Chipinga highlands, possibly formerly in Mount Selinda area. Mozambique, very local in Zambezia and Manica e Sofala; uncommon along Sul do Save coastal lowlands. Transvaal, rare and local: Woodbush, Haenertsburg and Tzaneen areas and near lower Sabie. KwaZulu-Natal, littoral plain, extending inland in N to Swaziland (where occurs marginally); recent range extension across Umtanvuna R., Transkei. Perhaps >5000 birds in Sul do Save, Mozambique, where population thought to have declined as result of removal of natural vegetation along coast and of persecution for the cage bird trade, for which c. 800 birds exported annually (Parker 1999). Description. E. p. perreini Vieillot: Zaı¨re, Angola, Zambia, ˆ i, intergrading with next race Tanzania and extreme N Malaw ˆ i. ADULT Y: forehead to hindneck, mantle, north of L. Malaw scapulars and upper back bluish grey; lower back, rump and uppertail-coverts red. Tail black, longish and graduated. Black streak through lores linked narrowly below eye to short black post-ocular streak. Cheeks, ear-coverts, chin and throat light grey, merging with bluish grey sides of neck and grading through grey breast and upper flanks to blackish grey lower flanks, lower belly, thighs and undertail-coverts. Flight feathers grey-brown, edges of primaries and outer webs of secondaries grey; rest of upperwing grey. Underwing-coverts and axillaries light grey. Bill shiny blue-grey with tip and cutting edges black; eyes red; legs black or greenish black. ADULT X: like ad. Y, but grey lower underparts typically less blackish. SIZE (10 YY, 10 XX): wing, Y 50–54 (515),

Estrilda perreini

? ? ?

X 47–53 (504); tail, Y 44–50 (459), X 42–48 (456); bill, Y 11–12 (116), X 11–125 (116); tarsus, Y 15–17 (157), X 145–165 (156). IMMATURE: juvenile slightly paler than ad. and lacks black eyestreak; crimson rump duller. NESTLING: hatchling flesh coloured with a little down on head and back; skin turns grey within 3–4 days. E. p. incana Sundevall (includes ‘poliogastra’ and ‘torrida’): S ˆ i, SE Zimbabwe, Mozambique and South Africa. Lighter Malaw grey below; red rump brighter, more scarlet. Slightly smaller, especially in Mozambique. Wing, Y (n ¼ 10) 47–52 (492). WEIGHT: E Zimbabwe, 2 YY 72, 76, 1 X 75, unsexed (n ¼ 3) 61–90 (74).

Field Characters. Length 105–11 cm. Similar to Cinderella Waxbill E. thomensis, which it meets in Angola, with red rump and uppertail-coverts and black line through eye, but darker grey, and has tiny black chin (hard to see in the field). No red on lower flanks, and black, graduated tail is longer; bill grey-blue, tip darker (no red at base). Immature duller, with paler bluish bill and vestigial eyeline. Yellowbellied Waxbill Cocccopygia quartinia and X Swee C. melanotis have grey head and breast but green back, yellow belly and red lower mandible.

Estrilda perreini BREEDING SUCCESS/SURVIVAL: 5 nests in one year and 3 in next all either destroyed by heavy rains or abandoned (Moore 1983).

289

Key References Brickell, N. and Konigkramer, T. (1997), Clement, P. et al. (1993), Goodwin, D. (1982), Hinze, I. (2000), Immelmann, K. et al. (1964), Kunkel, P. (1967), Moore, A. (1983).

Estrilda perreini (Vieillot). Grey Waxbill. Astrild a ` queue noire.

Plate 18

Fringilla perreini Vieillot, 1817. Nouv. Dict. Hist. Nat., nouv. ´ed., 12, p. 179; Malimbe, Portuguese Congo.

(Opp. p. 267)

Forms a superspecies with E. caerulescens and E. thomensis. Range and Status. Endemic resident, Gabon, only in Tchibanga area, where rare. Congo, only near Djambala and along lower Congo River. Zaı¨re, frequent around Boma near mouth of Congo R., extending upstream to Brazzaville, Kinshasa and Kwamouth; not uncommon in Katanga, north to Baraka; records from between Fizi and Mulala, NW shores of L. Tanganyika. Tanzania, near Kigoma; S Ufipa Plateau; Iringa Highlands, Marengo Highlands, near Jumbe, twice in Mikumi Nat. Park. Angola, Cabinda, Uı´ge, S Cunza Norte, Malanje, N Bie´, Cuanza Sul, Huambo and along escarpment to NW Huı´la; also in SW Lunda Sul, N Lunda Norte and N Moxico; probably much more widespread than mapped. Zambia, uncommon and local; Northern Prov. from near Muyombe and Old Fife, along Muchinga Escarpment from Muzyatama and Kolala to Danger Hill; also Lavushi Manda and north to Kasama and Mbala; in 26% of 300 ˆ i, only Zambia squares (R.J. Dowsett, pers. comm.). Malaw at Chiromo and Mlolo. Zimbabwe, very local, but can be common; up to 800 m in Honde and Nyamkwarara valleys, uncommon in Muneni valley, abundant up to 700 m in Lusitu-Haroni area; Buzi R. in Chipinga highlands, possibly formerly in Mount Selinda area. Mozambique, very local in Zambezia and Manica e Sofala; uncommon along Sul do Save coastal lowlands. Transvaal, rare and local: Woodbush, Haenertsburg and Tzaneen areas and near lower Sabie. KwaZulu-Natal, littoral plain, extending inland in N to Swaziland (where occurs marginally); recent range extension across Umtanvuna R., Transkei. Perhaps >5000 birds in Sul do Save, Mozambique, where population thought to have declined as result of removal of natural vegetation along coast and of persecution for the cage bird trade, for which c. 800 birds exported annually (Parker 1999). Description. E. p. perreini Vieillot: Zaı¨re, Angola, Zambia, ˆ i, intergrading with next race Tanzania and extreme N Malaw ˆ i. ADULT Y: forehead to hindneck, mantle, north of L. Malaw scapulars and upper back bluish grey; lower back, rump and uppertail-coverts red. Tail black, longish and graduated. Black streak through lores linked narrowly below eye to short black post-ocular streak. Cheeks, ear-coverts, chin and throat light grey, merging with bluish grey sides of neck and grading through grey breast and upper flanks to blackish grey lower flanks, lower belly, thighs and undertail-coverts. Flight feathers grey-brown, edges of primaries and outer webs of secondaries grey; rest of upperwing grey. Underwing-coverts and axillaries light grey. Bill shiny blue-grey with tip and cutting edges black; eyes red; legs black or greenish black. ADULT X: like ad. Y, but grey lower underparts typically less blackish. SIZE (10 YY, 10 XX): wing, Y 50–54 (515),

Estrilda perreini

? ? ?

X 47–53 (504); tail, Y 44–50 (459), X 42–48 (456); bill, Y 11–12 (116), X 11–125 (116); tarsus, Y 15–17 (157), X 145–165 (156). IMMATURE: juvenile slightly paler than ad. and lacks black eyestreak; crimson rump duller. NESTLING: hatchling flesh coloured with a little down on head and back; skin turns grey within 3–4 days. E. p. incana Sundevall (includes ‘poliogastra’ and ‘torrida’): S ˆ i, SE Zimbabwe, Mozambique and South Africa. Lighter Malaw grey below; red rump brighter, more scarlet. Slightly smaller, especially in Mozambique. Wing, Y (n ¼ 10) 47–52 (492). WEIGHT: E Zimbabwe, 2 YY 72, 76, 1 X 75, unsexed (n ¼ 3) 61–90 (74).

Field Characters. Length 105–11 cm. Similar to Cinderella Waxbill E. thomensis, which it meets in Angola, with red rump and uppertail-coverts and black line through eye, but darker grey, and has tiny black chin (hard to see in the field). No red on lower flanks, and black, graduated tail is longer; bill grey-blue, tip darker (no red at base). Immature duller, with paler bluish bill and vestigial eyeline. Yellowbellied Waxbill Cocccopygia quartinia and X Swee C. melanotis have grey head and breast but green back, yellow belly and red lower mandible.

290

ESTRILDIDAE

Voice. Tape-recorded (99, 104, F, GIB, GUT). Call high and thin, a long ‘seeeee’, often dropping at end, ‘seeeeeup’. In mild alarm, Y has plaintive, drawn out whistle, ‘fweeeee’, sometimes followed by up to 5 shorter notes; long whistle followed by short one given during side-toside movements that precede courtship display (Goodwin 1982). Apparent ‘song’, a variety of repeated long whistles followed by more abrupt notes (Clement et al. 1993). All vocalizations said to be distinct from those of Cinderella and Lavender Waxbills E. caerulescens (Gu ¨ ttinger and Nicolai 1973). General Habits. Inhabits open woodland with scattered large trees; shrubs and small trees with tufty, patchy grass between (S Zaı¨re); edges of primary and secondary forest and gallery forest (Angola); edges of lowland evergreen and riparian forests, thickets and secondary growth (Zimbabwe), grass and low herb growth on banks of streams and in shade of trees, e.g. plantation of rubber trees (W Zaı¨re); coastal woodlands, littoral plain forest edges and wooded valleys (Sul do Save, Mozambique, and KwaZulu-Natal, South Africa). Singly or in pairs; readily overlooked. Forages on ground, close to or underneath dense cover; also feeds at grass heads. Holds seeding grass head on ground under foot and extracts seeds with bill. Perches in trees, not only when nesting there, and said to feed sometimes in tree canopy. Sedentary. Food. Only small grass seeds in stomachs (Zaı¨re). Insects and nectar (Maclean 1993). In captivity, seeds of Panicum maximum, Casuarina equisetifolia, seeds in fruit of Sclerocarya caffra, and small insects, particularly termites. Breeding Habits. Solitary nester; monogamous. Courtship display (in captivity): Y flies conspicuously around aviary carrying long grass stem (c. 40 long) by one end in bill; he alights, adopts horizontal posture and swings body from side to side, sometimes whistling. Copulation does not follow this display but is initiated by X which, whilst pair is allopreening, suddenly crouches and quivers tail.

Greeting display: pair, with foreheads sleeked making heads look triangular, angled tails and slightly fluffed-out flanks, bow towards each other and synchronously wipe bills on perch; in captivity, pair ‘always displayed to each other at first sight when the light in their room came on in the morning’ (E. Po ¨hland in Goodwin 1982). NEST: retort-shaped or oval, with tubular entrance c. 45– 75 long, projecting slightly downward from nest top; robustly but roughly constructed with long, fine grass stems; decorated on top; unlined or with slight pad of soft, downy grass heads (Vincent 1949), or lined with fine grass and feathers (Maclean 1993). Ext. diam. c. 100  125. Placed in fork 15–5 (usually 2–4) m above ground in thick-foliaged evergreen bush or tree (once Acacia karoo) or bamboo clump, in open woodland often near stream; concealed or in full view. Occasionally relines and uses old nest of Dark-backed Weaver Ploceus bicolor or (in captivity) estrildids Uraeginthus cyanocephala and Amandava formosa. Nest built by both sexes, Y collecting material and X arranging lining (Brickell and Konigkramer 1997). EGGS: (n ¼ 15) 2–5, usually 3–4 in tropics and 4–5 in southern Africa where av. 43; up to 7 and 8 in captivity, with unlimited food supply. Pure white. SIZE: (n ¼ 10, Zaı¨re) 141–152  109–115 (147  111), (n ¼ 30, South Africa) 137–154  105–115 (146  114). LAYING DATES: Gabon, Apr; Zaı ¨re, Boma, (near to laying, ˆ i, Feb, Jan), Katanga, Apr; Zambia, Dec–Feb, Apr; Malaw Apr; Zimbabwe, Mar–Apr; Transvaal, Feb; KwaZuluNatal, Oct–Feb. INCUBATION: by both sexes. Period: in captivity, 12 days. DEVELOPMENT AND CARE OF YOUNG: young cared for by both parents. Brooding diminishes after 5 days and stops after 7 days. When begging, young do not adopt posture characterizing most congeners but stretch heads straight up (Immelmann et al. 1964). Period: in captivity, 19–21 days. Young return to roost in nest at night for c. 7 days. Key References Brickell, N. and Konigkramer, T. (1997), Goodwin, D. (1982), Immelmann et al. (1964), Po ¨hland, E. (1967, 1969), Vincent, A. W. (1949).

Plate 18

Estrilda thomensis Sousa. Cinderella Waxbill. Astrild cendrillon.

(Opp. p. 267)

Estrelda thomensis Sousa, 1888. Jorn. Sci. Math. Phys. Nat Lisboa, 12, p. 44; Deep Sloot, Benguella, Angola. Forms a superspecies with E. perreini and E. caerulescens. Range and Status. Endemic resident. Angola, locally common from SW Namibe north along escarpment to Deep Sloot in Benguela and Fazenda de Cuito in Huambo (12
270 S, 15
160 E), and W Huı´la east to Gambos (15
450 S, 14
050 E) (Dean 2000). Old record in S Bengo. Extreme NW Namibia along lower Cunene R. from Ruacana to about Baynes Mts, and south to Ehomba, Ondoto R. near Epembe, W side of Zebra Mts, and just north of Okongwati along Omuhongo River. The type specimen is said to have come from Sa˜o Tome´, but the bird has not been seen there

since, and probably never was native (de Naurois 1994); it may have been an imported bird. Classed as Near-Threatened by Collar et al. (1994), and stated to be included in Namibian Red Data Book as ‘Vulnerable’; however, the severe threat said by Harrison et al. (1997) to be posed by planned hydroelectric development on Cunene R. at Epupa Falls through destruction of riverine woodland is an overstatement. The bird is mainly associated with mopane, not riverine habitats; a much greater threat is the increase in human habitation at springs

290

ESTRILDIDAE

Voice. Tape-recorded (99, 104, F, GIB, GUT). Call high and thin, a long ‘seeeee’, often dropping at end, ‘seeeeeup’. In mild alarm, Y has plaintive, drawn out whistle, ‘fweeeee’, sometimes followed by up to 5 shorter notes; long whistle followed by short one given during side-toside movements that precede courtship display (Goodwin 1982). Apparent ‘song’, a variety of repeated long whistles followed by more abrupt notes (Clement et al. 1993). All vocalizations said to be distinct from those of Cinderella and Lavender Waxbills E. caerulescens (Gu ¨ ttinger and Nicolai 1973). General Habits. Inhabits open woodland with scattered large trees; shrubs and small trees with tufty, patchy grass between (S Zaı¨re); edges of primary and secondary forest and gallery forest (Angola); edges of lowland evergreen and riparian forests, thickets and secondary growth (Zimbabwe), grass and low herb growth on banks of streams and in shade of trees, e.g. plantation of rubber trees (W Zaı¨re); coastal woodlands, littoral plain forest edges and wooded valleys (Sul do Save, Mozambique, and KwaZulu-Natal, South Africa). Singly or in pairs; readily overlooked. Forages on ground, close to or underneath dense cover; also feeds at grass heads. Holds seeding grass head on ground under foot and extracts seeds with bill. Perches in trees, not only when nesting there, and said to feed sometimes in tree canopy. Sedentary. Food. Only small grass seeds in stomachs (Zaı¨re). Insects and nectar (Maclean 1993). In captivity, seeds of Panicum maximum, Casuarina equisetifolia, seeds in fruit of Sclerocarya caffra, and small insects, particularly termites. Breeding Habits. Solitary nester; monogamous. Courtship display (in captivity): Y flies conspicuously around aviary carrying long grass stem (c. 40 long) by one end in bill; he alights, adopts horizontal posture and swings body from side to side, sometimes whistling. Copulation does not follow this display but is initiated by X which, whilst pair is allopreening, suddenly crouches and quivers tail.

Greeting display: pair, with foreheads sleeked making heads look triangular, angled tails and slightly fluffed-out flanks, bow towards each other and synchronously wipe bills on perch; in captivity, pair ‘always displayed to each other at first sight when the light in their room came on in the morning’ (E. Po ¨hland in Goodwin 1982). NEST: retort-shaped or oval, with tubular entrance c. 45– 75 long, projecting slightly downward from nest top; robustly but roughly constructed with long, fine grass stems; decorated on top; unlined or with slight pad of soft, downy grass heads (Vincent 1949), or lined with fine grass and feathers (Maclean 1993). Ext. diam. c. 100  125. Placed in fork 15–5 (usually 2–4) m above ground in thick-foliaged evergreen bush or tree (once Acacia karoo) or bamboo clump, in open woodland often near stream; concealed or in full view. Occasionally relines and uses old nest of Dark-backed Weaver Ploceus bicolor or (in captivity) estrildids Uraeginthus cyanocephala and Amandava formosa. Nest built by both sexes, Y collecting material and X arranging lining (Brickell and Konigkramer 1997). EGGS: (n ¼ 15) 2–5, usually 3–4 in tropics and 4–5 in southern Africa where av. 43; up to 7 and 8 in captivity, with unlimited food supply. Pure white. SIZE: (n ¼ 10, Zaı¨re) 141–152  109–115 (147  111), (n ¼ 30, South Africa) 137–154  105–115 (146  114). LAYING DATES: Gabon, Apr; Zaı ¨re, Boma, (near to laying, ˆ i, Feb, Jan), Katanga, Apr; Zambia, Dec–Feb, Apr; Malaw Apr; Zimbabwe, Mar–Apr; Transvaal, Feb; KwaZuluNatal, Oct–Feb. INCUBATION: by both sexes. Period: in captivity, 12 days. DEVELOPMENT AND CARE OF YOUNG: young cared for by both parents. Brooding diminishes after 5 days and stops after 7 days. When begging, young do not adopt posture characterizing most congeners but stretch heads straight up (Immelmann et al. 1964). Period: in captivity, 19–21 days. Young return to roost in nest at night for c. 7 days. Key References Brickell, N. and Konigkramer, T. (1997), Goodwin, D. (1982), Immelmann et al. (1964), Po ¨hland, E. (1967, 1969), Vincent, A. W. (1949).

Plate 18

Estrilda thomensis Sousa. Cinderella Waxbill. Astrild cendrillon.

(Opp. p. 267)

Estrelda thomensis Sousa, 1888. Jorn. Sci. Math. Phys. Nat Lisboa, 12, p. 44; Deep Sloot, Benguella, Angola. Forms a superspecies with E. perreini and E. caerulescens. Range and Status. Endemic resident. Angola, locally common from SW Namibe north along escarpment to Deep Sloot in Benguela and Fazenda de Cuito in Huambo (12
270 S, 15
160 E), and W Huı´la east to Gambos (15
450 S, 14
050 E) (Dean 2000). Old record in S Bengo. Extreme NW Namibia along lower Cunene R. from Ruacana to about Baynes Mts, and south to Ehomba, Ondoto R. near Epembe, W side of Zebra Mts, and just north of Okongwati along Omuhongo River. The type specimen is said to have come from Sa˜o Tome´, but the bird has not been seen there

since, and probably never was native (de Naurois 1994); it may have been an imported bird. Classed as Near-Threatened by Collar et al. (1994), and stated to be included in Namibian Red Data Book as ‘Vulnerable’; however, the severe threat said by Harrison et al. (1997) to be posed by planned hydroelectric development on Cunene R. at Epupa Falls through destruction of riverine woodland is an overstatement. The bird is mainly associated with mopane, not riverine habitats; a much greater threat is the increase in human habitation at springs

Estrilda thomensis

Estrilda thomensis

grey, almost whitish on face and throat, with rosy tinge on back and breast, red lower flanks, pink and black bill. Immature with no red on flanks extremely similar to Grey Waxbill but paler, with pinkish white bill. Voice. Tape-recorded (99, 104, GUT). Song of Y, lasting c. 3 s, 3–4 high, thin, pure notes, first 1–2 short, others more drawn out, ‘seee-seee-seeeeeyo-syooooo’; last note slightly lower in pitch and with a sweet quality. Calls between songs, thin ‘tsit’ and hard, dry ‘brrt’. Song of X said to be single ‘seee’ (Goodwin 1982). Contact call during flight and when moving rapidly through vegetation while foraging, harsh ‘bzzt, bzzt’; more intense version used as alarm, rapidly repeated when birds are disturbed. Contact call used by perched birds when sitting together or preening after bathing or drinking, a soft, melodic ‘swee-swee-swee’, fairly drawn out. Also described are contact and locomotion intention calls, soft and variable with an ‘r’ quality, e.g. ‘tree’ or ‘kr’, uttered almost constantly when bird is moving about, lengthened and intensified when it is excited (Goodwin 1982).

and small tributaries south of Cunene R., with associated wood cutting, overgrazing and over-utilization of water. Description. ADULT Y: forehead to hindneck, mantle, scapulars and upper back pale grey; mantle and scapulars may show faint pinkish wash (not obvious in the field); lower back, rump and uppertail-coverts wine-red to crimson; tail black. Broad black loral area and narrow black post-ocular streak. Cheeks, chin and throat greyish white, becoming greyer on sides of neck, black spot on chin at base of lower mandible; breast and upper flanks grey (breast with pronounced pink wash in some birds), grading to blackish on centre of lower belly, thighs and undertail-coverts; some birds have distinct pale or red margin to tips of belly and vent feathers, which appear indistinctly barred in the field; lower flanks wine-red to crimson, red variable in extent. Flight feathers grey-brown, edges of primaries and outer webs of secondaries grey, rest of upper wing grey. Underwing-coverts and axillaries greyish white. Bill mauve to pinkish red with tip, culmen, cutting edges and underside of lower mandible blackish; eyes dark brown, thin orbital ring dull blackish slate; legs brownish black. ADULT X: similar to ad. Y but lower belly and undertail-coverts dark grey (less blackish), pink wash on breast and mantle fainter. SIZE: wing, YX (n ¼ 22) 485–53 (50), tail, YX (n ¼ 18) 48–525 (49); bill, YX (n ¼ 22) 8–91 (83); tarsus, YX (n ¼ 19) 131–149 (139). WEIGHT: Y (n ¼ 19) 6–8, X (n ¼ 10) 6–8, YX (n ¼ 22) 68–85 (76). IMMATURE: similar to ad. but lacks black eye-streak and pink tinge on breast and mantle; lower back and uppertail-coverts grey with red tips to feathers giving mottled appearance, cheeks and chin darker, not contrasting with mantle, flanks mottled wine-red and grey, not solid red, lower belly, thighs and undertail-coverts paler grey, not contrasting strongly with lower breast, upperwingcoverts brownish grey. Bill darker and duller, dull pink with some dark brown on lower mandible, cutting edges and tip. NESTLING: not described.

Field Characters. Length 11 cm. A grey waxbill with black line through lores to just behind eye, red rump and uppertail-coverts and black tail. Very similar to Grey Waxbill E. perreini, which it meets in Angola, but paler

General Habits. In Angola inhabits riverine vegetation in acacia and mopane woodland; in Namibia restricted to mopane and riverine woodlands of Cunene R. system and its drainages, rare in acacia. Definitely water-dependent, frequenting springs, hand-dug wells and shallow water scrapes in sandy river beds made by animals. Occurs singly, in pairs or small groups of up to 8, sometimes up to 20 or 25. Before and after visiting water, birds gather in same patch of vegetation, usually one not used by other species. Much socializing, preening and calling takes place at this site, and birds roost there during the heat of the day. First birds arrive at water around 08h00, peak of activity 09h00–11h00; very few birds come after 13h00 when heat most intense; small numbers come in late afternoon, about an hour before sunset. Roosts in old nests of its own or other species, e.g. White-browed Sparrow-Weaver Plocepasser mahali (9 birds seen entering nest at dusk). In captivity less aggressive and inquisitive than Grey Waxbill and much less so than Lavender Waxbill E. caerulescens (Goodwin 1982). Forages systematically through bush or tree canopy; picks grass seeds from flowering heads by climbing up and down stems or by hanging from twigs and branches of woody plants; not seen feeding on ground (pers. obs.), though said to do so by Ginn et al. (1989). Catches emerging termite alates on the wing. Apparently resident in Angola (Dean 2000), but subject to considerable local movements in Namibia after rain (in wet season absent or difficult to find in areas where known to be common). Food. Grass seeds and other small seeds; stamens of flowers of Euclea divinorum (and possibly insects thereon); feeds on flowers of Gymnosporia buxiifolia, either eating parts of plant or taking nectar; insects, including termite alates and honey-producing scale insects on mopane trees. In captivity chiefly dry millet, also soaked and sprouted seeds, sprays of half-ripe wild millet, seeding rye grass, honey water and pupae of ants, especially Lasius (Goodwin

291

292

ESTRILDIDAE

1982); seeds of Lolium perenne, Panicum laevifolium, P. maximum, P. coloratum, P. lanipes and Rhynchelytrum repens; spiders, termites, aphids and mealworms (Brickell and Konigkramer 1997). Breeding Habits. Nest-building in Namibia in late Nov– Dec, after first rains. Nothing further recorded in the wild. NEST: (in captivity) large ball of grass and coconut fibres, with narrow down-sloping entrance tube 120 long; sides and top 20–30 thick, floor 40–50 thick. A Y built ‘cock

nest’ on top of breeding nest and roosted in it regularly (Po ¨hland 1970). EGGS: (2 clutches) 3–4. White. INCUBATION: Period 125–14 days. DEVELOPMENT AND CARE OF YOUNG: young fed by both parents; start feeding themselves 2 days after fledging; no longer fed by parents after 10th day. Nestling period 17–21 days. Key Reference Goodwin, D. (1982).

Plate 18

Estrilda paludicola Heuglin. Fawn-breasted Waxbill. Astrild a` poitrine fauve.

(Opp. p. 267)

Estrilda paludicola Heuglin, 1863. J. Orn., 11, p. 166; middle course of Bahr el Ghazal. Forms a superspecies with E. poliopareia. Range and Status. Endemic resident, range fragmented, from Ethiopia to Angola and Zambia. Central African Republic, Manovo-Gounda-Saint Floris Nat. Park (Carroll 1988). Sudan, frequent in Bahr al Ghazal, El Buheyrat, Western Equatoria and W Eastern Equatoria; common around Boma; a few records of ochrogaster near Ethiopian border. W Ethiopia, uncommon to locally abundant, along streams at 1200–2100 m. Zaı¨re, paludicola the commonest waxbill in Uele, from Buta to L. Albert, common in E Ituri, south to L. Edward (paludicola), and in Kivu at 1830–1950 m, south to about Mwenga (roseicrissa); frequent in Katanga, east to Marungu Highlands (benguellensis); frequent along lower R. Congo from Lukolela to Bolobo, also away from river at Kunungu (ruthae). Gabon, frequent around Le´koni. Congo, Odzala Nat. Park (common), Le´fini Res. and Mpe´ area; and Mayombe Forest (Djambala, Mossaka). Gabon and Congo birds, and those southeast of Impfondo (Oubangui R., N Congo/Zaı¨re border) are probably all referable to ruthae. Uganda, locally common, roseicrissa in Toro, Ankole and Kigezi, paludicola widespread from Sango Bay to Kamchuru and Kidepo Valley Nat. Park. Kenya, uncommon and local, from Mt Elgon and Saiwa Nat. Park to Busia, Mumias and Kakamega Districts, south to Kisii, Kilgoris and NW Mara Game Reserve. Tanzania, roseicrissa in West Lake and Kibondo in NW and Shinyanga region south of L. Victoria; benguellensis in extreme SW at Tatanda; sight records in Katavi Nat. Park not certainly of this species; and marwitzi widespread in Iringa and Njombe Highlands. Rwanda, known in N, e.g. Gabiro; frequent (?) in NE. Burundi, scarce, in NE. Angola, common: Bengo, Uı´ge, Cuanza Norte, Cuanza Sul, Malanje, Lunda Norte, Lunda Sul, NE Moxico, Bie´, Huambo and N Huı´la Provs; probably more widespread than mapped and throughout Moxico. Zambia, as mapped, in 40% of 300 squares, S and E borders of range rather precisely delineated (Benson et al. 1971, R. J. Dowsett, pers. comm.), south to extreme N of Kafue Nat. Park (Ndulumina) and central Kabwe Rural, east to Muchinga Escarpment and SE Isoka. Description. E. p. paludicola Heuglin: S Sudan, NE Zaı¨re (Uele), Uganda, W Kenya. ADULT Y: forehead to nape brownish grey,

Estrilda paludicola

?

?

?

?

merging with olive-brown hindneck, sides of neck, mantle, scapulars and back; mantle feathers with faint darker barring; rump and uppertail-coverts crimson. Tail black, rather long and well graduated. Lores, cheeks and ear-coverts pale grey. Chin and throat buffy white, merging with pale yellowish buff breast and flanks, suffused olive-grey at sides; centre of belly brighter yellowish buff, some feathers tipped pink or red; undertailcoverts buffy white. Upperwing olive-brown; underwing coverts and axillaries creamy buff. Bill bright red to orange-red; eyes red or reddish brown; legs dark brown. ADULT X: like ad. Y, but underparts slightly paler, more grey-brown at sides, with little pink on belly feathers. SIZE (10 YY, 10 XX): wing, Y 46–52 (480), X 44–49 (468); tail, Y 42–47 (446), X 40–45 (428); bill, Y 9–105 (98), X 9–105 (99); tarsus, Y 14–155 (149), X 135–15 (142). WEIGHT: Kenya, Y (n ¼ 3) 67–74 (72), X (n ¼ 3) 74–79 (76); unsexed (n ¼ 26) 66–85 (76). IMMATURE: juvenile like ad. but paler below; bill black. NESTLING: naked at first; mouth pattern very like that of E. astrild (Goodwin 1982).

292

ESTRILDIDAE

1982); seeds of Lolium perenne, Panicum laevifolium, P. maximum, P. coloratum, P. lanipes and Rhynchelytrum repens; spiders, termites, aphids and mealworms (Brickell and Konigkramer 1997). Breeding Habits. Nest-building in Namibia in late Nov– Dec, after first rains. Nothing further recorded in the wild. NEST: (in captivity) large ball of grass and coconut fibres, with narrow down-sloping entrance tube 120 long; sides and top 20–30 thick, floor 40–50 thick. A Y built ‘cock

nest’ on top of breeding nest and roosted in it regularly (Po ¨hland 1970). EGGS: (2 clutches) 3–4. White. INCUBATION: Period 125–14 days. DEVELOPMENT AND CARE OF YOUNG: young fed by both parents; start feeding themselves 2 days after fledging; no longer fed by parents after 10th day. Nestling period 17–21 days. Key Reference Goodwin, D. (1982).

Plate 18

Estrilda paludicola Heuglin. Fawn-breasted Waxbill. Astrild a` poitrine fauve.

(Opp. p. 267)

Estrilda paludicola Heuglin, 1863. J. Orn., 11, p. 166; middle course of Bahr el Ghazal. Forms a superspecies with E. poliopareia. Range and Status. Endemic resident, range fragmented, from Ethiopia to Angola and Zambia. Central African Republic, Manovo-Gounda-Saint Floris Nat. Park (Carroll 1988). Sudan, frequent in Bahr al Ghazal, El Buheyrat, Western Equatoria and W Eastern Equatoria; common around Boma; a few records of ochrogaster near Ethiopian border. W Ethiopia, uncommon to locally abundant, along streams at 1200–2100 m. Zaı¨re, paludicola the commonest waxbill in Uele, from Buta to L. Albert, common in E Ituri, south to L. Edward (paludicola), and in Kivu at 1830–1950 m, south to about Mwenga (roseicrissa); frequent in Katanga, east to Marungu Highlands (benguellensis); frequent along lower R. Congo from Lukolela to Bolobo, also away from river at Kunungu (ruthae). Gabon, frequent around Le´koni. Congo, Odzala Nat. Park (common), Le´fini Res. and Mpe´ area; and Mayombe Forest (Djambala, Mossaka). Gabon and Congo birds, and those southeast of Impfondo (Oubangui R., N Congo/Zaı¨re border) are probably all referable to ruthae. Uganda, locally common, roseicrissa in Toro, Ankole and Kigezi, paludicola widespread from Sango Bay to Kamchuru and Kidepo Valley Nat. Park. Kenya, uncommon and local, from Mt Elgon and Saiwa Nat. Park to Busia, Mumias and Kakamega Districts, south to Kisii, Kilgoris and NW Mara Game Reserve. Tanzania, roseicrissa in West Lake and Kibondo in NW and Shinyanga region south of L. Victoria; benguellensis in extreme SW at Tatanda; sight records in Katavi Nat. Park not certainly of this species; and marwitzi widespread in Iringa and Njombe Highlands. Rwanda, known in N, e.g. Gabiro; frequent (?) in NE. Burundi, scarce, in NE. Angola, common: Bengo, Uı´ge, Cuanza Norte, Cuanza Sul, Malanje, Lunda Norte, Lunda Sul, NE Moxico, Bie´, Huambo and N Huı´la Provs; probably more widespread than mapped and throughout Moxico. Zambia, as mapped, in 40% of 300 squares, S and E borders of range rather precisely delineated (Benson et al. 1971, R. J. Dowsett, pers. comm.), south to extreme N of Kafue Nat. Park (Ndulumina) and central Kabwe Rural, east to Muchinga Escarpment and SE Isoka. Description. E. p. paludicola Heuglin: S Sudan, NE Zaı¨re (Uele), Uganda, W Kenya. ADULT Y: forehead to nape brownish grey,

Estrilda paludicola

?

?

?

?

merging with olive-brown hindneck, sides of neck, mantle, scapulars and back; mantle feathers with faint darker barring; rump and uppertail-coverts crimson. Tail black, rather long and well graduated. Lores, cheeks and ear-coverts pale grey. Chin and throat buffy white, merging with pale yellowish buff breast and flanks, suffused olive-grey at sides; centre of belly brighter yellowish buff, some feathers tipped pink or red; undertailcoverts buffy white. Upperwing olive-brown; underwing coverts and axillaries creamy buff. Bill bright red to orange-red; eyes red or reddish brown; legs dark brown. ADULT X: like ad. Y, but underparts slightly paler, more grey-brown at sides, with little pink on belly feathers. SIZE (10 YY, 10 XX): wing, Y 46–52 (480), X 44–49 (468); tail, Y 42–47 (446), X 40–45 (428); bill, Y 9–105 (98), X 9–105 (99); tarsus, Y 14–155 (149), X 135–15 (142). WEIGHT: Kenya, Y (n ¼ 3) 67–74 (72), X (n ¼ 3) 74–79 (76); unsexed (n ¼ 26) 66–85 (76). IMMATURE: juvenile like ad. but paler below; bill black. NESTLING: naked at first; mouth pattern very like that of E. astrild (Goodwin 1982).

Estrilda poliopareia E. p. ochrogaster Salvadori: Ethiopia, extreme E Sudan. Differs distinctively from nominate paludicola in having paler upperparts, more yellowish olive, top of head only slightly greyer than mantle; lores and cheeks tinged yellow, and chin to breast brighter yellowish buff, concolorous with belly; usually some pink on belly feathers. E. p. ruthae Chapin: Congo and adjacently in Gabon and W Zaı¨re. Similar to nominate paludicola but paler above, and with whole underparts whiter. E. p. roseicrissa Reichenow: E Zaı¨re, Rwanda, Burundi, SW Uganda, NW Tanzania. Differs from paludicola in having top of head rather pale olive-brown, uniform with mantle; buff and whitish below, with no yellow tinge; Y has rose-pink central belly patch (a pink tinge only in X). E. p. marwitzi Reichenow: SW Tanzania. Like roseicrissa, but darker, more tawny-olive above; greyer (less olive) on flanks; darker crimson on rump and uppertail-coverts. E. p. benguellensis Neumann: Angola, S Zaı¨re (Kasai, Katanga), N Zambia. Grey crown better demarcated than in nominate race and upperparts richer olive; uppertail-coverts brighter red with bases of tail feathers also edged crimson; underparts brighter buff and breast of Y often tinged pink; belly patch yellower, feathers more extensively tipped red in both sexes. TAXONOMIC NOTE: 6 subspecies in 5 isolates: only paludicola and roseicrissa are parapatric. Most are distinctive in appearance, ochrogaster yellowish, paludicola and benguellensis grey-headed, benguellensis and roseicrissa pink on flanks, roseicrissa and marwitzi grey-faced and brown-capped, and each of the 5 isolates is a candidate for upgrading to species status. Ethiopian ochrogaster was treated as an allospecies by Wolters (1979–1982) and Sibley and Monroe (1990) but as a subspecies by M. A. Traylor in Paynter (1968), Hall and Moreau (1970), White (1963), Goodwin (1982), Nikolaus (1987, 1989), Clement et al. (1993) and Dowsett and Dowsett-Lemaire (1993b). We prefer the latter course; its differences from other populations of E. paludicola are slighter than those between e.g. Coccopygia melanotis and bocagei (q.v.) which we treat as conspecific, and much slighter than those between E. paludicola and E. poliopareia.

Field Characters. Length 9–10 cm. A waxbill of damp places, longer-tailed than Swee Waxbill Coccopygia quartinia, with rich brown back and wings and all-red bill. Plumage varies with race: face grey, top of head grey to brown; throat creamy white, ventral patch pale pink, breast fawn-coloured to pale grey; underparts of Ethiopian race ochrogaster rich golden buff. Immature has black bill and whiter underparts with no pink on belly. Orangecheeked Waxbill E. melpoda is similar but with orange face. Voice. Tape-recorded (104, B, C, MOY). Chattering, chirping contact calls from group rather nasal, ‘jup’, ‘jip’, ‘jeep’, ‘jee-jee-jeep’, ‘jop-wick’. Alarm, ‘tsyee-krr’ or ‘kr-yee-eh’; song a hard ‘tek tek tek teketree teketree’ and variants (Goodwin 1982).

293

General Habits. Inhabits open damp grass areas within mature but thin woodland where shrubs interspersed with clumps of tall or tufty knee-high grass; savanna woodland with Terminalia laxiflora trees (Central African Republic), vegetation along streams between 1200 and 2100 m (Ethiopia), and grassy, overgrown cultivation at 1000– 2000 m (Uganda); rank grass and reeds near lowland swamps, marshes, rivers and streams, down to c. 400 m; dambos and inundated grassland; environs of villages. Sometimes occurs in dry grassland (Congo). Usually in flocks, often large (Weeks 1948, Chapin 1954), i.e. of 30 or more. Clings to seeding heads of tall grasses to feed; also forages on ground, sometimes in company of other waxbills e.g. Common E. astrild and Orange-cheeked E. melpoda (Zaı¨re). In captivity lively, not nervous or shy; feeding methods, gait, movements and flight like those of E. astrild and E. melpoda (Goodwin 1982). Food. Small grass seeds; some insects. In captivity, eats seeds of millet, Panicum maximum and P. laevifolium, and ant pupae, termites, wasp larvae and gnats. Breeding Habits. Solitary nester. NEST: spherical, made quite strongly of fine grass heads and fine, stiff grass, with funnel entrance at side, sloping down, 25 mm long. No lining. Well concealed, on ground or 10 cm above it at base of clump of rough grass 45 cm tall; in captivity, in bracken fern Pteridium aquilinum and ginger Aframomum sp. (Brickell and Konigkramer 1997). Nest diam. c. 90 mm. On top of 2 of the 3 nests described, a ‘cock’s-nest’ constructed, same size as nest proper but frail; both nests proper contained clutches (Sudan, Weeks 1948; S Zaı¨re, Vincent 1949). EGGS: 4–5, once 10. White. SIZE: (n ¼ 5) 126–138  97–100. LAYING DATES: Sudan, Boma, July-Aug; Uganda and Kenya, Jan–Mar, May, Oct-Nov. Zaı¨re, Lubumbashi, Dec; in general, breeds in rainiest season. Zambia, Jan–Apr. INCUBATION: period, in captivity (South Africa, n ¼ 4) 12–13 (1275) days. DEVELOPMENT AND CARE OF YOUNG: period, in captivity (South Africa, n ¼ 4) 19–21 (20) days. BREEDINGSUCCESS/SURVIVAL: once parasitized by Vidua sp., probably V. macroura (Uganda: Jackson and Sclater 1938). Key References Brickell, N. (1999), Brickell, N. and Konigkramer, T. (1997), Chapin, J. P. (1954), Clement, P. et al. (1993), Goodwin, D. (1982), Vincent, A. W. (1949), Weeks, J. T. (1948).

Estrilda poliopareia Reichenow. Anambra Waxbill. Astrild du Niger.

Plate 18

Estrilda poliopareia Reichenow, 1902. Orn. Monatsb., 10, p. 185; ‘‘Congo’’, restricted to S Nigeria by Chapin, 1950, Bull. Br. Orn. Club, 70, p. 24. Estrilda anambrae, 1907. J. Orn., 55, p. 624; Anambra Creek, Agoulerie, S Nigeria.

(Opp. p. 267)

Forms a superspecies with E. paludicola. Range and Status. Endemic resident, S Nigeria; known from 3–4 localities: Onitsha area (lower R. Niger),

Forcados (W coast of Niger delta) and Badagri (brackish creek and longshore island west of Lagos). Discovered in

Estrilda poliopareia E. p. ochrogaster Salvadori: Ethiopia, extreme E Sudan. Differs distinctively from nominate paludicola in having paler upperparts, more yellowish olive, top of head only slightly greyer than mantle; lores and cheeks tinged yellow, and chin to breast brighter yellowish buff, concolorous with belly; usually some pink on belly feathers. E. p. ruthae Chapin: Congo and adjacently in Gabon and W Zaı¨re. Similar to nominate paludicola but paler above, and with whole underparts whiter. E. p. roseicrissa Reichenow: E Zaı¨re, Rwanda, Burundi, SW Uganda, NW Tanzania. Differs from paludicola in having top of head rather pale olive-brown, uniform with mantle; buff and whitish below, with no yellow tinge; Y has rose-pink central belly patch (a pink tinge only in X). E. p. marwitzi Reichenow: SW Tanzania. Like roseicrissa, but darker, more tawny-olive above; greyer (less olive) on flanks; darker crimson on rump and uppertail-coverts. E. p. benguellensis Neumann: Angola, S Zaı¨re (Kasai, Katanga), N Zambia. Grey crown better demarcated than in nominate race and upperparts richer olive; uppertail-coverts brighter red with bases of tail feathers also edged crimson; underparts brighter buff and breast of Y often tinged pink; belly patch yellower, feathers more extensively tipped red in both sexes. TAXONOMIC NOTE: 6 subspecies in 5 isolates: only paludicola and roseicrissa are parapatric. Most are distinctive in appearance, ochrogaster yellowish, paludicola and benguellensis grey-headed, benguellensis and roseicrissa pink on flanks, roseicrissa and marwitzi grey-faced and brown-capped, and each of the 5 isolates is a candidate for upgrading to species status. Ethiopian ochrogaster was treated as an allospecies by Wolters (1979–1982) and Sibley and Monroe (1990) but as a subspecies by M. A. Traylor in Paynter (1968), Hall and Moreau (1970), White (1963), Goodwin (1982), Nikolaus (1987, 1989), Clement et al. (1993) and Dowsett and Dowsett-Lemaire (1993b). We prefer the latter course; its differences from other populations of E. paludicola are slighter than those between e.g. Coccopygia melanotis and bocagei (q.v.) which we treat as conspecific, and much slighter than those between E. paludicola and E. poliopareia.

Field Characters. Length 9–10 cm. A waxbill of damp places, longer-tailed than Swee Waxbill Coccopygia quartinia, with rich brown back and wings and all-red bill. Plumage varies with race: face grey, top of head grey to brown; throat creamy white, ventral patch pale pink, breast fawn-coloured to pale grey; underparts of Ethiopian race ochrogaster rich golden buff. Immature has black bill and whiter underparts with no pink on belly. Orangecheeked Waxbill E. melpoda is similar but with orange face. Voice. Tape-recorded (104, B, C, MOY). Chattering, chirping contact calls from group rather nasal, ‘jup’, ‘jip’, ‘jeep’, ‘jee-jee-jeep’, ‘jop-wick’. Alarm, ‘tsyee-krr’ or ‘kr-yee-eh’; song a hard ‘tek tek tek teketree teketree’ and variants (Goodwin 1982).

293

General Habits. Inhabits open damp grass areas within mature but thin woodland where shrubs interspersed with clumps of tall or tufty knee-high grass; savanna woodland with Terminalia laxiflora trees (Central African Republic), vegetation along streams between 1200 and 2100 m (Ethiopia), and grassy, overgrown cultivation at 1000– 2000 m (Uganda); rank grass and reeds near lowland swamps, marshes, rivers and streams, down to c. 400 m; dambos and inundated grassland; environs of villages. Sometimes occurs in dry grassland (Congo). Usually in flocks, often large (Weeks 1948, Chapin 1954), i.e. of 30 or more. Clings to seeding heads of tall grasses to feed; also forages on ground, sometimes in company of other waxbills e.g. Common E. astrild and Orange-cheeked E. melpoda (Zaı¨re). In captivity lively, not nervous or shy; feeding methods, gait, movements and flight like those of E. astrild and E. melpoda (Goodwin 1982). Food. Small grass seeds; some insects. In captivity, eats seeds of millet, Panicum maximum and P. laevifolium, and ant pupae, termites, wasp larvae and gnats. Breeding Habits. Solitary nester. NEST: spherical, made quite strongly of fine grass heads and fine, stiff grass, with funnel entrance at side, sloping down, 25 mm long. No lining. Well concealed, on ground or 10 cm above it at base of clump of rough grass 45 cm tall; in captivity, in bracken fern Pteridium aquilinum and ginger Aframomum sp. (Brickell and Konigkramer 1997). Nest diam. c. 90 mm. On top of 2 of the 3 nests described, a ‘cock’s-nest’ constructed, same size as nest proper but frail; both nests proper contained clutches (Sudan, Weeks 1948; S Zaı¨re, Vincent 1949). EGGS: 4–5, once 10. White. SIZE: (n ¼ 5) 126–138  97–100. LAYING DATES: Sudan, Boma, July-Aug; Uganda and Kenya, Jan–Mar, May, Oct-Nov. Zaı¨re, Lubumbashi, Dec; in general, breeds in rainiest season. Zambia, Jan–Apr. INCUBATION: period, in captivity (South Africa, n ¼ 4) 12–13 (1275) days. DEVELOPMENT AND CARE OF YOUNG: period, in captivity (South Africa, n ¼ 4) 19–21 (20) days. BREEDINGSUCCESS/SURVIVAL: once parasitized by Vidua sp., probably V. macroura (Uganda: Jackson and Sclater 1938). Key References Brickell, N. (1999), Brickell, N. and Konigkramer, T. (1997), Chapin, J. P. (1954), Clement, P. et al. (1993), Goodwin, D. (1982), Vincent, A. W. (1949), Weeks, J. T. (1948).

Estrilda poliopareia Reichenow. Anambra Waxbill. Astrild du Niger.

Plate 18

Estrilda poliopareia Reichenow, 1902. Orn. Monatsb., 10, p. 185; ‘‘Congo’’, restricted to S Nigeria by Chapin, 1950, Bull. Br. Orn. Club, 70, p. 24. Estrilda anambrae, 1907. J. Orn., 55, p. 624; Anambra Creek, Agoulerie, S Nigeria.

(Opp. p. 267)

Forms a superspecies with E. paludicola. Range and Status. Endemic resident, S Nigeria; known from 3–4 localities: Onitsha area (lower R. Niger),

Forcados (W coast of Niger delta) and Badagri (brackish creek and longshore island west of Lagos). Discovered in

294

ESTRILDIDAE

Estrilda poliopareia

Anambra Creek, Agoleri (Agoulerie), near mouth of R. Okulu (¼ Eruku), 30 km northeast of Onitsha; 24 birds collected there in 1905. Common at Onitsha, at least in 1950s; flock of 5 at Asaba, across Niger R. from Onitsha, in 1987 (Ash 1990). Small parties seen several times in Forcados area (Elgood et al. 1994) and in 1960s at Badagri (pers. obs., J. A. Button, pers. comm.). Several places in Tombia Ekpetiama area in Niger delta, with c. 40 birds in one from Mar 2001 (Bull. Afr. Bird Club 9, 2002, 147; World Birdwatch 24, Sept 2002, 2; Bull. Afr. Bird Club 10, 2003, 61). Classified as Vulnerable; possibly 180 localities as mapped (Lamarche 1993). Burkina Faso, uncommon in N Yatenga region in NW; 4 central localities (Holyoak and Seddon 1989); not uncommon in Ouagadougou area (Mauvais 1998); old records from Fada N’Gourma; rare, Arli Nat. Park in SE. Niger, frequent in ‘W’ Nat. Park; elsewhere, records at Gaya, Korogoungou, Gotheye, Mari, Niamey, Saga, Birni-Nkonni, Maradi, Zinder, and between Maradi and Tanout.

Guinea, common, Conakry. Liberia, frequent on Mt Nimba; 3, Monrovia, Mar 1989, perhaps escapes. Ivory Coast, very patchily distributed in N forest and S guinean savanna belt (Maroue´, Abengourou, uncommon in Comoe´ Nat. Park; also Abidjan golf-course, Demey and Fishpool 1991). Ghana, uncommon resident at Tumu, Karaga, Pong Tamale, Red Volta Valley, Kete Kratchi, Dodowa, Ada; frequent along coast from Cape Coast to Accra Plains. Togo, common in savannas north of Kara, 9
330 N; on coast, rare around Lome´ (seen on 5 days in 16 months, Browne 1980). Benin, uncommon in Pendjari Nat. Park and south to 10
170 N (Thonne´rieux 1988, Holyoak and Seddon 1989). Nigeria, locally common across N, from Kainji Lake Nat. Park to Sambisa (where abundant in dry season) and from close to Niger border (Sokoto, Saye and Farniso north of Kano, Maiduguri, Chad basin boreholes between Mongonu and Kauwa) south to Pategi and Shonga on Niger R., Gitata, Nindam (rare) and Yankari Nat. Park (rare). Cameroon, common in far N; occurs south to about Garoua and Bibe´mi. Chad, very common in soudanian zone, much

298

ESTRILDIDAE

ground, sometimes mixing with Common Waxbills E. astrild and mannikins Spermestes sp. Holds grass panicle in foot. Food. Seeds, including millet and (in captivity) Poa annua and Polygonum aviculare, and tips of young grass shoots; some invertebrates, including ant larvae, termites, aphids and spiders. Breeding Habits. Solitary nester; monogamous. In captivity, greeting, courtship and fluffed singing displays identical to those of Black-rumped Waxbill E. troglodytes, q.v. (Goodwin 1982). Nest built by both sexes, Y doing most work in early stages and carrying all larger pieces of material. Fixes decorative bits to top of nest, using shuddering side to side movements. Y’s behaviour in relation to cock’s nest identical to that of Black-rumped Waxbill; Y never roosted in it; twice, dead nestling removed from proper nest and placed in cock’s nest; Y spent minutes systematically killing every ant on ground near nest proper (Goodwin 1982). When collecting nest material, pulls grass head down, holds it under foot, and bites cleanly through stem. Parents excitable when nesting, fanning tails out wide and switching them from side to side (Timmis 1973). NEST: one was a sphere, made of grass heads, with side entrance; sited on ground in middle of quite short tuft of grass. One, in captivity, was large sphere made of grass leaves and stems, feathers, thin shreds of banana leaves, and hair, c. 50 mm from ground, resting on layer of dead leaves and grass; another was 9 m above ground, in thick

vegetation. Also in captivity, Y (paired for years with X Black-rumped Waxbill) built many nests, on floor of aviary or in a corner or in wall basket; floor nests pear-shaped, narrowing to tubular entrance with very small, concealed entrance hole surrounded by sharp ends of grass stems ‘so that it was impossible to insert a finger without some of them stabbing it quite sharply’ (Goodwin 1982); outer surface decorated with feathers, mainly white or glossy black ones, bits of thin paper, and small clumps of earth bound by rootlets made of grass stems with blades and/or seeding heads attached; walls 25 mm thick, compact, tough; lined with fine stems and flowering panicles of grass; all nests had a ‘cock’s nest’ built above, same inside dimensions as proper nest but walls thinner, with fairly large side entrance. EGGS: 4. White. SIZE: 1, 137  108. LAYING DATES: Eritrea, (birds at nests, Nov); Ethiopia, Aug; Zaı¨re, Sept; E Africa: Uganda, (‘attempted to breed’ Sept, juvs June: Carswell 1986); Region B, May, July, Region C, Jan, Mar–May, Region D, Mar–Apr, July, ˆ i, Feb, Apr. immediately after long rains; Malaw INCUBATION: by Y (and doubtless by X). Period: 12–14 days (Timmis 1973). DEVELOPMENT AND CARE OF YOUNG: at first young brooded mainly by X and fed mainly by Y. Young leave nest at 25 days and return to it to sleep at night for c. 7 days. BREEDING SUCCESS/SURVIVAL: parasitized by Pin-tailed Whydah Vidua macroura. Key References

Goodwin, D. (1982), Timmis, W. H. (1973).

Plate 19

Estrilda troglodytes Lichtenstein. Black-rumped Waxbill. Astrild cendre ´.

(Opp. p. 282)

F.[ringilla] troglodytes Lichtenstein, 1832. Verz. Doubl. zool. Mus. Berlin, p. 26; Senegambia. Forms a superspecies with E. rufibarba (Arabia), E. astrild and E. nigriloris. Range and Status. Endemic resident and wanderer, Senegal to SW Eritrea and W Kenya, partially migratory at some borders of range. Introduced successfully to Puerto Rico and Guadeloupe and unsuccessfuly to Hawaii (Oahu) and Tahiti; reported annually at Gibraltar since 1975 (I. Hinze, pers. comm.). Senegal, widespread but very patchy and local; can be common, e.g. in regions of Nioro-du-Rip, Kaolack and Kidira. Gambia, uncommon in W, locally very common inland. Mauritania, middle and upper Senegal R. valley; Karakoro valley; frequent at Nouakchott, thought to be escapes. Mali, locally quite common, >180 localities as mapped (Lamarche 1993). Burkina Faso, uncommon in N Yatenga region in NW; 4 central localities (Holyoak and Seddon 1989); not uncommon in Ouagadougou area (Mauvais 1998); old records from Fada N’Gourma; rare, Arli Nat. Park in SE. Niger, frequent in ‘W’ Nat. Park; elsewhere, records at Gaya, Korogoungou, Gotheye, Mari, Niamey, Saga, Birni-Nkonni, Maradi, Zinder, and between Maradi and Tanout.

Guinea, common, Conakry. Liberia, frequent on Mt Nimba; 3, Monrovia, Mar 1989, perhaps escapes. Ivory Coast, very patchily distributed in N forest and S guinean savanna belt (Maroue´, Abengourou, uncommon in Comoe´ Nat. Park; also Abidjan golf-course, Demey and Fishpool 1991). Ghana, uncommon resident at Tumu, Karaga, Pong Tamale, Red Volta Valley, Kete Kratchi, Dodowa, Ada; frequent along coast from Cape Coast to Accra Plains. Togo, common in savannas north of Kara, 9
330 N; on coast, rare around Lome´ (seen on 5 days in 16 months, Browne 1980). Benin, uncommon in Pendjari Nat. Park and south to 10
170 N (Thonne´rieux 1988, Holyoak and Seddon 1989). Nigeria, locally common across N, from Kainji Lake Nat. Park to Sambisa (where abundant in dry season) and from close to Niger border (Sokoto, Saye and Farniso north of Kano, Maiduguri, Chad basin boreholes between Mongonu and Kauwa) south to Pategi and Shonga on Niger R., Gitata, Nindam (rare) and Yankari Nat. Park (rare). Cameroon, common in far N; occurs south to about Garoua and Bibe´mi. Chad, very common in soudanian zone, much

Estrilda troglodytes NESTLING: skin yellowish, pale bluish down, conspicuous bluish white gape tubercles; mouth pattern like that of E. astrild, q.v.

Estrilda troglodytes

Field Characters. Length 95–105 cm. Like Common Waxbill E. astrild, with red bill and eye-stripe, but tiny, paler and greyer, barring very faint; black rump and tail (outer feathers edged white). Immature has black bill and smudgy dark eyeline.

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less so in sahel zone. Sudan, frequent, although recorded from 25 pairs per km2 in secondary forest; in logged forest, higher densities along roads, with 3 pairs per km2 in interior of mature forest (Gatter 1997); abundant in NE Gabon, 1 pair per 2–3 ha. Description. S. h. haematina (Vieillot): Sierra Leone to Ghana. ADULT Y: top and sides of head to narrow (2–3 mm) band across chin at base of mandible black; entire upperparts glossy black, tail black. Lower chin, throat, upper breast and flanks bright red; rest of underparts sooty black. Upperwings black, flight feathers brownish when worn; axillaries and underwing-coverts sooty black. Bill shiny blue, tip pinkish red; eyes brown, chestnutbrown or reddish, narrow pale bluish eye-ring; legs black or blackish brown. ADULT X: differs from ad. Y in having forehead, forecrown, lores and around eye to cheeks and ear-coverts maroon, forming large face patch; long uppertail-coverts crimson-maroon; rest of upperparts slate grey; chin to upper breast and flanks orange-red with some buffy feathers, merging into dark sides of head; lower breast to undertail-coverts blackish brown, densely spotted white. SIZE (10 YY, 10 XX): wing, Y 65– 71 (672), X 65–70 (666); tail, Y 50–57 (545), X 51–57 (543); bill, Y 18–20 (189), X 175–20 (186); tarsus, Y 205–215 (210), X 19–215 (204). WEIGHT: Liberia, Y (n ¼ 9) 199–256 (225), X (n ¼ 7) 181–237 (219); Nigeria, Y (n ¼ 4) 221–247 (230); Congo (unsexed, n ¼ 18) 20–26 (230±17). IMMATURE: juv. Y sooty black above, sooty brown below, browner on throat, tips of breast feathers tinged orange-red; large

Spermophaga haematina

? ? ?

Mali: see introduction.

uppertail-coverts dull orange-red; bill steely dark blue, eyes brown. Juv. X similar but browner above, dark tawny brown below. NESTLING: pale flesh-coloured with pale grey down on head, back and shoulders; swollen yellowish white tubercles on both mandibles; palate yellow with 3 large dark spots and 2 smaller ones behind them, a dark spot at corner of upper mandible at gape, narrow dusky bar across tongue broken in middle, crescentshaped dark mark on inside of lower mandible near tip (fig. p. 252). S. h. togoensis Neumann: Togo to SW Nigeria. Like nominate race, but Y has uppertail-coverts dark crimson. S. h. pustulata (Voigt): SE Nigeria to Central African Republic, Congo and Zaı¨re. Y has red extending from chin and throat to lower part of cheeks; X has face patch reddish (not dark maroon), extending to lores and above eye but not across forehead; uppertail-coverts brighter crimson than togoensis in both sexes. Nestling has 2 dark streaks on inside of upper mandible near tip.

Field Characters. Length 15 cm. Y told from other bluebills and from Crimson Seedcracker Pyrenestes sanguineus by mainly black head; in W (Gambia to Ghana) also lacks red on uppertail-coverts. Has been mistaken for Blue-billed Malimbe Malimbus nitens, which has red breast patch not extending onto flanks or throat, slimmer bill, and is noisy and readily observed. X of eastern race pustulata told from Red-headed Bluebill S. ruficapilla, which it overlaps in E Zaı¨re, by dark top of head. Best found in dark undergrowth by shiny pale blue bill. Shy and retiring; makes its presence known by sharp alarm calls or sweet song.

Spermophaga haematina Voice. Tape-recorded (104, B, CHA, ERA, GRI, KEI, LEM, LER). Varied, musical song of long, pure, sweet notes, up-slurred and down-slurred, tending to rise in pitch toward the end. Calls include high, thin ‘sseeeee’, sharp ‘chip’ or ‘tsip’ (contact), chaffinch-like ‘tswink-tswinktswink’ and ‘tac’ in alarm; another alarm call begins as a relatively slow ‘tsip-tsap’ and accelerates until notes run into each other; all alarm calls accompanied by violent wing and tail flicking (Goodwin 1982). An extended, rambling chatter may be a call or a different song type (Chappuis 2000). General Habits. Inhabits thick undergrowth in moist parts of primary and secondary lowland forest, especially clearings and edges with Marantaceae; in Gabon, old roads in logged forest overgrown with bushes of Scleria and Afromomum; gallery forest; prefers damp places and river edge, but in Gambia also in dry coastal palm forest and remnant forest thickets away from water; also occupies farmbush, forestry plantations and logged forest (Sierra Leone), oil palm plantations and mixed agriculture (Guinea-Bissau), young secondary growth around villages and in suburban areas (Nigeria) botanical gardens (Cameroon: Limbe), thick stands of elephant grass (Mt Cameroon). Catholic in its choice of habitats: in Liberia occurs in any dense undergrowth from coastal savanna scrub to mature high forest. In pairs or family parties, including young from previous brood; forages on ground or among low plants and lianas, typically below 5 m, sometimes to mid levels; joins mixed-species flocks of insectivores. Attracted to water to drink and bathe. Watchful, giving alarm calls to warn of approach of person. On ground holds tail in stiff broad fan (Barlow et al. 1997). 2 birds killed flying into windows suggest local movements may occur (Ghana); local movements evident in S Congo (Dowsett-Lemaire and Dowsett 1991). Moves around within its habitat: of 55 ringed birds, only 5 were recaptured at the same spot, more than a month later; one Y taken at same place only 4 times in 25 months, another 3 times in 16 months; one moved 1200 m in 7 months (Gabon: Brosset and Erard 1986).

NEST: large ball, with side entrance which lacks or has only a suggestion of an entrance tube, built of moss, ferns, dry leaves and grass stems and panicles, unlined or lined with grass or vegetable down, sometimes feathers; well hidden, 12–2 m above ground in thick brush, once in pile of sticks and debris left by flood in fork of tree (Gabon: Christy and Clarke 1994); built by both sexes in the wild, though in captivity only by Y. EGGS: 2–6; white, unglossed. SIZE: (n ¼ 3) 19– 195  135–14. LAYING DATES: Senegal, Nov–Dec; Sierra Leone (egg in oviduct June); Liberia (breeding condition July–Sept, nest-building June and Sept–Nov, dependent fledglings Oct, Dec–Jan, independent young Sept–Feb, immature plumage Oct–April); Ivory Coast (juv. Mar); Ghana, June (nestlings Sept); Benin (X with large brood patch Apr); Nigeria, Aug–Oct (breeding condition June); Cameroon, Jan, May, Aug, Oct (X with brood patch July, Oct–Dec, juv. July); breeding season not well marked (Eisentraut 1963); Central African Republic (breeding condition July– Nov); Gabon, Oct–Feb, May, in NE exceptionally during dry season in June; Zaı¨re, Itombwe, breeding condition Dec–Jan. INCUBATION: by X alone in the wild, in captivity also by Y during day. Period (captivity) 14–16 days; time from laying of eggs to fledging of young, 27 days (Brosset and Erard 1986). DEVELOPMENT AND CARE OF YOUNG: ad. plumage attained at 3 months; fledglings remain with parents a minimum of 1–2 months after leaving nest, joining them in mixed species flocks. In captivity, young brooded at night for first 5–6 days only, fledge at 18–22 days, returning to nest to roost for next few nights, begin feeding themselves after 2– 3 days, independent at 12 days. (NOTE: remarks supposedly pertaining to this species by Lowe (1982) may refer to S. ruficapilla, since he states that it occurs in eastern Africa; they are not included here.)

Key References (1982).

Brosset, A. and Erard, C. (1986), Goodwin, D.

Food. Berries, grass seeds, rice, husks of oil palm fruit; insects, including swarming termites and aphids; spiders. In captivity, soaked or sprouted grains, small dry seeds (refuses large hard ones); eggfood; plentiful insects, e.g. wax moths and their larvae, house fly maggots and pupae, small grasshoppers and crickets, whiteworms, newly moulted mealworms, smooth caterpillars (Goodwin 1982). Breeding Habits. In display Y with nest material in bill and uttering fluty song bobs up and down like displaying Estrilda and rhythmically opens wings; if X flies to him he drops material, mandibulates, spreads wings at shoulders in a ‘heraldic eagle’ manner and throws head back so red breast is fully displayed, meanwhile vibrating his tail (R. Neff in Goodwin 1982); X sits nearby but copulation does not usually follow. In another display, Y hops to and fro in front of X with grass stem in bill.

Grant’s Bluebill S. poliogenys from Mt Hoyo, Ituri, Zaı¨re, with black feathering around bare eyering (see p. 313). (This drawing and D on p. 321 by C.H. Fry from photos by H. Lehaen in Lippens and Wille, 1976).

315

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ESTRILDIDAE

Plate 20

Spermophaga ruficapilla (Shelley). Red-headed Bluebill. Se ´ne´gali `a teˆte rouge.

(Opp. p. 283)

Spermospiza ruficapilla Shelley, 1888. Proc. Zool. Soc. London, p. 30; Bellima, upper Uele district. Range and Status. Endemic resident. Angola, locally common from N Cuanza Norte and Uı´ge south through escarpment forests to Gabela in Cuanza Sul, and at Dundo in N Lunda Norte; Central African Republic (Baroua); Zaı¨re as mapped, chiefly in NE, west to Ibembo and south to Itombwe, also in Kasai, south to Dibaya; Rwanda (Nyungwe, Burunga, Akanyaru R.), Burundi (Musigati, Bujumbura, Bururi Forest); extreme S Sudan along Zaı¨re border and in Lotti Forest, up to 2400 m, fairly common; Uganda, widespread and usually common at 700–2400 m from Budongo, Bugoma, Bwamba, Kibale and Impenetrable (Bwindi) Forests to Malabigambo Forest, Entebbe, Mpumu, Mabira and Elgon; fairly common W Kenya from Elgon, Kapenguria and Saiwa Nat. Park through Kakamega and Nandi to W Mau, Trans-Mara, Kilgoris, Lolgorien and Rapogi, and in Chemorogok Forest, Eldama Ravine (Pearson and Turner 1998); uncommon in Meru and Embu Forests; Tanzania in NW (Minziro Forest and other localities in Bukoba District), W (Gombe and Mahari Mt) and NE (Usambaras). Description. S. r. ruficapilla (Shelley): range of species except NE Tanzania. ADULT Y: head and neck to upper breast and flanks bright crimson-red; narrow bluish white eye-ring, broader below eye than above (photo in Oatley 2001); long uppertail-coverts crimson; rest of upperparts black, tail black. Lower breast to undertail-coverts sooty black. Upperwing blackish brown, outer edges of tertials and greater coverts and tips of median and lesser coverts black, axillaries and underwing-coverts sooty black. Bill bright silvery blue, cutting edges and tip pinkish red; eyes brown; legs dark olive brown to blackish. ADULT X: like Y but buffy brown feathers show through on head and breast; upperparts dark slaty grey; belly sooty black densely spotted white; upperwing dark grey-brown, tertial edges and covert tips dark slaty grey. SIZE (10 YY, 10 XX): wing, Y 68–75 (721), X 68–71 (698); tail, Y 52–60 (567), X 48–58 (546); bill, Y 165–185 (178), X 17–185 (177); tarsus, Y 21–23 (224), X 21–225 (216). WEIGHT: Central African Republic, Y (n ¼ 7) 22–27 (243), X (n ¼ 6) 24–28 (248); Kenya, Y (n ¼ 21) 222–26 (241), X (n ¼ 24) 213–26 (232). Diurnal percentage weight change 174 (Kakamega, Kenya: Mann 1985). IMMATURE: juv. sooty grey above, long uppertail-coverts crimson-maroon; head to breast rich brown, paler, more tawny, from chin to upper breast, sides of upper breast tinged crimsonmaroon; lower underparts dark tawny-brown, barred sooty black. Wings and tail brownish black. Bill very dark bluish with a little red at tip, eyes dark brown, legs and feet dusky or greenish black. NESTLING: at hatching, yellowish flesh with a few down feathers on upperparts; bill starts to darken on day 2; mouth yellow, with 3 large black spots on palate and 2 smaller ones behind them (contra Chapin 1954: R.B. Payne, pers. comm.); 3 small yellow swellings at each side of gape; on day 6, 2 small black marks develop on inside of maxilla and mandible near tip. S. r. cana (Friedmann): Usambara Mts, NE Tanzania, sight record Meru forest, Kenya (see below). Y differs from nominate race in having slaty grey (not black) upperparts, often extending to nape; head and breast somewhat lighter shade of red; lower underparts dark grey-brown. X paler, more orange-red on head, upperparts (including nape) paler grey; only a tinge of crimson on uppertail-coverts; lower underparts dark grey-brown, spotted whitish. WEIGHT: Y (n ¼ 4) 213–236 (222), 1 X 217.

Spermophaga ruficapilla

TAXONOMIC NOTE: status of birds in Meru and Embu forests unclear; previously thought to be nominate race, but bird in Meru Forest reported as race cana.

Field Characters. Length 15 cm. Both sexes told from Black-bellied Seedcracker Pyrenestes ostrinus by black tail, smaller, curved bill with pink cutting edges and tip, and from the other bluebills by solid red head (nape black in Grant’s Bluebill S. poliogenys, top of head black in Western Bluebill S. haematina). Spotted X also has more extensive red on breast than Grant’s Bluebill, extending to flanks. Often detected by loud call. Voice. Tape-recorded (C, GREG, HOR, McVIC). Calls include loud hard ‘jip’ or ‘pik’, nasal ‘tewk’ and very high, thin ‘seep’, barely audible. Like Western Bluebill, appears to have 2 song types. Melodious song variously described as: 7 whistling or fluting notes, first 4 rising, last 3 falling in pitch, and ‘loud sequence of long-drawn fluting notes ending in a trill’ (Goodwin 1982); sibilant and whistling, beginning with short note, then short series of rising notes, and longer series of up-slurred notes, ‘si-sisi-su-wii si-sisswiswiswiswisiwiswi’, emphasis on last note (Stevenson and Fanshawe 2002); second song type also varied, with guttural trills, whistles, rattles, clucks and kissing sounds, given in an undertone; described as ‘a jumble of twitters and weak chirps’, mostly quite high and stuttery, varying suddenly in speed, rushing then slowing (Evans 1997). General Habits. Inhabits thick moist undergrowth in primary and secondary forest, especially along streams,

Pyrenestes

Food. Seeds, including Ipomoea (Convolvulaceae), Indigofera and other Leguminosae, Olyra latifolia (eaten especially by S. r. cana) and sword grass Imperator; insects, including weevils, other small beetles and winged termites. In captivity, seeds of white millet, hemp, sunflower and garden composite Tithonia diversifolia; greenstuff (chickweed, lettuce); mealworms, whiteworms, ant pupae, termites, crickets, wax moth larvae, minced meat, hard-boiled egg (Ellis 1977, Goodwin 1982); young fed mainly animal food.

back and sleeks down those on head and neck, producing hump-backed appearance, with tail angled toward X; he holds piece of grass, leaf or other nest material in bill, bows to one side toward X until bill is c. 45
below the horizontal, then throws back head until it is c. 70
above it, uttering second song type. Copulation may follow display; it is preceded by Y pecking X’s head, nape and back and/or by X soliciting with quivering tail; during copulation Y clings fluttering to X’s back for considerable time, then hops around her, bowing. In greeting ceremony after short absence, mates angle slightly spread tails towards each other. NEST: large untidy ball c. 200 diam. with side entrance but no entrance tube, built of dry leaves, ferns and coarse grass, lined with fine grass, plant down, flowers and a few feathers; one was placed in banana plant in abandoned field (Prigogine 1971), another was hidden under leaves in axils of drooping fronds at base of oil palm (van Someren and van Someren 1949); built by both sexes in the wild, in captivity mainly by Y. EGGS: 2–3; creamy white. Size (n ¼ 2) 210–219  14– 142. LAYING DATES: Angola (family parties of ads and young in May, July and Aug suggest egg-laying from Feb to May: Dean 2000); Sudan, Jan, Apr; Zaı¨re, (Itombwe) Oct–Mar, May; (lowlands north of equator, birds in breeding condition June–Nov); E Africa: Region B, Jan, Apr, June–July, Sept, Nov–Dec, Region D, Jan (nest-building Sept). INCUBATION: begins with last egg; by both sexes during day, by X only at night. Period: 17–18 days. DEVELOPMENT AND CARE OF YOUNG: young fledge at 20 days, are then called or led into cover by parents, but not back to nest; tended by both sexes, but if pair starts another brood while young still dependent, X stops feeding them before she begins to lay. BREEDING SUCCESS/SURVIVAL: ringed Y recaptured 8 years 10 months later; captive X lived >10 years.

Breeding Habits. In courtship display, given on ground or low perch, Y partly fluffs out feathers on belly, flanks and

Key References

thickets around clearings and along edges; also gallery forest, seasonal swamp forest, coffee forest (Angola), patches of tall grass, damp overgrown valleys, open areas with cultivation. In Tanzania occurred in sprawling thicket of shrubs and lianas not more than 6 m tall close to forest, and in area with tall dense shrub layer and 10 years.

Breeding Habits. In courtship display, given on ground or low perch, Y partly fluffs out feathers on belly, flanks and

Key References

thickets around clearings and along edges; also gallery forest, seasonal swamp forest, coffee forest (Angola), patches of tall grass, damp overgrown valleys, open areas with cultivation. In Tanzania occurred in sprawling thicket of shrubs and lianas not more than 6 m tall close to forest, and in area with tall dense shrub layer and 500 birds; considered threatened in the region (Parker 1999). Density in Zimbabwe (Dichwe Forest), 50 pairs per 10 ha. Formerly common in dense thickets on Zambezi floodplain in Mana Pools Nat. Park, Zimbabwe, but

Hypargos niveoguttatus

excessive concentrations of game animals have eliminated much of this habitat (A. J. Tree in Harrison et al. 1997).

Hypargos niveoguttatus Description. H. n. macrospilotus Mearns (includes ‘interior’, ‘idius’, ‘centralis’ and ‘baddeleyi’): range of species except nominate race. ADULT Y: forehead to nape ashy grey, tinged olive-brown; hindneck to rump tawny brown; uppertail-coverts dark crimson. Tail feathers blackish brown, T1 and outer webs of T2–T6 tinged crimson. Lores and above eye to ear-coverts, cheeks, sides of neck (extending toward hind neck) and chin to upper breast dark crimson. Rest of underparts sooty black, sides of breast and flanks with bold white spots (1 pair per feather). Median and lesser coverts warm brown; rest of upperwing blackish brown, secondaries and tertials broadly edged warm brown, greater coverts edged and tipped warm brown; axillaries and underwingcoverts white. Bill metallic violet-blue with black tip and culmen; bare eye-ring bright blue to whitish; eyes brown; legs blue-grey or slate grey. ADULT X: forehead to nape brownish grey; hindneck to rump warm olive-brown. Uppertail-coverts and tail as ad. Y, but crimson less intense. Lores and above eye to cheeks and ear-coverts greyish olive-green. Chin pale olive-yellow, merging into rufous-chestnut throat, upper breast and sides of neck. Underparts dark olive-grey, tinged green on flanks and undertail-coverts, flanks spotted white as in ad. Y. Upperwing as ad. Y; axillaries and underwing-coverts creamy white. SIZE (10 YY, 10 XX): wing, Y 55–60 (575), X 54–58 (557); tail, Y 49–55 (529), X 48–54 (511); bill, Y 13–145 (138), X 13–14 (134); tarsus, Y 165–18 (173), X 165–175 (170). WEIGHT: Kenya, Y (n ¼ 29) 124–169 (148), X (n ¼ 31) 129–163 (144); Zambia, unsexed (n ¼ 12) 12–15 (138); southern Africa, Y (n ¼ 23) 131– 171 (15), X (n ¼ 23) 125–179 (149). IMMATURE: juv. differs from ad. X in having olive-green (not reddish) breast and only trace of white spotting on sides; underparts sooty grey, sides and flanks olive-green. NESTLING: newly hatched bird flesh-coloured with some greyish down, skin becoming dark grey or black on day 1 (A); palate yellow with 3 dark spots, only a trace of dark crescent in lower mandible, tongue and inner and lower mouth pink and unmarked; gape with 2 large white tubercles on each side constricted by white gape corner (fig. p. 252) lower tubercle develops lateral spot that turns yellow by day 5 and orange at fledging (B, R. B. Payne, pers. comm.). H. n. niveoguttatus (Peters): E Zimbabwe and S Mozambique. Like macrospilotus, but Y slightly less rich red; X with red suffusion on chest reduced or lacking, olive-grey of lower underparts typically paler.

Field Characters. Length 125–13 cm. Y with red face, throat and breast and white-spotted black belly superficially like X Red-headed Bluebill Spermophaga ruficapilla but smaller, with rich brown upperparts and white spots confined to flanks; told from Pink-throated Twinspot H. margaritatus by deep red throat, grey crown. X has grey face, buffy yellow throat shading to orange on breast (X

Pink-throated entirely grey below). Juv. has buffy face and throat, brown underparts without spots, some black on belly. Voice. Tape-recorded (99, 103, 104, B, F, CART, GUT, LEM, PAY). Song variable; high thin notes accelerating into trill, or at different speeds interrupted by whistles, ‘titititititititi-way-twee-chachachacha-trrrrrrrrr’; often mixed with pure sweet note; sometimes a single note followed by a trill, or whistles may precede trill, ‘sip, tooo, tsssrrrrrr’ (Maclean 1993); in captivity a slower, more relaxed song with short trills at different pitches and slurred fluty notes. Close contact call soft ‘tseet’ or ‘tsee-et’, distance contact calls, louder, more drawn-out and emphatic ‘tseeet’, or rapid trill; alarm, harsh ‘tschee’. Nest call begins with close contact call, then quick series of notes, ‘beetbeetbeetbeet’ repeatedly interrupted by contact calls (Goodwin 1982). General Habits. Inhabits edges of evergreen forest with tangles of Afromomum and other shrubs, moist or dry riparian forest, dense thickets around pools in forest interior (Zaı¨re: Upemba Nat. Park), undergrowth inside more open forest, e.g. Sokoke, Kenya, where it occurs in rain forest, Afzelia and Cynometra habitats (Britton and Zimmerman 1979); dense coastal bush, moist bushland, woodland thickets, thick herbage by water, dense secondary scrub; in Zambia, ground stratum of wide variety of cover, from thickets on termite mounds in miombo woodland to drier riparian forest and thickets, especially in major river valleys; also Cryptosepalum forest and Marquesia thickets; undergrowth of plantations and other cultivation; prefers moister situations than Pink-throated Twinspot. Mainly lowlands, but in E Africa up to 2000 m. Pairs or small parties; forages on ground, often at edges of clearings or on path or road. Seen hawking insects in captivity (Bishop 1974). Quiet but not particularly shy, often tame and inquisitive, occurring near human habiˆ i). In captivity, Y tation, even foraging on lawns (Malaw more cautious than X. Food. Seeds of Olyra latifolia and other grasses; insects, including ants. In captivity, canary seed, soaked or sprouted millet, half ripe seeds of grasses and chickweed, high-protein seed mix, nectar food and pears; needs insects year round, e.g. live ant pupae, housefly larvae, aphids, chrysalids, blowflies, maggots, mealworms, hairless caterpillars; also takes worms and spiders (Brickell 1980). Breeding Habits. Territorial, competitive and aggressive. In communal display, several pairs congregate towards

343

344

ESTRILDIDAE

evening in patch of thick undergrowth; YY hop around, keeping several m apart and display as if to each other, uttering sharp ‘chip’ followed by long descending musical trill; XX simply sit and watch. In courtship display in captivity, Y dances around X, with or without feather or other nest material in bill, red feathers puffed up, tail spread and partly angled, uttering faint song; every s during display he performs bobbing movement, head alternately pointed upward at c. 70
angle and bowed to c. 30
; he starts with head elevated and body low to ground, then bows and throws head up, while hopping c. 1 cm off ground; he slowly bows head, and repeats the sequence. X remains quiet, or sometimes dances, hopping slowly with no material in bill, even singing quietly if mate absent; if she responds by soliciting with quivering tail, copulation follows (Harrison and Dormer 1962, Yantz 1988). Allopreening common between pair members. Possibly doublebrooded (Bishop 1974). NEST: ball or domed cup with side entrance, with or without tunnel, made of grass, rootlets, fern stalks, leaf skeletons, and dry moss, exterior covered with decomposing leaves, sometimes with incomplete ‘cock’s nest’ on top, lined with fine grass, dry moss or feathers; chamber c. 80 high, entrance diam. 40; on ground or low down in bush or tree. 1 nest was small domed cup, ext. diam. 120, with tunnel entrance 80 long, placed on ground between roots of tree growing out of low bank in riparian forest; floor of tunnel was bare ground; materials in order of dominance: small maidenhair fern stalks, fine hair roots, small leaf skeletons, dry moss (mainly as lining); almost no grass; top camouflaged by decomposing leaflets; possibly an old nest of Eastern Bearded Scrub-Robin Cercotrichas quadrivirgata covered and tunneled by the birds; another was sited in cavity in bank of dry stream bed, among roots of large tree (Zimbabwe: Edwards 1986, and nest record cards in National Museum, Bulawayo); built by both sexes. Unmated captive Y built several nests on his own (Yantz 1988).

EGGS: 3–6; white, unglossed. SIZE: (n ¼ 3) 157–160  122–125. LAYING DATES: E Africa, Region E, May, July; Zambia, ˆ i, Dec–May; Zimbabwe, all months Jan, Mar, May; Malaw except July–Aug, peak in Mar–June; Mozambique, Mar (partially fledged young July, breeding condition Dec). INCUBATION: by both sexes during day, by X only at night; at changeover, bird brings piece of material to nest entrance and calls softly, birds change places quickly; soft material brought more often after eggs hatch, and if no other material available, X even plucks feathers from flanks of Y, which tolerates this activity (captivity: Yantz 1988). Period: 12–13 days (12–14 in captivity). Birds sit very tight, almost refuse to leave nest on approach of person. DEVELOPMENT AND CARE OF YOUNG (data mainly for captive birds, from Yantz 1988): young fed by both parents; enough ‘live’ food must be available or parents will be unable to raise entire brood; live food consumption moderate at first, increases slightly at day 3, sharply at days 5–6, peaks at days 9–11 and tapers off until fledging; if insufficient food available, parents will remove and dispose of 1 or more nestlings until number that can be supported is reached. For first 6 days young produce low, sustained begging sounds which are apparently unrelated to actual begging for food; parents attempt to cover these up by calling loudly and giving alarm trills, drawing attention away from nest. Nestling period: 21 days (18–20 days in captivity). Fledged young cared for by parents for 12–14 days, even longer if they beg persistently; fully independent at c. 18 days; start assuming ad. colours 5 weeks after that; in the wild, sometimes return to roost in nest for c. 3 days after they are capable of flight. BREEDING SUCCESS/SURVIVAL: parasitized by Zambezi Indigobird Vidua codringtoni.

Key References Brickell, N. (1980), Goodwin, D. (1982), Maclean, G. L. (1993), Yantz, J. (1988).

Plate 20

Hypargos margaritatus Strickland. Pink-throated Twinspot. Se´ne ´gali de Verreaux.

(Opp. p. 283)

Spermophaga margaritata Strickland, 1844. Ann. Mag. Nat. Hist. 13, p. 148, p1. 10; Madagascar (¼ Coguno, Inhambane district, Mozambique, see W.L. Sclater, 1911, Ibis, p. 230). Forms a superspecies with H. niveoguttatus. Range and Status. Endemic resident with restricted range: S Mozambique (Save R., Sul do Save), E Swaziland and KwaZulu-Natal south to L. St Lucia, and marginally in NE and extreme SE Transvaal; 2 records further west in N Transvaal; recently found north of Save R. at Panda (Bull. Afr. Bird Club 3, 1996, 140). Uncommon to locally fairly common; population in Mozambique probably >80,000, although up to 2000 are exported annually for the cage bird trade (Parker 1999). In South Africa it occurs in substantial areas of protected habitat. Description. ADULT Y: forehead to rump warm brown; uppertailcoverts deep pink. Tail feathers dark grey-brown, T1 and outer

webs of T2–T6 tinged pink. Lores and above eye, cheeks, earcoverts and chin to upper breast deep pink; rest of underparts sooty black, with large paired pinkish white spots on sides of breast and flanks, and in narrow band bordering upper breast. Primaries, secondaries and primary coverts dark grey-brown, edged warm brown; tertials grey-brown with warm brown outer webs; alula and greater, median and lesser coverts warm brown. Axillaries and underwing-coverts white. Bill dark greyish blue; bare eye-ring mauve-pink to blue; eyes dark brown; legs blackish. ADULT X: upperparts as ad. Y. Lores and sides of head to above eye brownish grey; chin pale grey-buff; throat, upper breast and sides of neck pale grey, tinged brownish; centre of lower breast to centre of belly pale grey-buff; undertail-coverts blackish, broadly tipped white; sides of breast and flanks and narrow band

344

ESTRILDIDAE

evening in patch of thick undergrowth; YY hop around, keeping several m apart and display as if to each other, uttering sharp ‘chip’ followed by long descending musical trill; XX simply sit and watch. In courtship display in captivity, Y dances around X, with or without feather or other nest material in bill, red feathers puffed up, tail spread and partly angled, uttering faint song; every s during display he performs bobbing movement, head alternately pointed upward at c. 70
angle and bowed to c. 30
; he starts with head elevated and body low to ground, then bows and throws head up, while hopping c. 1 cm off ground; he slowly bows head, and repeats the sequence. X remains quiet, or sometimes dances, hopping slowly with no material in bill, even singing quietly if mate absent; if she responds by soliciting with quivering tail, copulation follows (Harrison and Dormer 1962, Yantz 1988). Allopreening common between pair members. Possibly doublebrooded (Bishop 1974). NEST: ball or domed cup with side entrance, with or without tunnel, made of grass, rootlets, fern stalks, leaf skeletons, and dry moss, exterior covered with decomposing leaves, sometimes with incomplete ‘cock’s nest’ on top, lined with fine grass, dry moss or feathers; chamber c. 80 high, entrance diam. 40; on ground or low down in bush or tree. 1 nest was small domed cup, ext. diam. 120, with tunnel entrance 80 long, placed on ground between roots of tree growing out of low bank in riparian forest; floor of tunnel was bare ground; materials in order of dominance: small maidenhair fern stalks, fine hair roots, small leaf skeletons, dry moss (mainly as lining); almost no grass; top camouflaged by decomposing leaflets; possibly an old nest of Eastern Bearded Scrub-Robin Cercotrichas quadrivirgata covered and tunneled by the birds; another was sited in cavity in bank of dry stream bed, among roots of large tree (Zimbabwe: Edwards 1986, and nest record cards in National Museum, Bulawayo); built by both sexes. Unmated captive Y built several nests on his own (Yantz 1988).

EGGS: 3–6; white, unglossed. SIZE: (n ¼ 3) 157–160  122–125. LAYING DATES: E Africa, Region E, May, July; Zambia, ˆ i, Dec–May; Zimbabwe, all months Jan, Mar, May; Malaw except July–Aug, peak in Mar–June; Mozambique, Mar (partially fledged young July, breeding condition Dec). INCUBATION: by both sexes during day, by X only at night; at changeover, bird brings piece of material to nest entrance and calls softly, birds change places quickly; soft material brought more often after eggs hatch, and if no other material available, X even plucks feathers from flanks of Y, which tolerates this activity (captivity: Yantz 1988). Period: 12–13 days (12–14 in captivity). Birds sit very tight, almost refuse to leave nest on approach of person. DEVELOPMENT AND CARE OF YOUNG (data mainly for captive birds, from Yantz 1988): young fed by both parents; enough ‘live’ food must be available or parents will be unable to raise entire brood; live food consumption moderate at first, increases slightly at day 3, sharply at days 5–6, peaks at days 9–11 and tapers off until fledging; if insufficient food available, parents will remove and dispose of 1 or more nestlings until number that can be supported is reached. For first 6 days young produce low, sustained begging sounds which are apparently unrelated to actual begging for food; parents attempt to cover these up by calling loudly and giving alarm trills, drawing attention away from nest. Nestling period: 21 days (18–20 days in captivity). Fledged young cared for by parents for 12–14 days, even longer if they beg persistently; fully independent at c. 18 days; start assuming ad. colours 5 weeks after that; in the wild, sometimes return to roost in nest for c. 3 days after they are capable of flight. BREEDING SUCCESS/SURVIVAL: parasitized by Zambezi Indigobird Vidua codringtoni.

Key References Brickell, N. (1980), Goodwin, D. (1982), Maclean, G. L. (1993), Yantz, J. (1988).

Plate 20

Hypargos margaritatus Strickland. Pink-throated Twinspot. Se´ne ´gali de Verreaux.

(Opp. p. 283)

Spermophaga margaritata Strickland, 1844. Ann. Mag. Nat. Hist. 13, p. 148, p1. 10; Madagascar (¼ Coguno, Inhambane district, Mozambique, see W.L. Sclater, 1911, Ibis, p. 230). Forms a superspecies with H. niveoguttatus. Range and Status. Endemic resident with restricted range: S Mozambique (Save R., Sul do Save), E Swaziland and KwaZulu-Natal south to L. St Lucia, and marginally in NE and extreme SE Transvaal; 2 records further west in N Transvaal; recently found north of Save R. at Panda (Bull. Afr. Bird Club 3, 1996, 140). Uncommon to locally fairly common; population in Mozambique probably >80,000, although up to 2000 are exported annually for the cage bird trade (Parker 1999). In South Africa it occurs in substantial areas of protected habitat. Description. ADULT Y: forehead to rump warm brown; uppertailcoverts deep pink. Tail feathers dark grey-brown, T1 and outer

webs of T2–T6 tinged pink. Lores and above eye, cheeks, earcoverts and chin to upper breast deep pink; rest of underparts sooty black, with large paired pinkish white spots on sides of breast and flanks, and in narrow band bordering upper breast. Primaries, secondaries and primary coverts dark grey-brown, edged warm brown; tertials grey-brown with warm brown outer webs; alula and greater, median and lesser coverts warm brown. Axillaries and underwing-coverts white. Bill dark greyish blue; bare eye-ring mauve-pink to blue; eyes dark brown; legs blackish. ADULT X: upperparts as ad. Y. Lores and sides of head to above eye brownish grey; chin pale grey-buff; throat, upper breast and sides of neck pale grey, tinged brownish; centre of lower breast to centre of belly pale grey-buff; undertail-coverts blackish, broadly tipped white; sides of breast and flanks and narrow band

Hypargos margaritatus

Hypargos margaritatus

bordering upper breast sooty black with large white spots (no pink tinge). Wings, tail and bare parts as ad. Y. SIZE: wing, Y (n ¼ 3) 53–54 (533), X (n ¼ 2) 52, 53; tail, Y (n ¼ 2) 50, 54, X (n ¼ 1) 47; bill, Y (n ¼ 3) 11–125 (117), X (n ¼ 1) 11; tarsus, Y (n ¼ 2) 165, 17, X (n ¼ 2) 155, 17; also: wing Y (n ¼ ?) 52–56, X (n ¼ ?) 52–54; (unsexed, n ¼ 8), tail 50–54, bill 11–12, tarsus 155–17 (Maclean 1993). WEIGHT: 1 Y 135, 2 XX 12, 13. IMMATURE: like X but underparts pale buff, without spots. NESTLING: pink, naked except for some down on head; palate yellow with 3 black triangular spots, gape tubercles white lined with black, 2 on each side, becoming blue-green within a few days of hatching and blue at fledging, rest of mouth and tongue reddish.

Field Characters. Length 12–13 cm. Very similar to Redthroated Twinspot H. niveoguttatus; Y paler, with pink, rather than deep red, face, throat and breast, brown crown, spots on underparts pale pink, not pure white; X more easily separated from X Red-throated, with pale grey throat and breast without any orange, white belly; juv. pale buff below. Voice. Tape-recorded (88, 99, B, F). Call a high thin trill on one pitch, variable in length; this may develop into a 2-part buzzy song type, ‘titidee-titidee, tidee-tidee-tideetidee-tidee’; also said to have a short whistled song like Red-billed Firefinch Lagonosticta senegala, q.v. Other calls include soft ‘seesee’ or ‘sisi’ and a low, trilling ‘zirrr’. At least some calls distinct from those of Red-throated Twinspot (Immelmann et al., in Goodwin 1982). General Habits. Inhabits dry woodlands with dense undergrowth and thickets and comparatively little grass; thorn and mixed scrub tangles in bushveld, palm scrub,

thorny cover at edges of evergreen forest, and along tracks, footpaths and little-used roads, mainly within East Coast Littoral and Arid Woodland vegetation types, but also adjacent to Afromontane Forest along Lebombo Range; in Kruger Nat. Park in northeastern sandy Nyandu Bush (Harrison et al. 1997). Very dense scrubby habitat on Lebombo Flats east of Komatipoort dominated by Acacia nilotica, A. borleae, Ziziphus mucronata, Maytenus heterophylla, Carissa bispinosa and 2 Euclea spp. (Edmonds 1984). Occurs in drier habitats than Red-throated Twinspot and Green Twinspot Mandingoa nitidula. Sea level to 600 m. In pairs and family parties. Usually close to ground; forages on ground, flying up into cover at first sign of danger, birds calling to one another while retreating (Clancey 1985); inconspicuous, keeping to dense cover, usually detected by soft call. Associates with other estrildines, especially firefinches Lagonosticta spp. and Blue Waxbills Uraeginthus angolensis. No seasonal movements suspected. Food. Seeds; probably also insects. In captivity, panicum millet, canary seed, soaked and sprouted millet and ant pupae. Breeding Habits. NEST: only 2 described: one oval, made of leaf ribs and skeletons, mixed with leaves, inflorescences and spider web, with side entrance 40–60 diam., lined with palm fibres, placed near ground at base of palm frond (Maclean 1993), the other an untidy ball-shaped structure of dry grass, lined with a few feathers, with side entrance (Ginn et al. 1989). EGGS: 3 (4 in captivity); white. SIZE (n ¼ 3) 15–156  12 (154  12). LAYING DATES: Mozambique ‘during the rains’; South Africa, Jan. DEVELOPMENT AND CARE OF YOUNG: fledgling almost uniform greyish brown; bill blackish, greyer basally and pale orange at gape, behind which is white Z-shaped flange (A); eye black, bare orbital ring broad, flat (not swollen as it is in ad.), pale mauve.

Key References Clancey, P.A. (1985), Goodwin, D. (1982), Harrison, J.A. et al. (1997), Maclean, G.L. (1993).

345

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ESTRILDIDAE

Genus Clytospiza Shelley

Single species of twinspot, related to twinspots Euschistospiza and Hypargos, but shown by means of molecular analysis to be closer to firefinches Lagonosticta (R.B. Payne, pers. comm.); with ‘a rather distinct colour pattern and its courtship display seems marginally closer to that of Hypargos, with which it is allopatric’ (Goodwin 1982). Y and X throat colour differ. Bill less robust, more pointed, than in Euschistospiza. P10 well developed, as in Euschistospiza and Spermophaga. Endemic.

Plate 18

Clytospiza monteiri (Hartlaub). Brown Twinspot. Se´ne ´gali brun.

(Opp. p. 267)

Pytelia monteiri Hartlaub, 1860. Proc. Zool. Soc. London. p. 111, pl. 161; Bembe, Congo district, Angola. Range and Status. Endemic resident, mesic savannas north and south of Congo basin. Nigeria, not uncommon around Serti and Abong; 1 record on Mambilla Plateau. Cameroon, scarce to frequent on Adamawa Plateau and south (in W) to border of forest zone and Yaounde´; west to Bamenda and Jauro Masali; frequent to common in upper valleys of Nun and Mbam rivers and around Tibati, Ngaounde´re´ and Banyo (R.B. Payne, pers. comm.); at Yaounde´, 24 birds seen in 21 months (Quantrill and Quantrill 1998). Chad, rare or overlooked; 3 records at about 8
300 N, others south to Bunda. Central African Republic, uncommon, from Chad border south to Bangui (where numerous), Luma I. and Mobaye on Ubangi R.; southwest to Carnot and Berbe´rati, east to Bangassou and Rafaı¨; single record in BaminguiBangoran region. Sudan, locally frequent in Bahr el Ghazal and Equatoria, north to Lado, Wau and Kafia Kingi. NE Zaı¨re, frequent to common in grasslands from 20
E to Uganda border. Uganda, frequent at 1000–1500 m from Wadelai south to Murchison Falls Nat. Park, Masindi, Budongo, Bugoma and shore of L. Victoria west to Masaka. Kenya, uncommon and local, from Busia and Mumias Districts to Siaya, Ng’iya and Akala. Gabon, confined to Nyanga District in S (Rand et al. 1959); recently, flock of 10, Moukalaba (Sargeant 1993). Congo, uncommon in Kouilou (Tchissanga, Pointe Noire), common in Brazzaville/Stanley Pool region. Angola, uncommon: Cabinda, Zaı¨re, Cuanza Norte, W Malanje; NE Lunda Norte; Cuanza Sul (Gabela and Quitondo). W Zaı¨re, frequent near Congo R. upstream to about Bolobo; S-central Zaı¨re, frequent in Kasai and W Kivu from NE Angola border to Lualaba R., north to near border of Haute Zaı¨re Prov.; common around Kananga (Luluabourg). Description. ADULT Y: top and sides of head to chin, sides of throat and sides of neck ashy grey, merging above with greyish brown mantle to rump and scapulars; uppertail-coverts crimson; spot on centre of throat red. Tail blackish brown. Underparts chestnut-brown; upper breast, sides and flanks with paired whitish spots, centre of lower breast and belly and undertailcoverts with whitish bars and broad white feather tips. Upperwing greyish brown. Axillaries and underwing-coverts greyish white, barred chestnut-brown. Bill black with steel-blue base; eyes brown, dull red or blood red; narrow bare eye-ring pale blue; legs reddish brown to greyish. ADULT X: like ad. Y, but throat

Clytospiza monteiri

?

?

?

spot white. SIZE (10 YY, 10 XX): wing, Y 56–60 (578), X 56–60 (581); tail, Y 47–53 (496), X 45–51 (479); bill, Y 135–15 (141), X 13–14 (138); tarsus, Y 165–175 (168), X 16–18 (171). WEIGHT: (Cameroon) Y (n ¼ 12) 135–16 (147), X (n ¼ 6) 14–16 (154), 1 unsexed 15, juv. (n ¼ 3) 130–133 (131); (Kenya) 2 YY 138, 147, 2 XX 144, 154, unsexed (n ¼ 7) 135–147 (141). IMMATURE: juv. differs from ad. X in having top of head and upperparts browner; uppertail-coverts more orange-red; chin to throat paler grey, no central spot; underparts plain rufous, without bars or spots (R. B. Payne, pers. comm.); later, underparts chestnut, belly with diffuse broad barring. NESTLING: down whitish; bill black, greyish below mandible; iris dark greyish brown; feet pale purplish grey; bilobed swelling at corner of mouth, whitish outside (A), yellow with 2 black spots inside; palate yellow with 5 black spots, lateral ones largest, posterior pair small; tongue flesh colour with dusky band across it; beneath tongue a blackish crescent (fig. p. 252).

348

ESTRILDIDAE

Genus Clytospiza Shelley

Single species of twinspot, related to twinspots Euschistospiza and Hypargos, but shown by means of molecular analysis to be closer to firefinches Lagonosticta (R.B. Payne, pers. comm.); with ‘a rather distinct colour pattern and its courtship display seems marginally closer to that of Hypargos, with which it is allopatric’ (Goodwin 1982). Y and X throat colour differ. Bill less robust, more pointed, than in Euschistospiza. P10 well developed, as in Euschistospiza and Spermophaga. Endemic.

Plate 18

Clytospiza monteiri (Hartlaub). Brown Twinspot. Se´ne ´gali brun.

(Opp. p. 267)

Pytelia monteiri Hartlaub, 1860. Proc. Zool. Soc. London. p. 111, pl. 161; Bembe, Congo district, Angola. Range and Status. Endemic resident, mesic savannas north and south of Congo basin. Nigeria, not uncommon around Serti and Abong; 1 record on Mambilla Plateau. Cameroon, scarce to frequent on Adamawa Plateau and south (in W) to border of forest zone and Yaounde´; west to Bamenda and Jauro Masali; frequent to common in upper valleys of Nun and Mbam rivers and around Tibati, Ngaounde´re´ and Banyo (R.B. Payne, pers. comm.); at Yaounde´, 24 birds seen in 21 months (Quantrill and Quantrill 1998). Chad, rare or overlooked; 3 records at about 8
300 N, others south to Bunda. Central African Republic, uncommon, from Chad border south to Bangui (where numerous), Luma I. and Mobaye on Ubangi R.; southwest to Carnot and Berbe´rati, east to Bangassou and Rafaı¨; single record in BaminguiBangoran region. Sudan, locally frequent in Bahr el Ghazal and Equatoria, north to Lado, Wau and Kafia Kingi. NE Zaı¨re, frequent to common in grasslands from 20
E to Uganda border. Uganda, frequent at 1000–1500 m from Wadelai south to Murchison Falls Nat. Park, Masindi, Budongo, Bugoma and shore of L. Victoria west to Masaka. Kenya, uncommon and local, from Busia and Mumias Districts to Siaya, Ng’iya and Akala. Gabon, confined to Nyanga District in S (Rand et al. 1959); recently, flock of 10, Moukalaba (Sargeant 1993). Congo, uncommon in Kouilou (Tchissanga, Pointe Noire), common in Brazzaville/Stanley Pool region. Angola, uncommon: Cabinda, Zaı¨re, Cuanza Norte, W Malanje; NE Lunda Norte; Cuanza Sul (Gabela and Quitondo). W Zaı¨re, frequent near Congo R. upstream to about Bolobo; S-central Zaı¨re, frequent in Kasai and W Kivu from NE Angola border to Lualaba R., north to near border of Haute Zaı¨re Prov.; common around Kananga (Luluabourg). Description. ADULT Y: top and sides of head to chin, sides of throat and sides of neck ashy grey, merging above with greyish brown mantle to rump and scapulars; uppertail-coverts crimson; spot on centre of throat red. Tail blackish brown. Underparts chestnut-brown; upper breast, sides and flanks with paired whitish spots, centre of lower breast and belly and undertailcoverts with whitish bars and broad white feather tips. Upperwing greyish brown. Axillaries and underwing-coverts greyish white, barred chestnut-brown. Bill black with steel-blue base; eyes brown, dull red or blood red; narrow bare eye-ring pale blue; legs reddish brown to greyish. ADULT X: like ad. Y, but throat

Clytospiza monteiri

?

?

?

spot white. SIZE (10 YY, 10 XX): wing, Y 56–60 (578), X 56–60 (581); tail, Y 47–53 (496), X 45–51 (479); bill, Y 135–15 (141), X 13–14 (138); tarsus, Y 165–175 (168), X 16–18 (171). WEIGHT: (Cameroon) Y (n ¼ 12) 135–16 (147), X (n ¼ 6) 14–16 (154), 1 unsexed 15, juv. (n ¼ 3) 130–133 (131); (Kenya) 2 YY 138, 147, 2 XX 144, 154, unsexed (n ¼ 7) 135–147 (141). IMMATURE: juv. differs from ad. X in having top of head and upperparts browner; uppertail-coverts more orange-red; chin to throat paler grey, no central spot; underparts plain rufous, without bars or spots (R. B. Payne, pers. comm.); later, underparts chestnut, belly with diffuse broad barring. NESTLING: down whitish; bill black, greyish below mandible; iris dark greyish brown; feet pale purplish grey; bilobed swelling at corner of mouth, whitish outside (A), yellow with 2 black spots inside; palate yellow with 5 black spots, lateral ones largest, posterior pair small; tongue flesh colour with dusky band across it; beneath tongue a blackish crescent (fig. p. 252).

Clytospiza monteiri

Field Characters. Length 11–13 cm. Unique among twinspots in having rich chestnut-brown underparts spotted and barred with white; head light grey, small red (Y) or white (X) patch in centre of throat; red uppertail-coverts at base of short, broad dusky tail conspicuous as bird flits into cover (Zimmerman et al. 1996). Juv. like X but duller and browner above, uppertail-coverts orange-red, chestnut below with only a hint of pale barring except on lower belly and undertail-coverts. Voice. Tape-recorded (104, B, PAY). Close contact call, which may lead into song, a harsh, nasal ‘vaa’ or ‘vay’, with whining quality (Payne and Payne 1994), intensified and uttered in longer series as distance contact call (Goodwin 1982). Song a bright, bustling medley of chatters, trills and chuckles at varying speeds, with rather hard quality, interspersed with single sparrow-like ‘chip’ and ‘cherp’, and with some whistled notes, high, descending ‘peeeea’ or ‘pee-pee-pee–pee-pew’; somewhat reminiscent of Atlantic Canary Serinus canaria but without the sweetness. Alarm a hard, Sylvia-like ‘tek’ (Goodwin 1982).

remain passive (Goodwin 1982). X solicits with quivering tail, and copulation follows. Display is identical with that of Peters’s Twinspot Hypargos niveoguttatus, q.v., except that Y Brown Twinspot’s movements are far less graceful and more jerky (Harrison and Dormer 1962). NEST: uses abandoned nests of estrildines; once in large mass of dry leaves, possibly old nest of seedcracker Pyrenestes or coucal Centropus, adding slight lining of matted hair collected from droppings of large carnivore, feathers, soft vegetable material and, often, pieces of snake skin. One sited on bough 2 m above water in swampy wood; 3 nests in old nests of Bronze Mannikins Spermestes cucullatus in single mango tree; one in old estrildid nest in grass on riverbank. In captivity builds own nest, roofed, globular or oval with large side entrance; made mainly of broad blades of grass and wheat, thickly lined with feathers; only Y collects material, carrying in bill up to 5 feathers at a time; if X is in nest being built, she takes feathers from Y and positions them (Neff 1975, Goodwin 1982). EGGS: 6; in captivity (Germany) 4–6. Pure white. SIZE: (n ¼ ?) 144–155  122–127. LAYING DATES: Nigeria, (juvs Feb); Cameroon, (oviduct egg, Oct; dependent young, Dec); Central African Republic, (gonads active Sept–Dec); Sudan, Sept; NE Zaı¨re, Aug– Dec; Uganda, (said to breed June–Oct: Mackworth-Praed and Grant 1960); Gabon, Apr. INCUBATION: in captivity, by X; Y takes feathers and pieces of green grass into nest during incubation and brooding periods; large nest entrance always screened or closed by a feather. Period: 13 days (in captivity, Germany).

General Habits. Inhabits moist bushy grasslands, tall, rank grass and thickets in mature woodland and gardens, and riverine bushes; tangled vegetation in scattered bush and forest patches (Uganda, from suburbs of Kampala to lake shore); overgrown cultivation; Lantana scrub. Keeps low down in vegetation. In pairs and family parties; once a flock of 10. Forages on ground, on patches of bare earth amongst tall grass, once on lichen-covered boulders, once on roadside. On ground moves in long quick hops, in horizontal stance. Shy. Y seems to sing only during courtship display. Pair commonly allopreens. Food. Small grass seeds and arthropods, particularly termites; once a spider. In captivity eats millet and canary seed and, when rearing young, ant pupae, whiteworms and half-ripe grass seeds (Goodwin 1982). Breeding Habits. Solitary nester; evidently monogamous in captivity. In courtship display, Y takes feather or grass blade in bill and sings whilst making long hops around or half around X on ground, his tail spread and angled towards her, bobbing, his head alternately angled up at c. 70
and bowed towards X at angle of c. 30
. He starts, crouching close to ground with head elevated, slowly lowers head then throws it up quickly whilst hopping ‘halfan-inch off the ground’ (Harrison and Dormer 1962). X may perform same movements, without anything in bill and without singing, synchronously with Y, or she may

DEVELOPMENT AND CARE OF YOUNG: parents do not remove nestlings’ faeces from nest, which becomes very dirty by time of fledging. Period: 19–21 days (in captivity, Germany, Neff 1975, 1977). Young recently out of nest hide in thick leafy cover and attract parents to them with loud ‘eek, eek, eek’ calls. Whitish flange at corner of mouth becomes smaller and less conspicuous (B). BREEDING SUCCESS/SURVIVAL: thought to be brood parasitized by Cameroon Indigobird Vidua camerunensis, q.v.

Key References Goodwin, D. (1982), Harrison, C.J.O. and Dormer, B.P. (1962), Neff, R. (1975, 1977).

349

350

ESTRILDIDAE

Genus Pytilia Swainson

A distinctive, endemic genus of 5 estrildid finches in wooded grassland, YY mainly grey or olive-green, face red (3 spp.) or grey or yellow, rump and tail red (4 spp.) or olive, wings red, orange, yellow or green, flanks barred; bill red (3 spp.) or black. XX like YY but not red-faced. Eat grass seeds; make loosely built grass ball nest in bush. Parasitized by paradise whydahs Vidua interjecta, V. togoensis, V. orientalis and V. paradisea. The 5 species treated as 2 superspecies by Sibley and Monroe (1990), phoenicoptera/lineata/afra and melba/hypogrammica. In appearance phoenicoptera/lineata, hypogrammica, afra and melba from a series, in which the closest relationship would seem to be between the grey hypogrammica and green afra; however, molecular analysis shows that phoenicoptera and hypogrammica are very closely related, lineata not quite so close and afra less close again, the 4 forming a sister clade to melba (R. B. Payne, pers. comm.). We therefore treat them as a superspecies (hypogrammica/phoenicoptera/lineata) and 2 independent species. A pytilia lacking red in plumage, known from Djibouti and N Cape, may be a mutant P. melba or a new taxon (see under that species); see also TAXONOMIC NOTES under P. hypogrammica and P. lineata.

Pytilia phoenicoptera superspecies

1,2 1 2

3

2 1

1,2

1,2

1,2 1,2?

1 P. phoenicoptera 2 P. hypogrammica 3 P. lineata

Plate 17

Pytilia hypogrammica Sharpe. Yellow-winged Pytilia. Beaumarquet `a ailes jaunes.

(Opp. p. 266)

Pytelia hypogrammica Sharpe, 1870. Ibis, p. 56; Fantee, Gold Coast. Forms a superspecies with P. phoenicoptera and P. lineata. Range and Status. Endemic resident, Guinea-Bissau to W Central African Republic. Guinea-Bissau, Dulombi-Bafata (R.B. Payne, pers. comm.). Guinea, uncommon in Kindia (Mambia, Foulayah, Kolente´), Dalaba, and Beyla/Leleko areas. Sierra Leone, records at Kabala and Musaia. Liberia, rare: small flock Wologizi, Dec 1984 (Gatter 1997). Ivory Coast, frequent in Marahoue´ (Maroue´) Nat. Park (Bull. Afr. Bird Club 7, 2000, 147) and Comoe´ Nat. Park (Salewski

1997) with several records in between; south to Toumodi. Burkina Faso/Benin, uncommon to rare, Arli/Pendjari Nat. Parks (Green and Sayer 1979). Ghana, uncommon to rare, local; in N (Gambaga), mid latitudes and SE (Akwapim Hills, Opintin Forest, Teshie, Abokobi, Accra Plains). Togo, uncommon in N (Binaparba, Kpayando, Ketao, Fazao) and centre (Ebe´va, Edifou). Nigeria, uncommon: Kishi and Kaduna/Zaria areas, Abuchi (Abugi), and from

350

ESTRILDIDAE

Genus Pytilia Swainson

A distinctive, endemic genus of 5 estrildid finches in wooded grassland, YY mainly grey or olive-green, face red (3 spp.) or grey or yellow, rump and tail red (4 spp.) or olive, wings red, orange, yellow or green, flanks barred; bill red (3 spp.) or black. XX like YY but not red-faced. Eat grass seeds; make loosely built grass ball nest in bush. Parasitized by paradise whydahs Vidua interjecta, V. togoensis, V. orientalis and V. paradisea. The 5 species treated as 2 superspecies by Sibley and Monroe (1990), phoenicoptera/lineata/afra and melba/hypogrammica. In appearance phoenicoptera/lineata, hypogrammica, afra and melba from a series, in which the closest relationship would seem to be between the grey hypogrammica and green afra; however, molecular analysis shows that phoenicoptera and hypogrammica are very closely related, lineata not quite so close and afra less close again, the 4 forming a sister clade to melba (R. B. Payne, pers. comm.). We therefore treat them as a superspecies (hypogrammica/phoenicoptera/lineata) and 2 independent species. A pytilia lacking red in plumage, known from Djibouti and N Cape, may be a mutant P. melba or a new taxon (see under that species); see also TAXONOMIC NOTES under P. hypogrammica and P. lineata.

Pytilia phoenicoptera superspecies

1,2 1 2

3

2 1

1,2

1,2

1,2 1,2?

1 P. phoenicoptera 2 P. hypogrammica 3 P. lineata

Plate 17

Pytilia hypogrammica Sharpe. Yellow-winged Pytilia. Beaumarquet `a ailes jaunes.

(Opp. p. 266)

Pytelia hypogrammica Sharpe, 1870. Ibis, p. 56; Fantee, Gold Coast. Forms a superspecies with P. phoenicoptera and P. lineata. Range and Status. Endemic resident, Guinea-Bissau to W Central African Republic. Guinea-Bissau, Dulombi-Bafata (R.B. Payne, pers. comm.). Guinea, uncommon in Kindia (Mambia, Foulayah, Kolente´), Dalaba, and Beyla/Leleko areas. Sierra Leone, records at Kabala and Musaia. Liberia, rare: small flock Wologizi, Dec 1984 (Gatter 1997). Ivory Coast, frequent in Marahoue´ (Maroue´) Nat. Park (Bull. Afr. Bird Club 7, 2000, 147) and Comoe´ Nat. Park (Salewski

1997) with several records in between; south to Toumodi. Burkina Faso/Benin, uncommon to rare, Arli/Pendjari Nat. Parks (Green and Sayer 1979). Ghana, uncommon to rare, local; in N (Gambaga), mid latitudes and SE (Akwapim Hills, Opintin Forest, Teshie, Abokobi, Accra Plains). Togo, uncommon in N (Binaparba, Kpayando, Ketao, Fazao) and centre (Ebe´va, Edifou). Nigeria, uncommon: Kishi and Kaduna/Zaria areas, Abuchi (Abugi), and from

Pytilia hypogrammica

Pytilia hypogrammica

?

TAXONOMIC NOTE: a Y with orange-red wings from Bunda, Chad, was described in 1906 as Pytilia lopezi; a Nigerian Y had red wings and some breast feathers tipped red, and a X had red feathers in supercilium (Serle 1957). 6 young bred from a pair of red-winged ‘lopezi’ in captivity all had red wings, so red-winged form may be a colour morph or a valid subspecies.

upper Assop (Plateau State) and Wamba to Enugu, Okigwi Hills and Serti region; frequent around Enugu, east to Abakaliki; old record from Abuochichi ( ¼ Aboutschi, west of Obudu). Cameroon, frequent between Tibati and Ngaounde´re´ and elsewhere on Adamawa Plateau; from 10
E to Chad border. S Chad and NW Central African Republic, known from 5 or more localities in Chari, upper Logone and upper Man Dadji valleys, Ouham R. and Bozoum; W Bamingui-Bangoran Nat. Park; Kaga Bandoro; said to occur in Lobaye Pre´fecture. Description. ADULT Y: forehead, forecrown and above eye to earcoverts, cheeks and chin bright red or scarlet, enclosing grey lores; hindcrown to rump and scapulars brownish grey. Lower rump and uppertail-coverts bright red. Tail feathers dark grey-brown, upperside of T1 and of outer webs of T2–T5 washed crimson. Throat and sides of neck to upper breast brownish grey, upper breast with fine dark brown bars; rest of underparts brownish grey, narrowly barred whitish and dark brown; undertail-coverts dark grey, tipped white. Upperwing grey-brown; primaries, secondaries, primary coverts and alula edged olive-yellow; greater, median and lesser coverts broadly fringed olive-yellow. Axillaries and underwing-coverts whitish, spotted dusky grey. Bill dark grey; eyes dark red to scarlet; legs pale brown or grey-brown. ADULT X: lacks red on face; top of head uniform with upperparts, side of head slightly paler; upperparts browner than in ad. Y. Underparts greyish olive, paler on chin, throat and centre of belly; barred buffish white, finely so from chin to upper breast, more broadly and prominently across lower breast and belly; undertail-coverts broadly tipped buffish white. Wings, tail and bare parts as ad. Y. SIZE (10 YY, 10 XX): Y 56–60 (583), X 54–58 (564); tail, Y 34–41 (370), X 33–40 (362); bill, Y 125–135 (129), X 12–135 (128); tarsus, Y 145–16 (154), X 15–155 (151). WEIGHT: 1 Y 143. IMMATURE: juv. similar to ad. X but upperparts browner, yellow edges to wing feathers narrower; underparts barred throughout. NESTLING: skin dark, with long pale down (A).

Field Characters. Length 125–13 cm. A pytilia of W savannas, not overlapping Orange-winged P. afra. Grey, with fine pale bars on flanks and belly; told from Redwinged Pytilia P. phoenicoptera by red face, black outer tail feathers, yellow or orange-red wings, from Green-winged Pytilia P. melba by overall grey colour; X lacks red face and is light brown below barred dull white, told from Greenwinged by grey-brown upperparts and black bill, from Green-winged and Red-winged by yellow wings. Juv. browner than X, barring on underparts less distinct, more like juvs of the other 2 species, but still has some yellow in wings, and no hint of green on upperparts. Voice. Tape-recorded (104, B, GUT, PAY). Call a plaintive ‘tsee’, ‘tsew’ or ‘tseea’ (sonagram in Guttinger and Nicolai 1973); courtship song of Y described as ‘vee-vee-vee’. Vocalizations said to be very like those of Red-winged Pytilia (Nicolai 1977). General Habits. Inhabits mature, undisturbed S-guinean savanna woodland and derived savanna bushland; edges of gallery forest, coastal thickets and dense woody growth around bases of inselbergs (Ghana); abandoned cultivation and orchard bush, often near streams (Nigeria). Singly, in pairs; sometimes in parties of up to 5. Forages on ground, seeking grass seeds, often in company of other estridines; but often perches in topmost branches of trees and shrubs. Flight undulating and very fast. When disturbed by person commonly flies to some distance, often out of sight (Goodwin 1982). Food. Grass seeds and insects. Of 19 stomachs, seeds in 16 and insects in 3 (Serle 1957). In captivity readily eats millet seed, canary seed and lettuce seed; very fond of ant pupae and fed hatchlings on them entirely for first few days, later giving them dry millet seeds (Eckl 1974). Breeding Habits. Solitary nester; presumably monogamous; not conspicuously territorial. In courtship display,

351

352

ESTRILDIDAE

on ground, Y at side of X bows repeatedly with tail angled, or hops around her bowing and uttering ‘vee-vee-vee’ calls. Once Y of pair perched in bush, sang in erect posture, bowed to X who solicited with quivering tail, and they copulated (Eckl 1974). Captive Y and X may bow to each other and Y, with ‘nesting symbol’ in bill, jumps up and down on perch, letting go of it momentarily, and throwing head up as he does so (Burkard 1964, Goodwin 1982). NEST: domed oval or globular structure with circular opening at one side; very loosely constructed, with 3 clearly defined layers: outer envelope of broad grass strips, middle layer of grass stems, and lining of flowering grass tops and a few feathers, sometimes of guineafowl; 120 mm from top to bottom, entrance 50 wide. Sited 05–4 m above ground in fork of small shrub in recently burnt shrubby savanna. EGGS: 3–4. White, ovate with smooth, slightly glossy surface. SIZE: (n ¼ 3, 1 clutch) 148–155  115–119 (152 

117) (a 4th egg in this clutch, 172  119, was of brood parasite Exclamatory Paradise-Whydah Vidua interjecta.) LAYING DATES: Nigeria, Oct–Jan (Nicolai 1977) (and enlarged gonads, Mar). INCUBATION: in captivity, Y and X incubate by day in turns. X sat close and when flushed flew straight away out of sight; on return 90 min later she flew directly to nest. Period: 12–14 days (in captivity). BREEDING SUCCESS/SURVIVAL: brood-parasitized by Vidua interjecta; 3 parasitized nests, SE Nigeria, held 3, 4 and 4 pytilia eggs and respectively 1, 2 and 3 V. interjecta eggs; probably parasitized also by Togo Paradise-Whydah V. togoensis. In Nigeria many nests and young destroyed by man-made grass fires (Nicolai 1977). Key References Eckl, G. (1974), Goodwin, D. (1982), Gu ¨ ttinr, H.R. and Nicolai, J. (1973), Nicolai, J. (1977), Serle, W. (1957).

Plate 17

Pytilia phoenicoptera Swainson. Red-winged Pytilia. Beaumarquet aurore.

(Opp. p. 266)

Pytilia phoenicoptera Swainson, 1837. Birds West Africa, 1, p. 203, pl. 16; West Africa (¼ Gambia, Hartlaub, 1857, Syst. Orn. Westafrika, p. 145). Forms a superspecies with P. hypogrammica and P. lineata. Range and Status. Endemic resident and probably partial migrant, Senegal to N Uganda. Senegal, uncommon, near Gambian border and in SE. Gambia, widespread, uncommon, but locally frequent in large forest parks, particularly in Kiang West; in Central River Division, only south of river, near Kudang; 3 times as common as Green-winged Pytilia P. melba. Guinea-Bissau, old records near coast; inland at Farim (Rodwell 1996). Guinea, Labe´, Le´louma, Dalaba (R.B. Payne, pers. comm.); uncommon in Kindia area (Mambia, Foulayah: Demey 1995). Sierra Leone, recorded (Dowsett 1993). Mali, uncommon in Bamako and Koulikoro areas. Ivory Coast, frequent throughout Comoe´ Nat. Park (Salewski 1997), rare west to Odienne´. Burkina Faso, uncommon north to Ouaga (Ouagadougou); Burkina Faso/Benin, frequent in Arli/Pendjari Nat. Parks. Ghana, uncommon in NE (Tumu, Sugu, Gambaga, Tamale); old record at Accra (Bannerman 1949). Togo, not uncommon in N (Mare aux Lions, Kpayando); occurs south to Ayagba (7
46’N). Benin, frequent in Pendjari Nat. Park, south to Be´te´rou area where rare. Niger, frequent in ‘W’ Nat. Park in all months except Sept–Oct, on Niger, Mekrou and Tapoa rivers. Nigeria, uncommon to frequent, from Kainji Lake Nat. Park east through middle latitudes including Kaduna and Zaria (where uncommon) and Yankari Nat. Park (where much commoner than P. melba) and Jos Plateau (sparse), southeast to Yola, north to Sokoto, Falgore, Kano, Kari and Potiskum, south to Kafanchan. Cameroon, local on Benue Plain (Touroua area) and E Adamawa Plateau (Tello area); north to 11
N, south to 4
N. Central African Republic, common throughout year in Bamingui-Bangoran Nat. Park; uncommon in ManovoGounda-Saint Floris Nat. Park and Vakaga region; occurs in Lobaye Pre´fecture. Sudan, uncommon in S: Tambura,

Pytilia phoenicoptera

Wau, Lado, Kajo Kaji. NE Zaı¨re, frequent: Garamba Nat. Park, Faradje, Aba, also Mahagi Port. Uganda, uncommon at 600–1200 m in West Nile, Acholi, Kidepo Valley Nat. Park, Murchison Falls Nat. Park, Lango and Teso. Description. P. p. emini Hartert: Cameroon to Uganda. ADULT Y: head and upperparts dark grey; lower rump and uppertail-coverts bright crimson. Tail feathers dark grey-brown, upperside of T1 and of outer webs of T2–T5 washed crimson. Chin to upper breast

352

ESTRILDIDAE

on ground, Y at side of X bows repeatedly with tail angled, or hops around her bowing and uttering ‘vee-vee-vee’ calls. Once Y of pair perched in bush, sang in erect posture, bowed to X who solicited with quivering tail, and they copulated (Eckl 1974). Captive Y and X may bow to each other and Y, with ‘nesting symbol’ in bill, jumps up and down on perch, letting go of it momentarily, and throwing head up as he does so (Burkard 1964, Goodwin 1982). NEST: domed oval or globular structure with circular opening at one side; very loosely constructed, with 3 clearly defined layers: outer envelope of broad grass strips, middle layer of grass stems, and lining of flowering grass tops and a few feathers, sometimes of guineafowl; 120 mm from top to bottom, entrance 50 wide. Sited 05–4 m above ground in fork of small shrub in recently burnt shrubby savanna. EGGS: 3–4. White, ovate with smooth, slightly glossy surface. SIZE: (n ¼ 3, 1 clutch) 148–155  115–119 (152 

117) (a 4th egg in this clutch, 172  119, was of brood parasite Exclamatory Paradise-Whydah Vidua interjecta.) LAYING DATES: Nigeria, Oct–Jan (Nicolai 1977) (and enlarged gonads, Mar). INCUBATION: in captivity, Y and X incubate by day in turns. X sat close and when flushed flew straight away out of sight; on return 90 min later she flew directly to nest. Period: 12–14 days (in captivity). BREEDING SUCCESS/SURVIVAL: brood-parasitized by Vidua interjecta; 3 parasitized nests, SE Nigeria, held 3, 4 and 4 pytilia eggs and respectively 1, 2 and 3 V. interjecta eggs; probably parasitized also by Togo Paradise-Whydah V. togoensis. In Nigeria many nests and young destroyed by man-made grass fires (Nicolai 1977). Key References Eckl, G. (1974), Goodwin, D. (1982), Gu ¨ ttinr, H.R. and Nicolai, J. (1973), Nicolai, J. (1977), Serle, W. (1957).

Plate 17

Pytilia phoenicoptera Swainson. Red-winged Pytilia. Beaumarquet aurore.

(Opp. p. 266)

Pytilia phoenicoptera Swainson, 1837. Birds West Africa, 1, p. 203, pl. 16; West Africa (¼ Gambia, Hartlaub, 1857, Syst. Orn. Westafrika, p. 145). Forms a superspecies with P. hypogrammica and P. lineata. Range and Status. Endemic resident and probably partial migrant, Senegal to N Uganda. Senegal, uncommon, near Gambian border and in SE. Gambia, widespread, uncommon, but locally frequent in large forest parks, particularly in Kiang West; in Central River Division, only south of river, near Kudang; 3 times as common as Green-winged Pytilia P. melba. Guinea-Bissau, old records near coast; inland at Farim (Rodwell 1996). Guinea, Labe´, Le´louma, Dalaba (R.B. Payne, pers. comm.); uncommon in Kindia area (Mambia, Foulayah: Demey 1995). Sierra Leone, recorded (Dowsett 1993). Mali, uncommon in Bamako and Koulikoro areas. Ivory Coast, frequent throughout Comoe´ Nat. Park (Salewski 1997), rare west to Odienne´. Burkina Faso, uncommon north to Ouaga (Ouagadougou); Burkina Faso/Benin, frequent in Arli/Pendjari Nat. Parks. Ghana, uncommon in NE (Tumu, Sugu, Gambaga, Tamale); old record at Accra (Bannerman 1949). Togo, not uncommon in N (Mare aux Lions, Kpayando); occurs south to Ayagba (7
46’N). Benin, frequent in Pendjari Nat. Park, south to Be´te´rou area where rare. Niger, frequent in ‘W’ Nat. Park in all months except Sept–Oct, on Niger, Mekrou and Tapoa rivers. Nigeria, uncommon to frequent, from Kainji Lake Nat. Park east through middle latitudes including Kaduna and Zaria (where uncommon) and Yankari Nat. Park (where much commoner than P. melba) and Jos Plateau (sparse), southeast to Yola, north to Sokoto, Falgore, Kano, Kari and Potiskum, south to Kafanchan. Cameroon, local on Benue Plain (Touroua area) and E Adamawa Plateau (Tello area); north to 11
N, south to 4
N. Central African Republic, common throughout year in Bamingui-Bangoran Nat. Park; uncommon in ManovoGounda-Saint Floris Nat. Park and Vakaga region; occurs in Lobaye Pre´fecture. Sudan, uncommon in S: Tambura,

Pytilia phoenicoptera

Wau, Lado, Kajo Kaji. NE Zaı¨re, frequent: Garamba Nat. Park, Faradje, Aba, also Mahagi Port. Uganda, uncommon at 600–1200 m in West Nile, Acholi, Kidepo Valley Nat. Park, Murchison Falls Nat. Park, Lango and Teso. Description. P. p. emini Hartert: Cameroon to Uganda. ADULT Y: head and upperparts dark grey; lower rump and uppertail-coverts bright crimson. Tail feathers dark grey-brown, upperside of T1 and of outer webs of T2–T5 washed crimson. Chin to upper breast

Pytilia phoenicoptera and flanks dark grey, concolorous with head; lower breast and belly brownish grey with fine white barring, broadest on centre of belly; undertail-coverts dark grey, tinged red and tipped white. Remiges dark grey-brown; outer primaries narrowly edged rufous-cinnamon, inner primaries and outer secondaries edged bright red, outer secondaries and tertials edged rufous-cinnamon; primary coverts and alula dark grey-brown, narrowly edged cinnamon; greater, median and lesser coverts red. Axillaries and underwing-coverts whitish, spotted dusky grey; inner borders of undersides of flight feathers greyish white. Bill black; eyes red; legs greyish brown to flesh. ADULT X: similar to ad. Y but browner grey above; paler, more olive-grey below, with narrow whitish barring more prominent, extending higher on breast. SIZE (8 YY, 5 XX): wing, Y 57–59 (577), X 57–60 (580); tail, Y 35–39 (373), X 33–39 (362); bill, Y 125–15 (133), X 12–14 (129); tarsus, Y 15–165 (158), X 145–15 (147). IMMATURE: juv. has upperparts, head to upper breast and flanks dark greyish brown; lower breast and belly buffish white, barred brown at sides; undertail-coverts grey-brown, broadly tipped buffish white; long uppertail-coverts and outer edges of tail feathers tinged red; upperwing grey-brown, primaries and secondaries edged pale buffy brown. NESTLING: skin black, down grey; bill black; 2 lobes at corner of mouth, bluish white, each with black spot on inner side; roof of mouth whitish anteriorly, becoming bright rose, with large light bluish or violet area at each side of internal nares; no black spots on palate; tongue rosy at base, flesh at tip (fig. p. 252). Eyes brownish grey; legs greyish pink. P. p. phoenicoptera Swainson: Senegal to Nigeria. Y slightly browner grey than emini; white barring on underparts broader, extending to upper breast. X paler below than emini, with more pronounced breast barring. WEIGHT: (Ghana) unsexed (n ¼ 5) 137–149 (145); (Nigeria) Y (n ¼ 5) 148–152 (150), X (n ¼ 4) 111–153 (138), unsexed (n ¼ 8) within range 111–153, av. of all 17, 145 (Fry 1971).

Field Characters. Length 125–13 cm. Grey with fine pale barring on belly, like Yellow-winged Pytilia P. hypogrammica which it overlaps in W Africa; differs in grey face, red wings, and wholly red tail (outer feathers black on Yellowwinged). X same colours as Yellow-winged except for red wing; brown juvs even more alike, but both show some diagnostic wing colour. Very like Red-billed Pytilia. P. lineata of Ethiopia, except for black bill, but their ranges do not overlap. Voice. Tape-recorded (104, B, PAY). Song from captive bird (PAY) a rather disjointed medley of whistles and scratchy notes; frequent is a pure up-slurred whistle, ‘toowee’, sometimes 3-syllable ‘too-wee-oo’, and a harsh, descending ‘jaaaa’ or ‘jit-jaaa’, which can be combined with whistle, ‘tee-tloo-waaa’; other notes include burry ‘tuweea’ and whistled ‘preep’; song punctuated by short, hard ‘jit’, ‘chut’ or ‘jrrt’. Transcriptions of songs in Goodwin (1982) suggest individual and/or geographic variation. Contact call said to be fluting or whistling in tone; another call (function?) ventriloquial ‘eek’ or ‘lkyerk’; during copulation a soft ‘geegeegeegee’ (hard ‘g’) (Goodwin 1982). Sonagram of call in Gu ¨ ttinger and Nicolai (1973). General Habits. Inhabits mature, dry savanna woodlands with large trees and bushy thickets; also moister coastal areas in Gambia and Guinea-Bissau; in Nigeria often around villages and in cultivation clearings; in Zaı¨re occurs also on roadsides and in cleared and burnt places. Usually in pairs;

sometimes singly, or in flocks of c. 4, once 8. Unobtrusive. Forages on ground, sometimes mixing with waxbills, mannikins and weavers but far more retiring than them, keeping close to bushy cover. When not feeding spends much time in bushes and trees; often perches conspicuously on dead branches high up in emergent tree. Regularly drinks towards sunset at puddles in woods, at roadsides and in laterite quarries, where occurs well out in the open. When alarmed, crouches motionless with tail raised. Resident, but in Gambia more commonly seen in rains (July–Oct). May be migratory in Nigeria; at Zaria, 17 birds caught only from mid May to late June, in netting study from Oct to July (Fry 1971). Evidently a breeding visitor from mid-Oct to Jan in Uele, Zaı¨re, but further north, in Garamba Nat. Park, also occurs in June (Chapin 1954). Food. Grass seeds and insects. Of 8 stomachs, 4 contained only grass seeds and 4 only worker (un-winged) termites (Chapin 1954). Breeding birds in captivity need abundant supplies of ant pupae to feed to their young. Breeding Habits. Solitary nester; presumably monogamous; not conspicuously territorial. In captivity very unaggressive. In display, Y hops around X, bowing and bobbing up and down, holding piece of nesting material in bill and uttering penetrating ‘tseet, tseet’ call. Apparently also hops up and down on perch, just letting go with feet, like Yellow-winged Pytilia (q.v.). NEST: bulky, globular, made almost entirely of flowering or seeding heads of grasses 30–120 long and put together loosely and irregularly, the seeds not stripped off, with a few old grass blades and a few feathers, once of francolin Francolinus bicalcaratus; with round entrance at one side, diam. 35; foundation of loosely accumulated grass blades and stems and herb flower heads; 2 nests thickly lined with feathers. Sited in middle or near top of bush, between 075 and 2 m above ground. EGGS: 2–4 (usually 3–4). Pure white; rounded ovate, slightly glossy, or matt. SIZE: (n ¼ 9) 150–167  120–132 (156  124). LAYING DATES: Mali, (carrying nest material Dec, Jan); Gambia, Sept; Nigeria, Nov, Dec, Feb (and oviduct egg Sept); Togo, (young 2 days out of nest, early Mar); Zaı¨re, (oviduct egg Nov; nestlings mid Nov). INCUBATION: by Y and X, taking turns; Y flying in to nest to relieve X often carries feather in its bill. Incubating bird sits close; when flushed by person, either flies to nearby tree or disappears for 30 min. Period: 12–13 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: young brooded by both parents, taking turns; fed on seeds (Bannerman 1949); nestling begging for food opens mouth wide, raises base of tongue unusually high and wags head from side to side ‘as though strangling’ (Chapin 1954). Nestlings’ faeces not removed by parents but accumulate on nest floor. Nestling period: c. 21 days (in captivity). BREEDING SUCCESS/SURVIVAL: brood-parasitized by Exclamatory Paradise-Whydah Vidua interjecta. Key References Bannerman, D.A. (1949), Clement, P. et al. (1993), Cotterel, R. (1960), Goodwin, D. (1982).

353

354

ESTRILDIDAE

Plate 17

Pytilia lineata Heuglin. Red-billed Pytilia. Beaumarquet a` bec rouge.

(Opp. p. 266)

Pytelia lineata Heuglin, 1863. J. Orn., 11, p. 17; Dembea, Abyssinia. Forms a superspecies with P. hypogrammica and P. phoenicoptera. Range and Status. Endemic resident, Ethiopia and E Sudan. Rare in Sudan: 2–3 old records close to Ethiopian border between 8
and 12
N. Ethiopia, common to abundant in W and in central highlands, known from c. 30 localities with up to 26 birds seen in a day (J. S. Ash, pers. comm.); rare in NW, where only 1 encountered in 9 years (at Waldagadana, 1675 m: Cheesman and Sclater 1935–1936). Description. ADULT Y: top of head and upperparts dark brownish grey; lower rump and uppertail-coverts crimson. Tail feathers blackish brown, upperside of T1 and of outer webs of T2–T5 washed crimson. Cheeks, ear-coverts, sides of neck and chin to upper breast dark grey; lower breast, belly and flanks barred dark brownish grey and buffy white; undertail-coverts dark brownish grey, tipped whitish. Upperwing feathers dark grey-brown, remiges edged crimson; primary coverts narrowly edged crimson, outer webs of greater coverts and tips of median and lesser coverts crimson. Axillaries and underwing-coverts buffy white, barred dusky grey. Bill red; eyes brown. ADULT X: similar to ad. Y, but browner grey above; slightly paler below, whitish barring below extending higher on breast. SIZE (3 YY): wing, Y 58–60 (593), 1 unsexed 61; tail, Y 38–39 (387); bill, Y 125–135 (132); tarsus, Y 16. WEIGHT: unsexed (n ¼ 99) 110– 168 (143) (J.S. Ash, pers. comm.). IMMATURE: juv. has upperparts, head to upper breast and flanks dark brown; lower breast and belly buffy white, barred brown at sides; undertail-coverts dark brown, tipped buffish white; uppertail-coverts and edges of tail feathers tinged red; upperwing dark grey-brown, primaries and secondaries edged buffish brown. Bill greyish or brown, becoming pinkish before turning red. TAXONOMIC NOTE: usually treated as a race of P. phoenicoptera, but in captivity lineata and phoenicoptera do not interbreed and are in ‘rigid reproductive isolation’, behaving as different species even though voices and displays are almost identical; as shown experimentally, the strikingly different bill colour functions as an isolating mechanism (Nicolai 1968). Recognized as a species distinct from P. phoenicoptera by Sibley and Monroe (1990) but not by Dowsett and Dowsett-Lemaire (1993a). Shown to be slightly less closely related to phoenicoptera and hypogrammica than the 2 are to each other (R. B. Payne, pers. comm.). Ranges of phoenicoptera and lineata are c. 375 km apart (Nikolaus 1987).

Field Characters. Length 125–13 cm. Grey and red; like Red-winged Pytilia P. phoenicoptera except for red bill, but identification not a problem since their ranges do not overlap. Readily separated by lack of green in plumage from all plumages, including juvs, of sympatric Orange-winged

Pytilia lineata

and Green-winged Pytilias P. afra and P. melba, from YY also by grey, not red, face. Voice. Not tape-recorded. Said to be like Red-winged Pytilia (Nicolai 1968). Call in flight and when disturbed, ‘chwick’ (J.S. Ash, pers. comm.). General Habits. Inhabits open wooded grassland and open bush; almost entirely confined to Combretum-Terminalia woodland; occurs only occasionally in other woodlands and Acacia bush (J. S. Ash, pers. comm.). Food. In captivity, seeds, mealworms and mosquito larvae. Breeding Habits. Not known in wild except for a nest in May; in captivity, Y display and nest same as in Redwinged Pytilia P. phoenicoptera (though details not given: Nicolai 1968, Goodwin 1982). Key References

Goodwin, D. (1982), Nicolai, J. (1968).

Plate 17

Pytilia melba (Linnaeus). Green-winged Pytilia; Melba Finch. Beaumarquet melba.

(Opp. p. 266)

Fringilla Melba Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 180; China (¼Angola, Zedlitz, 1916, J. Orn., 64, p. 31; restricted to Luanda by Clancey, 1962, Bull. Br. Orn. Club, 82, p. 4). Range and Status. Endemic resident. Mauritania, uncommon, >20 localities north to 17
N; Senegal, frequent, widespread; absent south of 13
N. Gambia, uncommon

and local: Kiang; north of Kaur, mainly on N side of river; Basse; east of Farfenni. Guinea-Bissau, records in W. Guinea, single old record in N. Mali, widespread but

354

ESTRILDIDAE

Plate 17

Pytilia lineata Heuglin. Red-billed Pytilia. Beaumarquet a` bec rouge.

(Opp. p. 266)

Pytelia lineata Heuglin, 1863. J. Orn., 11, p. 17; Dembea, Abyssinia. Forms a superspecies with P. hypogrammica and P. phoenicoptera. Range and Status. Endemic resident, Ethiopia and E Sudan. Rare in Sudan: 2–3 old records close to Ethiopian border between 8
and 12
N. Ethiopia, common to abundant in W and in central highlands, known from c. 30 localities with up to 26 birds seen in a day (J. S. Ash, pers. comm.); rare in NW, where only 1 encountered in 9 years (at Waldagadana, 1675 m: Cheesman and Sclater 1935–1936). Description. ADULT Y: top of head and upperparts dark brownish grey; lower rump and uppertail-coverts crimson. Tail feathers blackish brown, upperside of T1 and of outer webs of T2–T5 washed crimson. Cheeks, ear-coverts, sides of neck and chin to upper breast dark grey; lower breast, belly and flanks barred dark brownish grey and buffy white; undertail-coverts dark brownish grey, tipped whitish. Upperwing feathers dark grey-brown, remiges edged crimson; primary coverts narrowly edged crimson, outer webs of greater coverts and tips of median and lesser coverts crimson. Axillaries and underwing-coverts buffy white, barred dusky grey. Bill red; eyes brown. ADULT X: similar to ad. Y, but browner grey above; slightly paler below, whitish barring below extending higher on breast. SIZE (3 YY): wing, Y 58–60 (593), 1 unsexed 61; tail, Y 38–39 (387); bill, Y 125–135 (132); tarsus, Y 16. WEIGHT: unsexed (n ¼ 99) 110– 168 (143) (J.S. Ash, pers. comm.). IMMATURE: juv. has upperparts, head to upper breast and flanks dark brown; lower breast and belly buffy white, barred brown at sides; undertail-coverts dark brown, tipped buffish white; uppertail-coverts and edges of tail feathers tinged red; upperwing dark grey-brown, primaries and secondaries edged buffish brown. Bill greyish or brown, becoming pinkish before turning red. TAXONOMIC NOTE: usually treated as a race of P. phoenicoptera, but in captivity lineata and phoenicoptera do not interbreed and are in ‘rigid reproductive isolation’, behaving as different species even though voices and displays are almost identical; as shown experimentally, the strikingly different bill colour functions as an isolating mechanism (Nicolai 1968). Recognized as a species distinct from P. phoenicoptera by Sibley and Monroe (1990) but not by Dowsett and Dowsett-Lemaire (1993a). Shown to be slightly less closely related to phoenicoptera and hypogrammica than the 2 are to each other (R. B. Payne, pers. comm.). Ranges of phoenicoptera and lineata are c. 375 km apart (Nikolaus 1987).

Field Characters. Length 125–13 cm. Grey and red; like Red-winged Pytilia P. phoenicoptera except for red bill, but identification not a problem since their ranges do not overlap. Readily separated by lack of green in plumage from all plumages, including juvs, of sympatric Orange-winged

Pytilia lineata

and Green-winged Pytilias P. afra and P. melba, from YY also by grey, not red, face. Voice. Not tape-recorded. Said to be like Red-winged Pytilia (Nicolai 1968). Call in flight and when disturbed, ‘chwick’ (J.S. Ash, pers. comm.). General Habits. Inhabits open wooded grassland and open bush; almost entirely confined to Combretum-Terminalia woodland; occurs only occasionally in other woodlands and Acacia bush (J. S. Ash, pers. comm.). Food. In captivity, seeds, mealworms and mosquito larvae. Breeding Habits. Not known in wild except for a nest in May; in captivity, Y display and nest same as in Redwinged Pytilia P. phoenicoptera (though details not given: Nicolai 1968, Goodwin 1982). Key References

Goodwin, D. (1982), Nicolai, J. (1968).

Plate 17

Pytilia melba (Linnaeus). Green-winged Pytilia; Melba Finch. Beaumarquet melba.

(Opp. p. 266)

Fringilla Melba Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 180; China (¼Angola, Zedlitz, 1916, J. Orn., 64, p. 31; restricted to Luanda by Clancey, 1962, Bull. Br. Orn. Club, 82, p. 4). Range and Status. Endemic resident. Mauritania, uncommon, >20 localities north to 17
N; Senegal, frequent, widespread; absent south of 13
N. Gambia, uncommon

and local: Kiang; north of Kaur, mainly on N side of river; Basse; east of Farfenni. Guinea-Bissau, records in W. Guinea, single old record in N. Mali, widespread but

Pytilia melba

Pytilia melba

? ?

?

?

?

?

?

? ? ?

?

uncommon, between 13
and 17
N. Ivory Coast, uncommon in SW Comoe´ Nat. Park, 4 old records in extreme N, 1 in 1990 south of Niakaramandougou (8
270 N, 5
120 W), and single YY 4 times near there in 2000 (Rheindt et al. 2002). Burkina Faso, uncommon: N Yatenga area (20 records in 20 months), around Ouagadougou, and 5 records in E and SW. Togo, once, Mango, 10
210 N. Niger, frequent in extreme SW, otherwise only a few records, north to Tahoua, Zinder, Nguigmi and Dillia de Lagane (14
460 N). Nigeria, locally frequent in N, south to Sokoto, Kano, Kari and Maiduguri; recent record of pair in Gashaka-Gumti Nat. Park requires confirmation. Cameroon, frequent in Waza Nat. Park and occurs in Waza town (Scholte et al., 1999); south to Mora in W and Yagoua in E; 2 once on Mt Oku (Fotso 2001). Chad, common throughout sahel zone and Ouaddaı¨; widespread but not common in Wadi Rime-Wadi Achim Faunal Res.; uncommon in soudanian zone, south to c. 10
N; outlying population in Ennedi Mts. Central African Republic, uncommon in N Vakaga region (Bretagnolle 1993). Sudan, 5 races: citerior, clanceyi, belli and soudanensis common, from north to south respectively, in regions mapped; jessei uncommon, near Eritrean border. Eritrea, uncommon, below 1500 m; absent from coastal plains and south of c. 14
N; known from several localities east of 38
E; uncertain whether occurs west of 38
E. Ethiopia, frequent in regions mapped; perhaps much more widespread, but few recent records. Djibouti, Goda Massif (Toha, Dittilou, Randa-Bankouale´, 11
47–500 N, 42
42– 430 E) and Mabla Mts (Goula, 11
570 N, 43
000 E), at 500– 850 m. Somalia, frequent in NW and south of 3–5
N, mainly at 600–900 m; 3 records in NE. Uganda, frequent only in SW and extreme NE. Kenya, widespread from coast (south to Malindi) up to 1400 m but scarce and local; mainly east and south of central highlands. Tanzania, sparse and local in N, widespread east of Iringa highlands,

from Mikindani (where frequent) and Nachingwea (common) to Songea and S Njombe; common in Kibondo, uncommon to frequent in Zanzibar. Frequent in Rwanda and W half of Burundi. Congo, Brazzaville region (Landana, Manyanga) only. Zaı¨re, in W, along lower Congo valley east to near Kinshasa; just enters NE and E Zaı¨re in Garamba Nat. Park and Kivu Highlands east of Kasindi, around Luvungi and south to NW shores of L. Tanganyika (but not in Itombwe or Murungu Highlands); in SE, locally frequent in Katanga north on Luapulu R. to Kasenga. Angola, common, Cabinda, Luanda, Cuanza Norte and Malanje; and from S Bengo and coastal N Benguela through W Huı´la (where very common) to W Cunene; probably more widespread than shown. Zambia, widespread as mapped; in ˆ i, widespread 54% of 299 squares; rare north of 16
S. Malaw and common at least locally, below 1500 m; in places commoner and in others less common than Orangewinged Pytilia P. afra. Mozambique, common in Zambe´zia and Nampula Provs (Vincent 1936) but probably absent from much of Niassa and Cabo Delgado Provs, as shown. Uncommon in Namibia but in remainder of southern Africa common as mapped, with outlying records/populations in Botswana at Morwa Pan, Nossob, and along Molopo R. to Bray, in N Cape in Kalahari Gemsbok Nat. Park, Molopo and Kuruman drainage lines, and Kathu and Gamagara valley near Kuruman, in Free State between Bloemfontein and Senekal, and in KwaZulu-Natal east of Estcourt and near Mpendle. Population density in mixed Acacia-Dichrostachys thorn savanna of 025–125 breeding pairs per ha (Transvaal: Barnard and Markus 1990), i.e. pair per 08–40 ha; or 039– 079 breeding pairs per ha, i.e. pair per 13–26 ha (calculated from ringing data, Transvaal: Skead 1975). Density in Acacia savannas, Swaziland, 12, 32, 69 and 97 birds per 100 ha (Monadjem 2002). Est. 200,000 birds in Sul do Save, Mozambique. A popular cagebird, caught and exported in large numbers; during 1992–1993 drought, declined dramatically in SE Botswana ‘to the extent that bird-export companies failed to make up their usually sustainable quotas’ (Harrison et al. 1997). Mortality during shipment is high and in captivity this is regarded as a difficult and delicate species (Goodwin 1982). Description. P. m. citerior Strickland (includes ‘clanceyi’): Senegal to Nile Valley north of c. 10
N, intergrading in S Sudan with soudanensis. ADULT Y: forehead and forecrown to lores, around eye, forecheeks, chin and upper throat bright red or scarlet; hindcrown to hindneck and to ear-coverts, hindcheeks and sides of neck pale brownish grey; mantle to rump and scapulars pale yellowish olive-green; uppertail-coverts dull orange-red. Central tail-feathers orange-red above, T2–T5 dark brown with orangered outer webs, T6 dark brown with grey-buff outer web tinged orange near shaft. Underside of T1 grey-brown, of T2–T6 blackish brown with grey-brown outer web, palest on T6. Lower throat and upper breast olive-yellow with narrow grey-brown barring showing through on upper breast; lower breast and flanks buffish white with narrow grey-brown barring, merging into unbarred whitish lower belly and undertail-coverts. Flight feathers grey-brown, outer edges of primaries and outer webs of secondaries pale yellowish olive-green; primary coverts and alula grey-brown; tertials and greater, median and lesser coverts yellowish olive-green. Axillaries and underwing-coverts pale

355

356

ESTRILDIDAE

buffish white; inner borders of undersides of flight feathers pale grey-buff. Bill dark red; eyes red or orange-red; legs light brown or flesh. ADULT X: differs from ad. Y in having no red on face, and entire top and side of head brownish grey, slightly darker on lores and around eye; chin to upper breast pale grey-buff with narrow grey-brown barring, like lower breast and flanks; olive upperparts rather paler; red on uppertail-coverts duller, more restricted to posterior feathers. SIZE (10 YY, 10 XX): wing, Y 56– 61 (579), X 55–59 (574); tail, Y 45–52 (478), X 47–51 (481); bill, Y 135–15 (142), X 13–15 (140); tarsus, Y 15–17 (161), X 15–165 (159). WEIGHT: (n ¼ 34, Chad, unsexed) 10–17 (135). IMMATURE: juv. similar to X, but upperparts browner, head and throat washed olive, underparts pale unbarred greyish brown, whiter on centre of belly and undertail-coverts; red on uppertailcoverts restricted to largest feathers; bill blackish; eyes brown; legs greyish. NESTLING: skin dark, with sandy white down (A); mouth with median black palate spot, 2 lateral blue spots, whitish gape tubercles; lower region blackish; tongue partly dark (fig. p. 252). P. m. soudanensis (Sharpe) (includes ‘jubaensis’): Sudan (south of c. 10
N and east of Nile), S and E Ethiopia, Somalia, NE Uganda, N and E Kenya and NE Tanzania (south to Tarangire and Masai Steppe). Intergrades with citerior in S Sudan, jessei in Djibouti, and grotei in NE Tanzania. Olive upperparts brighter than in citerior, grey head darker, red on rump and uppertail-coverts slightly darker; underparts more heavily and darkly barred, white bars broken into spots on sides of breast. In Y, red face patch does not extend behind middle of eye; olive breast band greener than in citerior, lower border invaded by white spotting. X has chin to breast darker grey than in citerior. WEIGHT: (SE Kenya) Y (n ¼ 24) 132–162 (146), X (n ¼ 14) 12–162 (148). P. m. jessei Shelley: NE Sudan, Eritrea, NW Somalia. Intergrades with soudanensis in west and southeast. Y like soudanensis but lores grey; forehead and chin to throat lighter, more orange-red; upperparts paler and tinged greyish, underparts slightly paler. P. m. ‘flavicaudata’ [described by Welch and Welch (1988, 1998); although colour photographs have been depositied in Nat. Hist. Mus. (London) no specimen has been collected and so taxon is technically unnamed (Payne 1989). See also colour photo in Brickell (1999), and TAXONOMIC NOTE below.]: Goda Massif and Mabla Mts, Djibouti; also near Kathu and along 70 km of Gamagara R., N Cape Prov., South Africa. Y like Y jessei and Y soudanensis but forehead, front half of ear-coverts, chin, throat, upper breast and sides of neck bright golden yellow (or yellow with orange wash: Payne 1989), lores grey, rump rich golden yellow, tail the same with central feathers greenish; underparts from lower breast to undertail-coverts white with black barring, fine on lower breast, becoming coarser and broader on lower belly and flanks. Bill pink with grey culmen or saddle of grey near base of culmen; eyes deep red; legs and feet pale pink (Welch and Welch 1998, Hand 1998, Brickell 1999). X like Y but rump and tail not so bright yellow. Juv. and imm. like juv. and imm. P. melba (Brickell 2000).

P. m. percivali van Someren: central and SW Kenya, N and central Tanzania (east to Moshi, south to Iringa). Y differs from soudanensis in having lores and around eye grey, separating red forehead from red chin; upperparts slightly darker. X has darker grey head and breast and darker green upperparts. P. m. belli Ogilvie-Grant: W Uganda, Rwanda, Burundi, Zaı¨re (Kivu) and W Tanzania (south to Kigoma and Mpanda, east to Mwanza and Serengeti). Y like percivali, with grey lores, but red extends from chin and throat to centre of breast, invading olive band. Upperparts slightly darker and greener than in soudanensis, underpart barring darker and heavier. WEIGHT: (W Uganda) Y (n ¼ 9) 139–164 (152), X (n ¼ 6) 111–162 (145). P. m. grotei Reichenow: E Tanzania (inland to Kilosa, Liwale and Songea), Zanzibar, Mozambique south to lower Zambezi and ˆ i east of Rift Valley. Y like belli but red of throat and breast Malaw darker, invading centre of breast band more completely; sides of breast greener. Some YY with tinge of red on scapulars and upperwing-coverts. P. m. melba (Linnaeus) (includes ‘thamnophila’ and ‘damarenˆ i west of sis’): Congo and Zaı¨re (Katanga) to SW Tanzania, Malaw Rift, lower Zambezi and South Africa. Y brightly and darkly coloured like belli, and with grey lores, but red confined to forehead, chin and throat; upper breast olive-green. WEIGHT: Y (n ¼ 280) 134–179 (155), X (n ¼ 284) 128–184 (154), juv. Y (n ¼ 316) 127–174 (149), juv. X (n ¼ 228) 128–170 (148) (Skead 1975). P. m. hygrophila Benson and Irwin: N Zambia (N and W Luapala ˆ i (Dowa and L. Kasuni). Prov., N Northern Prov.) and N Malaw Like nominate race, but Y slightly darker red on face, darker grey on crown and nape and brighter olive-green on mantle; breast band more olive-green (less golden), underpart barring heavier. X generally darker throughout than nominate race. TAXONOMIC NOTE: in Djibouti ‘flavicaudata’ divides 2 populations of jessei, to northwest and southeast of it. Differences in appearance between P. m. ‘flavicaudata’ and other populations of P. melba are greater than those between P. phoenicoptera and P. lineata and are at least as great as those between P. hypogrammica and P. afra, so ‘flavicaudata’ may prove to be a distinct species. A population evidently identical to the Djibouti one, in Gamagara valley and Kathu, N Cape Prov., South Africa, might suggest that ‘flavicaudata’ is a genetic mutant of P. melba. Greatly outnumbered by typical red pytilias, it ‘breeds alongside Green-winged Pytilia Pytilia melba, but does not occur in mixed family groups, preferring to feed and drink separately’ (Brickell 1999) – indicating that they may indeed be separate taxa.

Field Characters. Length 12–14 cm. The most widespread and common pytilia, with green wings and back, blackand-white barred underparts. Y has bright red patches on forehead and throat, otherwise variable: rest of head grey, extending around eye to lores, breast green (nominate race, southern Africa), elsewhere red of throat may extend onto yellow or orange breast, and in W African citerior lores and face are red and barring on underparts less distinct. X larger than X Orange-winged Pytilia, with grey head, no orange on wings or back, more pronounced barring on undersides, extending onto throat; drab brown juv. has plain olive wings without orange. Newly hatched chick told from chick of brood-parasitic Long-tailed Paradise-Whydah V. paradisaea by: tubercles at corner of mouth same size (in parasite upper one larger than lower); skin dark but not black (parasite black); down sandy white (parasite grey-white); bill pointed (parasite broad and conical). Chick ready to leave nest: olive-grey with rump reddish (uniform grey-brown in parasite); 8 scales on tarsus (6 in parasite); parasite larger (Skead 1975).

Pytilia melba Voice. Tape-recorded (59, 86, 88, 99, 104, B, F, LEM, PAY). Song in southern Africa a quiet medley of hollow popping notes, thin chips and buzzes, low chuckles, double ‘woy-woy’ or ‘wooy-hyoo’ and long ascending whistle ‘wooooey’ often followed by long descending ‘shrrreeeeoo’; see examples in Goodwin (1982) and Maclean (1993). Song in Kenya (soudanensis) rather harsher and more disjointed, series of thin hisses, rattles, whistles and low chuckles; song of citerior ‘noticeably different’ (Goodwin 1982), a series of single ‘veet’ notes followed by whistles and gurgles, also said to lack the lengthy trilling songs and dripping water sounds noted in E and S Africa (Barlow et al. 1997). Contact call a loud, down-slurred, plaintive ‘tseeoo’ (sonagram in Gu ¨ ttinger and Nikolai 1973); other calls include low ‘wit’, sharp ‘tsip’ and soft, insect-like ‘kwik-kweek’ and ‘tsit-tsit’ in alarm. General Habits. Inhabits Acacia-Dichrostachys thorn savanna, in southern Africa especially A. tortilis and A. erioloba; Acacia mosaic in otherwise broad-leaved woodland with thorn thickets and open grassy patches; dense, dry savanna woodlands with tangled thickets; thin ironstone bush (Senegal, Gambia); wadis with plenty of Boscia senegala (Chad). Needs to drink regularly and availability of surface water clearly affects local distribution (Harrison et al. 1997). Usually in pairs. Family parties of ads and very young juvs occur for short time after nesting. Concentrates at feeding and drinking places but not gregarious and pairs go their separate ways afterward. Shy, unobtrusive and inconspicuous; very difficult to watch for any length of time. Forages for seeds exclusively on ground, often in company of firefinches Lagonosticta spp., waxbills Estrilda spp., cordon-bleus Uraeginthus spp. and Speckle-fronted Weavers Sporopipes frontalis. Takes termites by picking them from ground, moving sticks or bits of dung under which termites are working, perching on termitaria and taking termites from inside chimneys, and by hawking them from ground and from perch on bush (Skead 1975). Forages mainly in morning and late afternoon; sometimes feeds after sunset; drinks regularly, mainly in morning (in South Africa in winter); works its way through tangled vegetation overhanging stream, drinks quickly and flies off. Once seen to bathe in rain puddle. Resident but wanders locally; for instance, numbers at Malamfatori, NE Nigeria, increase markedly from mid Sept (start of breeding season). Much seasonal variation at Rurutse, SE Botswana, from c. 30 birds netted per 10 km in Apr–Sept to 200 localities as mapped, north to near In Zekouane (18
450 N). Ivory Coast, once Ouangolodougou. Burkina Faso, frequent in N Yatenga area, uncommon in Ouagadougou area (Gonse´, Koubri). Ghana, 3 singles in far N and flock of 19 at Bolgatanga. Togo, twice, Kpani and Parc Nat. de la Ke´ran. Benin, several, Pendjari Nat. Park, Apr 1984. Niger, frequent across middle from Tahoua to Tanout, Zinder, Baban Tubki (where can be abundant) and L. Chad; in dry season in ‘W’ Nat. Park and Niger valley north to Saga. Nigeria, locally common across N in sahel and soudanian zones; in dry season south to Kainji Lake Nat. Park and Zaria; became much commoner at Zaria in 1970s, perhaps as a consequence of the sahel drought then; 5 near Lagos, Jan 1984, possibly escapes from captivity (Alexander-Marrack et al. 1985). Cameroon, common in Waza Nat. Park and south to Garoua. Chad, uncommon in soudanian zone, common in sahel zone, north to Ouadi Rime Faunal Res. (northern limit, Ouadi Haddat) and Ennedi. Sudan, frequent in middle belt, south to Bahr al Ghazal, north on Nile to Berber and once near Wadi Halfa, also to Ashraf area near Red Sea coast; alexanderi uncommon in East Equatoria. Eritrea, widespread in W and N, below 1520 m; occurs on central plateau up to 2130 m; probably occurs further south than shown. Ethiopia, frequent to common below 1200 m, probably more widespread in NE, SE and S than shown. Somalia, frequent at 900–1500 m in NW, and in S in lowlands south of 5
N. Uganda, only in NE, in Moroto and Kidepo Valley Nat. Park. Kenya, common and widespread below 1300 m; local movements result in

Amadina fasciata

?

sporadic occurrences in L. Victoria basin and towards coast. Tanzania, frequent to common in NE and on dry central plateau; common around Dar es Salaam, probably escaped cagebirds. Angola, record at Ca´fu, Cunene. Namibia, rare; several recent records. Zambia, in 12% of 300 squares, mainly in S (Choma, Monze); frequent in Zambezi valley from Livingstone to Fira; in Luangwa ˆ i, from Kasache and Salima on SW valley at Mfuwe. Malaw ˆ i littoral, south along Shire valley; L. Chilwa. L. Malaw Botswana, Zambia, Transvaal and extreme NE Cape Prov., uncommon to locally frequent and nomadic, as mapped (Harrison et al. 1997); in Zambia, recent range extensions to Kasama (flock of 20, Sept) and Kabwe; vagrant in Free

380

ESTRILDIDAE

State and KwaZulu-Natal (Cyrus and Robson 1980, Earle´ and Grobler 1987). Mozambique; records from Nampula coast near Moambique town and Inhambane coast near Imhambane town, and a few in river valleys in W Sul do Save; estimated to be >5000 birds (Parker 1999). Description. A. f. fasciata (Gmelin): Senegal to Sudan (except extreme SE). ADULT Y: forehead to hindneck and down to eye buff, barred or scaly; each feather with blackish subterminal bar and more central V-shaped bar, producing scaly effect. Upper mantle cinnamon-buff, narrowly barred blackish; lower mantle to back and scapulars deeper cinnamon or pinkish buff, usually spotted black, sometimes with extensive black barring; cinnamonbuff rump and paler buff uppertail-coverts barred black, each feather with black subterminal V-bar and smaller central V-mark. Tail feathers grey-brown, darkening toward tip; T2–T6 with progressively larger white spot at tip, T6 also with white outer web. Lores pale buff; from immediately behind eye to forecheeks and chin buffy white; patch from ear-coverts across rear cheeks to upper throat dark red, often bordered behind by narrow buff and black speckled band; behind it a buffy white collar on lower side of neck and around lower throat. Upper breast pale cinnamonbuff, bordered below by variable black-speckled or black-barred band; lower breast and upper belly darker chestnut-brown; sides of breast and flanks cinnamon-buff, flanks with variable blackish barring, boldest at rear; lower belly buffy white; undertail-coverts pale buff with a few crescentic tawny-brown scales. Remiges dull dark brown, edged buffish, tertials tipped cinnamon-buff; primary coverts dark brown; greater coverts edged and tipped cinnamon-buff, inner feathers with blackish sub-terminal bar; median and lesser coverts buff, barred blackish. Underwingcoverts and axillaries pale cinnamon-buff. Bill pale horn-grey, pinkish grey or bluish grey, often darker on culmen; eyes light brown to reddish; legs pale flesh. ADULT X: lacks red face patch; cheeks and ear-coverts to chin and throat buff, finely spotted blackish brown. Slightly paler, more sandy above than ad. Y, barring rather browner, less heavy; paler buff, less cinnamon below with weaker band of bars on breast, and little or no chestnut on upper belly. SIZE (10 YY, 10 XX): wing, Y 65–69 (674), X 65–69 (667); tail, Y 39–43 (414), X 36–42 (380); bill, Y 125–14 (132), X 125–13 (128); tarsus, Y 15–175 (162), X 15– 16 (156). WEIGHT: Nigeria, Y (n ¼ 8) 161–196 (178), 1 X 180; Chad, unsexed (n ¼ 95) 150–320 (180). IMMATURE: juv. similar to ad. X, but tertials with large pale buff tips and blackish subterminal marks, greater, median and lesser coverts tipped pale buff. Juvenile Y has band of red-tipped feathers around throat, and poorly marked patch on belly. NESTLING: skin bluish black; down pale, profuse on head and underparts; inside of mouth yellow, palate with 5 black spots (2 medial and 2 lateral near centre, 1 at mandible tip); swollen gape flanges pale yellow or white (fig. p. 252). A. f. alexanderi Neumann: Eritrea, Ethiopia and Somalia to extreme SE Sudan, NE Uganda, Kenya and Tanzania. Rather darker and browner, less cinnamon above than nominate race, with bolder barring on head and entire mantle, scapulars and back; underparts more extensively barred, all breast and flank feathers with V marks, chestnut patch more confined to upper belly. A. f. meridionalis Neunzig: S Angola, N Namibia, Zambia, ˆ i, NE Mozambique, W Zimbabwe and Botswana. Like Malaw alexanderi, but mantle darker, more tawny brown, top of head darker and greyer with broader barring. Bill smaller: Y (n ¼ 10) 11–12 (117). A. f. contigua Clancey: S Zimbabwe, S Mozambique, South Africa and Swaziland. Y differs from meridionalis in having ground colour of head top buffier (less whitish); mantle, rump and scapulars with pale brown cross barring and apical spotting buffier; chin buffy (not clear white); red throat bar lighter carmine; lower throat buffier (less white), less well demarcated from breast; brown abdominal patch warmer and slightly paler. X

slightly more cinnamon (less greyish) above than meridionalis, more pinkish cinnamon on breast and sides of underparts. WEIGHT: Transvaal, 2 YY 177, 179, 2 XX 174, 174.

Field Characters. Length 12 cm. Fine zebra barring on head and body diagnostic; Y has red ‘cut-throat’ slash extending onto sides of neck, contrasting with barred brown head (head entirely red in Red-headed Finch A. erythrocephala). X told from X Red-headed by barring on head and upperparts, browner tone to back and breast, irregular broken dark barring on underparts rather than white spots and black scalloping. Immature like X but less barring on underparts, none on back; Y has broken red throat band mixed with black. Voice. Tape-recorded (99, 104, B, GREG, GUT, McVIC, PAY). Calls higher-pitched than in Red-headed Finch A. erythrocephala, without House Sparrow quality: grating ‘zheep’ or ‘pzeep’, lower ‘zherk’ and ‘chizh’, dry ‘bzzt’, upslurred ‘weezht’. In flight, thin ‘eee-eee-eee’; loud plaintive ‘kee-air’ given by nesting birds (Goodwin 1982). Delightful song a continuous rather acrocephaline medley of sweet whistles, bubbling trills, grating churrs, chuckles and cheeps, with a few hard notes suggestive of quailfinch Ortygospiza; not loud, but with a feeling of suppressed power, like Nightingale Luscinia megarhynchos. Other vocalizations, see below. General Habits. Inhabits Acacia and other wooded savannas; arid lowland thornbush, scrub on sandy soils in dry woodland, coastal broad-leaved open woodlands, Colophospermum mopane woods, large rural and urban gardens, wadis in semi-desert, cultivation including harvested millet fields (Chad) and riparian scrub (Tacazze ˆ i). valley, W Eritrea); often under Hyphaene palms (Malaw Usually in pairs, but habitually forms flocks outside breeding season with weavers and estrildids (e.g., in Chad, Ploceus vitellinus, P. luteolus, Uraeginthus bengalus and Sporopipes frontalis) and freely associates with Red-billed Queleas Quelea quelea; sometimes in large monospecific flocks. Readily flocks with Red-headed Finch in Makgadikgadi Pans area, NE Botswana. Forages on sandy patches of bare ground; hops, walks and runs short distances. Pair often perches on bare branch at top of tall tree. Often roosts (as well as breeding) in old weaver nests; sometimes roosts in natural cavities. Very partial to dust bathing. Regularly comes to drink at pans and waterholes, about mid-day. Mainly resident but wanders, and occurrence in many areas unpredictable. Migratory in W Africa, however: resident in sahel zone; dry season visitor to Gambia, Nov– May, mainly Dec–Mar; evidently not breeding, although once seen emerging from nest of Village Weaver Ploceus cucullatus in Nov; dry season visitor to Kano, N Nigeria, Oct–May (latest, 5th June); presumably a dry season visitor throughout Nigerian range – certainly so in Borgu (visits Mar–Apr) and Zaria (visits early Dec to early May, breeds in Jan); all records in Ivory Coast, Ghana, Togo, Benin and SW Niger are in mid or late dry season, except for 1 in Togo in July (Demey 1998); most or all records from N Cameroon (Le´re´, Yagoua, Waza) are dry season; passage

384

ESTRILDIDAE

‘spectacles’ and large white chinspot, making them radically different in appearance, if not from ugandae (which is intermediate in character, spectacled in Uganda, often not so in Sudan), then from atricollis and in particular ansorgei. DNA sequencing of 5 taxa shows that atricollis and ansorgei are very closely allied; that fuscata and gabonensis are very closely allied to each other and quite closely to muelleri; but that atricollis-ansorgei and fuscata-gabonensis/muelleri are about as distant from each other as are several other pairs of African estrildid species. That is our interpretation of M. D. Sorenson and R. B. Payne’s unpublished estrildid phylogenetic tree, although Sorenson and Payne (2001) recognize only the single species, O. atricollis. To us, it still seems best to separate O. gabonensis and to divide O. atricollis into 2 species, following White (1963), O. atricollis sens. str. and O. fuscocrissa. In Uganda O. gabonensis is parapatric with O. atricollis and resembles it, both being separated by L. Victoria from the dissimilar-looking O. fuscocrissa. The nomadic tendency of the superspecies may account for ‘spectacled’ gene flow into Sudan. Further vocal and molecular researches of all populations are needed. Endemic. 3 species, composing a superspecies.

Ortygospiza atricollis superspecies

1 1

1 or 3

2

7

1 or 2 1

O. atricollis 1 atricollis 2 ansorgei 3 'ugandae' O. gabonensis 4 gabonensis 5 dorsostriata 6 fuscata O. fuscocrissa 7 fuscocrissa 8 muelleri 9 smithersi 10 pallida 11 digressa

3

5

4 6

8 6 9

8 8 10 8 11

Areas of overlap solid red.

Plate 21

Ortygospiza atricollis (Vieillot). Black-faced Quailfinch. Astrild-caille a` face noire.

(Opp. p. 330)

Fringilla atricollis Vieillot, 1817. Nouv. Dict. Hist. Nat., nouv. ´ed., 12, p. 182; Senegal. Forms a superspecies with O. gabonensis and O. fuscocrissa. Range and Status. Endemic resident and wanderer, Senegal to N Cameroon. Mauritania, uncommon in Senegal R. valley and Karakoro drainage; record near Nouakchott. Senegal, locally frequent to common in Senegal R. valley, south to Richard-Toll, and in SW and S. Gambia, frequent throughout. Occurs near coast in Guinea-Bissau, Guinea (e.g. Conakry Airport), Sierra Leone (formerly common on Freetown golf course) and Liberia, where not uncommon resident along coast and coastal plain below 100 m, east to

about Buchanan; in Guinea, occurs in NE south on Milo R. to Kankan, and to Haut Niger Nat. Park (where frequent: Nikolaus 2000). Mali, uncommon to frequent on Niger R. floodplain and higher ground between Bandiagara and Hombori; several records in Kirane-Tango area in NW, near Bafoulabe´ and in Res. de Fina; old records near Sikasso in SE. Ivory Coast, not uncommon from Korhogo to Comoe´ Nat. Park, records south to Man and Bouake´, and Assinie on coast (Bull. Afr. Bird Club 2001, 118). Burkina

Ortygospiza atricollis

Ortygospiza atricollis

Faso, known only from Loumbila and Gonse´, northeast of Ouagadougou, and from Arli Nat. Park. Niger, old record from Tillabe´ri. Ghana, in N, records only at Tugu and Gambaga; in S, not uncommon on Accra Plains and coastal thicket zone. Togo, locally common, Tantigou south to Ayengre´; record on coast near Lome. Benin, frequent in Pendjari Nat. Park. Nigeria, locally common in N from Kainji and New Bussa to Mongonu, north to Sokoto, Kazaure, Kirikasama and Malamfatori, south to Jos Plateau, Kafanchan, Lokoja and Kiri on Gongola R.; uncertain whether it occurs on Niger and Benue R. floodplains, but in SW, occurs locally from Lagos to Iwo, e.g. 6 records at Lagos and Tarkwa in 2 years; in SE, common round Okigwi, north to Enugu, and once near Wukari. Cameroon, Waza Nat. Park, Sir and Rhumsiki area, very common around Garoua; 2 records further south (M. Languy, pers. comm.). Chad, old record on SE corner of L. Chad. Absent from Central African Republic. Sudan, rare in S Kordofan, uncommon in E Sobat from Nile to Ethiopian border, uncommon in White Nile and Equatoria Provs (Malakal, Aweil, Tonj, Rumbek, Juba, Kajo Kaji). Zaı¨re, only around Kasenye, SW shore of L. Albert. Uganda, records from Moyo, Albert Nile, Murchison Falls Nat. Park and Entebbe; breeding range uncertain. Kenya, old record at Mumias. Description. O. a. atricollis (Vieillot) (includes ‘ugandae’? – see below): Senegal to Cameroon (‘ugandae’ further east). ADULT Y: forehead blackish; rest of upperparts drab or greyish brown with faint darker mottling. Tail short, dark grey-brown, tip and outer web of T6 white. Lores grey-brown; rarely, a narrow white line above lore and eye; cheeks (back to rear of eye), chin and throat blackish, forming a dark patch merging with grey-brown ear coverts, hindcheeks and sides of neck; often a small white area on upper chin at base of lower mandible. Upper breast greyish brown, finely barred white; sides of breast, flanks, and sides of

belly greyish brown, tinged tawny posteriorly, with broader, more widely spaced barring, each feather crossed by a blackishbordered white bar and with a V-shaped buffish white bar nearer base; centre of lower breast and belly chestnut-brown, grading to tawny-buff on vent, thighs and undertail-coverts, the last broadly streaked dark grey-brown. Upperwing grey-brown; underwingcoverts and axillaries pale tawny buff. Bill red, upper mandible dusky or brown in non-breeding condition; eyes light hazel; legs yellowish flesh. ADULT X: differs from ad. Y in having entire head and neck greyish brown; upperparts slightly paler and browner; breast and flanks paler, more buffish grey, barring tinged buff and less sharply defined; belly patch tawny buff (less chestnut). SIZE (10 YY, 10 XX): wing, Y 48–55 (522), X 50–56 (517); tail, Y 24–27 (257), X 24–27 (252); bill, Y 85–95 (90), X 85–10 (92); tarsus, Y 135–155 (143), X 135–155 (146). IMMATURE: juv. similar above to ad. X; dull tawny-brown below with paler grey-brown chin and throat, upper breast and flanks with faint dusky barring. Bill dusky. NESTLING: skin pale orange-pink; long, pale grey down on crown, back, rump and thighs; in mouth, palate pinkish white with ring of 6 black spots (2 round ones on each side with midline pair in front); tongue pink with broken black bar; corner of gape swollen and magenta, with 2 pale blue papillae on maxilla (front one half size of back one) and one on mandible, all 3 surrounded by black, giving checkerboard pattern with pale lower papilla fitting under black between upper pair (Payne and Payne 1994). O. a. ansorgei Ogilvie-Grant: coastal Guinea-Bissau to W Liberia and coastal Ghana; probably this race in S Nigeria. Intergrades with atricollis in S Senegal and SE Mali. Upperparts much darker and greyer than in nominate race, with black on head extending further onto ear-coverts and onto upper breast, reducing the extent of barring; flanks darker and greyer. O. a. ‘ugandae’ van Someren 1921: S Sudan (White Nile and Equatoria Provs), NW Uganda and NE Zaı¨re (Kasenye, where breeds), vagrant(?) to L. Victoria (Entebbe) and W Kenya (Mumias, including type specimen) (Traylor 1963). Distinguished from atricollis only in having fine (not wide) line around lore and eye, lacking in some Sudan birds, and slightly larger white chinspot. Described from 2 birds at Mumias (van Someren 1921) but status uncertain. Synonymized with O. (fuscocrissa) muelleri by White (1963); recognized by Paynter et al. (1968); not recognized by Nikolaus in Sudan (1987, 1989: only atricollis, south to Uganda border at Kajo Kaji and Moyo), nor by Britton in Uganda (1980: only atricollis), nor by Zimmerman et al. in Kenya (1996: only muelleri, west to near Uganda border). TAXONOMIC NOTE: see generic diagnosis.

Field Characters. Length 9–10 cm. Typical quailfinch behaviour, well described by Barlow et al. (1997): ‘almost always first seen after flushing from underfoot, springing steeply up from rice stubble or swamp grassland, showing dumpy, bobbing silhouette, circling against sky uttering small tinny cries’. Black extends farther back on forehead and face than in Y Black-chinned Quailfinch O. gabonensis, ear-coverts grey; in both sexes upperparts are grey-brown, largely unstreaked. Juv. browner below, barring muted, bill mainly black. Race ansorgei much darker, head and breast blackish, upperparts dark grey. Voice. Tape-recorded (104, 108, MOR, PAY). Extremely similar to other quailfinches. Social contact call given by pairs feeding out of sight of each other a harsh ‘klek’, also used by parents and young calling back and forth (Payne and Payne 1994); fledgling separated from its parents gave harsher version. Variants of ‘klek’ given in excitement or alarm. Flight calls ‘tink’ or plaintive ‘seep’. Songs as described under African Quailfinch O. fuscocrissa. Begging

385

386

ESTRILDIDAE

and other nestling calls described by Payne and Payne (1994), and see sonagrams therein. General Habits. Inhabits treeless, short-grass savannas often inundated in wet season, floodplains, coastal plains, low grassy hills (SE Nigeria), tussocky drying edges of floodplain marshes, burnt grassland and dried-out rice fields after harvesting, farmland stubble, airstrips and airports, large games fields, vegetated margins of coastal sandflats (Gambia) and vegetable fields on sandy soils (Liberia). Occurs singly and in pairs in breeding season, in flocks of dozens all year, sometimes up to 80–100. Forages on open ground, often at edge of drying-out pond, bird standing in shallows and taking items from surface; perches on low vegetation very rarely and only when ground flooded. Flighty and elusive; hard to see on ground and harder to watch; flock put to flight quickly rises high in air, circles in wide radius, and often alights on ground near original spot, birds keeping still and well camouflaged in lee of tussocks. Sings on ground. Food. Not known. Breeding Habits. Solitary nester. NEST: made of grass, spherical or pear-shaped, with side entrance (A). Placed on ground at base of grass tussock, or sheltered by weeds in depression at side of furrow. EGGS: 2–5 (5 complete clutches, Nigeria). White. SIZE: (n ¼ 4) 137–143  110–112 (140  1115). LAYING DATES: Mauritania, Sept–Feb; Senegal, Sept– Dec, mainly Nov (11 out of 25 clutches), once Feb;

Gambia, Nov; Mali, Jan–Feb; Ghana, Aug; Nigeria, Sept– Dec, Feb; Zaı¨re, Kasenye, (oviduct egg Sept). INCUBATION: period: 18 days (Serle 1940, 1943b). DEVELOPMENT AND CARE OF YOUNG: in captivity young retain blue papillae on gape for 3 weeks and black spots on palate for 3 months after fledging. BREEDING SUCCESS/SURVIVAL: parasitized by Quailfinch Indigobird Vidua nigeriae (for evidence, see that species).

Plate 21

Ortygospiza gabonensis Lynes. Black-chinned Quailfinch. Astrild-caille a` gorge noire.

(Opp. p. 330)

Ortygospiza gabonensis Lynes, 1914. Bull. Br. Orn. Club, 33, p. 131; Gabon. Forms a superspecies with O. atricollis and O. fuscocrissa. Range and Status. Endemic resident and wanderer, Mbini and Congo to Zambia; E Zaı¨re and Uganda. Mbini, Bata to Benito. Gabon, in W locally common from Port Gentil to Gamba and probably along entire length of coastal lowlands; frequent in Re´s. de la Lope´; locally common in Le´koni. Congo, local in Odzala Nat. Park; occurs on coast; widespread in Le´fini and lowlands along Congo R. where also on Zaı¨re bank (frequent around Kinshasa; upstream to Lukolela). NE Zaı¨re, S Lendu Plateau (Dele and Bogoro, 1370 m, which are 15 km west of and 750 m higher than Kasenye, SW L. Albert, where O. atricollis ‘ugandae’ occurs), immediately north and south of L. Edward (Kasindi, Lubilia Valley, Kabare, Rutshuru Plain); also in Itombwe highlands around Luiko, at 1860–2010 m. Uganda, widespread and sometimes common at 900–1500 m from Kigezi, Ankole and Toro east to Masaka, Entebbe and Mpumu. Tanzania, north of Bukoba on L. Victoria and around Ilabiro near Burundi border; also just into extreme SW (S end of L. Tanganyika). Rwanda, Kagera. Burundi, abundant

in Kumoso region (Kininya, Kiharo, Magera, Giofi). Angola, known only from Huambo and SW Lunda Sul; quailfinches around Luando town thought to be O. atricollis. S Zaı¨re, scattered records in Kasai Occidental and SW Katanga, common around Kananga, and probably common and far more widespread in the region than shown (Chapin 1954); in SE, frequent in Pepa area in Marungu highlands. Zambia, widespread in Northern and Luapula Provs (Mbala, Mporokoso, Kasam, Mansa); Copperbelt (Ndola, Kitwe); N North-Western Prov. (Solwezi, Mwinilunga) south to Kabompo and Balovale Provs (Mayau, Mongu, South Luete R.); occurs in 18% of 300 Zambia squares. Description. O. g. dorsostriata van Someren: Uganda and E Zaı¨re to Burundi and NW Tanzania. ADULT Y: forehead black; crown to uppertail-coverts, including scapulars, dark brown with warm buffish brown streaks produced by pale feather edges. Tail short, blackish brown, T6 with whitish fringe at tip and whitish outer edge, T5 with small whitish mark at tip. Lores brown; forecheeks to entire chin and throat black, merging with warm brown rear

386

ESTRILDIDAE

and other nestling calls described by Payne and Payne (1994), and see sonagrams therein. General Habits. Inhabits treeless, short-grass savannas often inundated in wet season, floodplains, coastal plains, low grassy hills (SE Nigeria), tussocky drying edges of floodplain marshes, burnt grassland and dried-out rice fields after harvesting, farmland stubble, airstrips and airports, large games fields, vegetated margins of coastal sandflats (Gambia) and vegetable fields on sandy soils (Liberia). Occurs singly and in pairs in breeding season, in flocks of dozens all year, sometimes up to 80–100. Forages on open ground, often at edge of drying-out pond, bird standing in shallows and taking items from surface; perches on low vegetation very rarely and only when ground flooded. Flighty and elusive; hard to see on ground and harder to watch; flock put to flight quickly rises high in air, circles in wide radius, and often alights on ground near original spot, birds keeping still and well camouflaged in lee of tussocks. Sings on ground. Food. Not known. Breeding Habits. Solitary nester. NEST: made of grass, spherical or pear-shaped, with side entrance (A). Placed on ground at base of grass tussock, or sheltered by weeds in depression at side of furrow. EGGS: 2–5 (5 complete clutches, Nigeria). White. SIZE: (n ¼ 4) 137–143  110–112 (140  1115). LAYING DATES: Mauritania, Sept–Feb; Senegal, Sept– Dec, mainly Nov (11 out of 25 clutches), once Feb;

Gambia, Nov; Mali, Jan–Feb; Ghana, Aug; Nigeria, Sept– Dec, Feb; Zaı¨re, Kasenye, (oviduct egg Sept). INCUBATION: period: 18 days (Serle 1940, 1943b). DEVELOPMENT AND CARE OF YOUNG: in captivity young retain blue papillae on gape for 3 weeks and black spots on palate for 3 months after fledging. BREEDING SUCCESS/SURVIVAL: parasitized by Quailfinch Indigobird Vidua nigeriae (for evidence, see that species).

Plate 21

Ortygospiza gabonensis Lynes. Black-chinned Quailfinch. Astrild-caille a` gorge noire.

(Opp. p. 330)

Ortygospiza gabonensis Lynes, 1914. Bull. Br. Orn. Club, 33, p. 131; Gabon. Forms a superspecies with O. atricollis and O. fuscocrissa. Range and Status. Endemic resident and wanderer, Mbini and Congo to Zambia; E Zaı¨re and Uganda. Mbini, Bata to Benito. Gabon, in W locally common from Port Gentil to Gamba and probably along entire length of coastal lowlands; frequent in Re´s. de la Lope´; locally common in Le´koni. Congo, local in Odzala Nat. Park; occurs on coast; widespread in Le´fini and lowlands along Congo R. where also on Zaı¨re bank (frequent around Kinshasa; upstream to Lukolela). NE Zaı¨re, S Lendu Plateau (Dele and Bogoro, 1370 m, which are 15 km west of and 750 m higher than Kasenye, SW L. Albert, where O. atricollis ‘ugandae’ occurs), immediately north and south of L. Edward (Kasindi, Lubilia Valley, Kabare, Rutshuru Plain); also in Itombwe highlands around Luiko, at 1860–2010 m. Uganda, widespread and sometimes common at 900–1500 m from Kigezi, Ankole and Toro east to Masaka, Entebbe and Mpumu. Tanzania, north of Bukoba on L. Victoria and around Ilabiro near Burundi border; also just into extreme SW (S end of L. Tanganyika). Rwanda, Kagera. Burundi, abundant

in Kumoso region (Kininya, Kiharo, Magera, Giofi). Angola, known only from Huambo and SW Lunda Sul; quailfinches around Luando town thought to be O. atricollis. S Zaı¨re, scattered records in Kasai Occidental and SW Katanga, common around Kananga, and probably common and far more widespread in the region than shown (Chapin 1954); in SE, frequent in Pepa area in Marungu highlands. Zambia, widespread in Northern and Luapula Provs (Mbala, Mporokoso, Kasam, Mansa); Copperbelt (Ndola, Kitwe); N North-Western Prov. (Solwezi, Mwinilunga) south to Kabompo and Balovale Provs (Mayau, Mongu, South Luete R.); occurs in 18% of 300 Zambia squares. Description. O. g. dorsostriata van Someren: Uganda and E Zaı¨re to Burundi and NW Tanzania. ADULT Y: forehead black; crown to uppertail-coverts, including scapulars, dark brown with warm buffish brown streaks produced by pale feather edges. Tail short, blackish brown, T6 with whitish fringe at tip and whitish outer edge, T5 with small whitish mark at tip. Lores brown; forecheeks to entire chin and throat black, merging with warm brown rear

Ortygospiza gabonensis

Ortygospiza gabonensis

? ? ?

?

can occur together. Voices similar but not identical; some calls different enough to be learned with experience. Voice. Tape-recorded (86, 102, 104, B, GBG, LEM, MOY). Flight call a hard ‘tuk’, ‘tewk’ or ‘tutup’, almost identical to African Quailfinch but higher-pitched; said by R.J. Stjernstedt (in Penry 1986) to be ‘very similar to the hard, grating chirp of African Quailfinch but softer and less clipped’; described by Penry (1986) as ‘drink drink’, but as with African Quailfinch there is no ringing quality. Other calls (context?) are drier, partly rolled, ‘cheep’, ‘chrrt’, ‘krrep’, some with squeaky overtone. The ‘song’ described by Penry (1986) as ‘take it away’ or slower ‘dukit-du-way’, or repeated in rapid succession when bird is excited, ‘dukituway dukit dukituway’ is classified by Nuttall (1993) as the X contact phrase. For sonagrams and further discussion see Penry (1986).

?

cheeks and ear-coverts; sides of neck dark brown. Upper breast, sides of breast and flanks barred blackish and white, narrowly on breast, more broadly on flanks where warm brown feather bases show through. Centre of lower breast pale orange-chestnut, grading to pale tawny buff on belly and undertail-coverts, the latter streaked and barred blackish. Upperwing feathers dark brown, tertials and wing-coverts tinged paler brown; underwing coverts and axillaries pale rufous-chestnut. Bill bright red; eyes pale, dull orange; legs pale brown. ADULT X: differs from ad. Y in having forehead, cheeks, chin and throat dull greyish brown; brown ground colour more prominent on breast and flanks; lower breast patch slightly paler, more orange-buff. Bill dark brown or blackish with reddish edges to mandibles; eyes light hazel; legs dull brown. SIZE (4 YY, 3 XX): wing, Y 50–53 (518), X 50–52 (513); tail, Y 25–27 (260), X 25–26 (257); bill, Y 95–10 (99), X 9–10 (95); tarsus, 145–155 (150), X 14–15 (145). IMMATURE: no information. NESTLING: mouth markings as in O. atricollis. O. g. gabonensis Lynes: Mbini, Gabon, Congo. Warmer brown above than dorsostriata; belly paler, orange-chestnut patch confined more to anterior part of lower breast. Smaller; wing, Y (n ¼ 2) 48, 48, X (n ¼ 2) 47, 47. O. g. fuscata Sclater: W and S Zaı¨re (Middle Congo R. to Kasai and Katanga), central Angola and Zambia. Y darker, blacker above than in dorsostriata, pale streaks narrower; ground colour of breast and flanks blacker, less brown showing through; central underparts darker, more chestnut. X darker above, greyer brown on sides of head, chin and throat. Larger; wing, Y (n ¼ 4) 53–56 (540).

Field Characters. Length 9–10 cm. Very similar to Blackfaced Quailfinch O. atricollis but browner above with dark mottling; Y has warm brown patch on cheeks and earcoverts, reduced black on face (narrowly on forehead, chin and lores); X with no black extremely similar to X Black-faced, best told by spotted upperparts, including crown. Race fuscata overlaps African Quailfinch O. fuscocrissa in central Africa but is blacker above and lacks any white on face. Prefers damper habitats, but the 2

General Habits. Inhabits short grass level plains (grass 30–40 cm tall) in lowlands and plains and grassy and gravelly hillsides in highlands; swampy, sedgy grasslands and fields, sometimes surrounded by woods; burnt grass; tussocky ground; permanently moist dambos and flood plains (Zambia). In pairs and small flocks of c. 12 birds; sometimes in threes (2 YY and 1 X; Clement et al. 1993). Forages on ground, keeping itself well hidden from view amongst tufts of grass. Does not perch on vegetation. Moves between tussocks in creeping manner, walk-hopping in short steps. Undemonstrative and difficult to observe. When flushed, flock typically rises almost vertically and circles often at considerable height, before plunging back to ground, alighting amongst vegetation and becoming instantly ‘invisible’ (Christy and Clarke 1994). Calls in flight, but may give flight call or part of song from ground, especially when alarmed (Penry 1986). Mainly resident, but at any one locality prone to absences and influxes; nomadic, making considerable local movements, e.g. in Kitwe region of Zambia, where much commoner in Feb–Apr than in June–Nov (Penry 1986), amounting to migration (Winterbottom 1959). Food. Tiny grass and herb seeds and occasional tiny spiders. Breeding Habits. Solitary nester; presumably monogamous; no evidence of territoriality. Vigorous chase of one bird by another seen only once, low over ground. At start of breeding season, majority of birds in an area continue to forage and fly as a flock, but some pairs separate from the flock and their vocalisations change. NEST: ball made of old fine-stemmed grass, with side entrance; lined with finer, green grasses; 1 nest (out of 5) also with 7 feathers of Helmeted Guineafowl Numida meleagris. Ext. diam. 90–115, ext. height 110, entrance diam. 30–50. Placed at edge of inconspicuous, flattened tuft of grazed-down or otherwise short grass, with base of nest on or very near wet ground or touching water 2 cm deep. EGGS: 3–4. White, stained orange with oily ooze from damp ground within 3 days. SIZE: (n ¼ 9) 137–152  110– 123 (145  117).

387

388

ESTRILDIDAE

LAYING DATES: NE Zaı ¨re, (juvs Sept); Uganda, Aug; Zambia, Chingola Mar–Apr (when no evidence of nesting at Kitwe), elsewhere Jan–Feb (and active gonads Mar).

INCUBATION:

Key References

evidence that both sexes incubate. Penry, E.H. (1986), Traylor, M.A. (1963).

Plate 21

Ortygospiza fuscocrissa Heuglin. African Quailfinch. Astrild-caille a ` lunettes.

(Opp. p. 330)

Ortygospiza fuscocrissa Vieillot, 1863. J. Orn., 11, p. 18; Dembea and Tigre´, Abyssinia. Forms a superspecies with O. atricollis and O. gabonensis. Range and Status. Endemic resident and nomad, Eritrea to South Africa. Eritrea, uncommon and sporadic (possibly migratory – not seen in winter) above 1830 m. Ethiopia, frequent to common above 1500 m and up to at least 2440 m, north to L. Tana, east to Somali border, south to Alghe; also in Rift Valley. Somalia, old record from Hargeysa. Kenya, frequent but local, above 1500 m; commonest in Laikipia Plateau, Nairobi Nat. Park, Mara Game Res., L. Baringo, L. Nakuru; up to 3000 m in Mau Narok; rare as far east as Tsavo East. Tanzania, locally abundant from Kenya border to 3
300 S between 34
300 E and 37
300 E; rare or absent in interior; scarce on Ufipa Plateau (where some observations may refer to Black-chinned Quailfinch O. gabonensis), Iringa Highlands and in Mikumi Nat. Park, resident on coast north of Dar es Salaam (N.E. Baker, pers. comm.). Angola, uncommon, Luanda, Huambo to SW Namibe, east to Cunene and along Namibian border to SE Cuando Cubango, and E Moxico. Zambia, in Northern Prov. from Mweru Marsh and Mbala through Kasama to S border of Bangweulu Swamp, partly sympatric with O. gabonensis; absent from most of N Zambia but widespread in S, north to Ndola, S Kabompo and Balovale (where overlaps O. gabonensis); absent from Luangwa and middle Zambezi ˆ i, Valleys; occurs in 34% of 300 Zambia squares. Malaw only in Karonga District. Botswana, locally common in N and E; common at Orapa and Jwaneng, very common at Phitsane Molopo; in SE, frequent to common, north to Salajwe; frequent in Makgadikgadi Pans region, though rare or absent in drought years when thought to move to central and W Kalahari and to W Zimbabwe. Elsewhere in southern Africa locally frequent to common as mapped, distribution centred on Etosha Pan, Namibia, central plateau of Zimbabwe and highveld grasslands of NE Cape Prov. and W KwaZulu-Natal, South Africa (Harrison et al. 1997). Population in Sul do Save, Mozambique, estimated to be >50,000 birds (Parker 1999). Description. O. f. muelleri Zedlitz (includes ‘bradfieldi’ and ˆ i, Zambia ‘minisucla’): central and S Kenya, Tanzania, Malaw (except range of smithersi), Angola, W Zimbabwe, Namibia, SE Botswana, N Cape Prov. and W Transvaal (South Africa). ADULT Y: upperparts greyish brown with diffuse broad dark brown streaks, grading to blackish on front of forehead. Tail short, blackish brown; T6 with white tip and wedge along inner web and white outer web, T5 (and often T4) with white tip. Lores dark grey-brown; cheeks, ear-coverts and sides of neck greybrown; broad white lines above and below lores and around eye. Chin and throat blackish brown, with small white patch on upper chin at base of bill; upper breast boldly and narrowly barred

Ortygospiza fuscocrissa

?

? ? ?

blackish and white; sides of breast and flanks with more widely spaced blackish and white barring, tinged tawny, each feather crossed by a black bordered white bar with another white or pale tawny bar near base; lower breast deep tawny brown, grading to pale tawny buff on belly and undertail-coverts, the last with blackish brown bases and edges. Upperwing dark grey-brown, tertials and greater, median and lesser coverts broadly edged paler greyish brown. Underwing-coverts and axillaries pale tawny buff. Bill dark red, upper mandible blackish or brown in non-breeding season; eyes light ochreous brown; legs light brown to pinkish flesh. ADULT X: differs from ad. Y in having head and neck greyish brown, top of head with darker brown mottling; white lines above and below lores and around eye narrower; chin patch buffy white; barring on underparts browner, forming narrower band across upper breast. SIZE (8 YY, 6 XX): wing, Y 53–56 (548), X 52–55 (535); tail, Y 24–28 (269), X 23–27 (257); bill, Y 95–105 (103), X 95–105 (101); tarsus, Y 14–155 (149), X 14–15 (147). IMMATURE: juv. similar above to ad. X; tawny buff below, with paler chin and throat, upper breast faintly barred and flanks more strongly barred dark brown; upperwing-coverts fringed buff. NESTLING (O. f. digressa): skin pink-orange; eyes dark and bulbous; long white or grey-white tufts of down on supraorbital, occipital, spinal, humeral and femoral feather tracts; inside mandible orange-pink, palate very pale pink with ring of 6 small black spots; when bill closed, gape tubercles appear as alternating

388

ESTRILDIDAE

LAYING DATES: NE Zaı ¨re, (juvs Sept); Uganda, Aug; Zambia, Chingola Mar–Apr (when no evidence of nesting at Kitwe), elsewhere Jan–Feb (and active gonads Mar).

INCUBATION:

Key References

evidence that both sexes incubate. Penry, E.H. (1986), Traylor, M.A. (1963).

Plate 21

Ortygospiza fuscocrissa Heuglin. African Quailfinch. Astrild-caille a ` lunettes.

(Opp. p. 330)

Ortygospiza fuscocrissa Vieillot, 1863. J. Orn., 11, p. 18; Dembea and Tigre´, Abyssinia. Forms a superspecies with O. atricollis and O. gabonensis. Range and Status. Endemic resident and nomad, Eritrea to South Africa. Eritrea, uncommon and sporadic (possibly migratory – not seen in winter) above 1830 m. Ethiopia, frequent to common above 1500 m and up to at least 2440 m, north to L. Tana, east to Somali border, south to Alghe; also in Rift Valley. Somalia, old record from Hargeysa. Kenya, frequent but local, above 1500 m; commonest in Laikipia Plateau, Nairobi Nat. Park, Mara Game Res., L. Baringo, L. Nakuru; up to 3000 m in Mau Narok; rare as far east as Tsavo East. Tanzania, locally abundant from Kenya border to 3
300 S between 34
300 E and 37
300 E; rare or absent in interior; scarce on Ufipa Plateau (where some observations may refer to Black-chinned Quailfinch O. gabonensis), Iringa Highlands and in Mikumi Nat. Park, resident on coast north of Dar es Salaam (N.E. Baker, pers. comm.). Angola, uncommon, Luanda, Huambo to SW Namibe, east to Cunene and along Namibian border to SE Cuando Cubango, and E Moxico. Zambia, in Northern Prov. from Mweru Marsh and Mbala through Kasama to S border of Bangweulu Swamp, partly sympatric with O. gabonensis; absent from most of N Zambia but widespread in S, north to Ndola, S Kabompo and Balovale (where overlaps O. gabonensis); absent from Luangwa and middle Zambezi ˆ i, Valleys; occurs in 34% of 300 Zambia squares. Malaw only in Karonga District. Botswana, locally common in N and E; common at Orapa and Jwaneng, very common at Phitsane Molopo; in SE, frequent to common, north to Salajwe; frequent in Makgadikgadi Pans region, though rare or absent in drought years when thought to move to central and W Kalahari and to W Zimbabwe. Elsewhere in southern Africa locally frequent to common as mapped, distribution centred on Etosha Pan, Namibia, central plateau of Zimbabwe and highveld grasslands of NE Cape Prov. and W KwaZulu-Natal, South Africa (Harrison et al. 1997). Population in Sul do Save, Mozambique, estimated to be >50,000 birds (Parker 1999). Description. O. f. muelleri Zedlitz (includes ‘bradfieldi’ and ˆ i, Zambia ‘minisucla’): central and S Kenya, Tanzania, Malaw (except range of smithersi), Angola, W Zimbabwe, Namibia, SE Botswana, N Cape Prov. and W Transvaal (South Africa). ADULT Y: upperparts greyish brown with diffuse broad dark brown streaks, grading to blackish on front of forehead. Tail short, blackish brown; T6 with white tip and wedge along inner web and white outer web, T5 (and often T4) with white tip. Lores dark grey-brown; cheeks, ear-coverts and sides of neck greybrown; broad white lines above and below lores and around eye. Chin and throat blackish brown, with small white patch on upper chin at base of bill; upper breast boldly and narrowly barred

Ortygospiza fuscocrissa

?

? ? ?

blackish and white; sides of breast and flanks with more widely spaced blackish and white barring, tinged tawny, each feather crossed by a black bordered white bar with another white or pale tawny bar near base; lower breast deep tawny brown, grading to pale tawny buff on belly and undertail-coverts, the last with blackish brown bases and edges. Upperwing dark grey-brown, tertials and greater, median and lesser coverts broadly edged paler greyish brown. Underwing-coverts and axillaries pale tawny buff. Bill dark red, upper mandible blackish or brown in non-breeding season; eyes light ochreous brown; legs light brown to pinkish flesh. ADULT X: differs from ad. Y in having head and neck greyish brown, top of head with darker brown mottling; white lines above and below lores and around eye narrower; chin patch buffy white; barring on underparts browner, forming narrower band across upper breast. SIZE (8 YY, 6 XX): wing, Y 53–56 (548), X 52–55 (535); tail, Y 24–28 (269), X 23–27 (257); bill, Y 95–105 (103), X 95–105 (101); tarsus, Y 14–155 (149), X 14–15 (147). IMMATURE: juv. similar above to ad. X; tawny buff below, with paler chin and throat, upper breast faintly barred and flanks more strongly barred dark brown; upperwing-coverts fringed buff. NESTLING (O. f. digressa): skin pink-orange; eyes dark and bulbous; long white or grey-white tufts of down on supraorbital, occipital, spinal, humeral and femoral feather tracts; inside mandible orange-pink, palate very pale pink with ring of 6 small black spots; when bill closed, gape tubercles appear as alternating

Ortygospiza fuscocrissa presumed Y (‘contact phrases’: Nuttall 1993), a series of dry notes strung out unevenly, ‘tyerk-tyik-tyok-tyerktikree-tikrok . . . ’, or in shorter repeated phrases, ‘tuk, tuk-ree, tuk-roo’ or ‘klee-to-klee, kler-to-klee’; X’s phrases distinguishable, usually just 4 notes, ‘gru grie grit-ty’. True song used by Y during courtship is low-intensity ‘burbling’: staccato notes uttered in random sequence and covering broad frequency range (Nuttall 1993). For sonagrams and full discussion of these and other calls and their contexts, see Nuttall (1993). pale purplish blue and black areas; when bill open they appear as distinct black and white spots along edges of gape. At 4–5 days, front end of maxilla/palate dark, 2 halves separated by pale area; inside of mandible dark near tip, with thin black V mark just inside; tongue arrow-shaped, pink, with 3 black patches in triangle, the front one (tip of tongue) largest; gullet orange-pink (fig. p. 252). Large, fleshy, pale pink swellings at corner of mouth, 2 at each side; in front of them, maxilla has 2 white swellings on black background and mandible has single white swelling on black background (A – Nuttall 1992); when bill closed white mandible swelling falls below black gap between 2 maxillary swellings, forming checkerboard pattern. O. f. fuscocrissa Heuglin: Eritrea and Ethiopia. Upperparts slightly darker and browner than in muelleri; dark barring on breast and flank feathers browner (less black) and band of barring across upper breast somewhat narrower; belly deeper tawny. Larger: wing, Y (n ¼ 3) 55–59 (570), X (n ¼ 4) 53–56 (550). O. f. smithersi Benson: NE Zambia (Northern Prov. except Mbala district). Darker above than muelleri, blacker on cheeks, with narrower white ring round eye. O. f. digressa Clancey: E Zimbabwe, S Mozambique, South Africa (except N Cape Prov. and W Transvaal), Swaziland and Lesotho. Like muelleri, pale and well-mottled above, but slightly browner. WEIGHT: (southern African races) Y (n ¼ 4) 105–120 (114), X (n ¼ 8) 100–115 (106). O. f. pallida Roberts: Makgadigadi Pans, N Botswana; migrant to adjacent NW Zimbabwe. Greyer than muelleri, paler throughout. TAXONOMIC NOTE: see generic diagnosis.

Field Characters. Length 9–10 cm. Tiny; keeps to ground in grassland. Seldom seen until flushed, when it rises steeply and erratically with characteristic call; plump and short-tailed in silhouette. Plain grey wings distinguish it in flight from orange-winged Locust-Finch Paludipasser locustella. Conspicuous zebra barring on breast and flanks; underparts vary from white with touch of orange on breast to mainly orange. Separated from the other 2 quailfinches by white spectacles (both sexes), small white patch on chin; variable amount of black on face of Y, none on X; rump grey (red in X Zebra Waxbill Sporaeginthus subflavus and Locust-Finch); immature like X but barring faint (heavy in immature Locust-Finch, which also has orange-brown wings; young Orange-breasted Waxbill yellowish below with no barring). Voice. Tape-recorded (86, 88, 99, B, F, LEM). Common calls given on take-off and in flight have a clipped, dry, hard quality difficult to describe but unlike that of any other bird, enabling instant identification, ‘tyuk’, tyek’, ‘puk’, ‘pik’; have been transcribed as ‘djink’ or ‘tirrilink’ and said to be tinny or bell-like, but they lack any ringing quality and are better described as a ‘hard, grating chirp’ (R.J. Stjernstedt in Penry 1986). Song-like utterances of

General Habits. Inhabits short grass and sorrels near highland streams, also farmyards where grain has been threshed (Eritrea); dry or wet swamps with short grass, often patchy, tufty and wiry, especially on flat, black cotton soils (Kenya); grassland and short-grass dambos in woodland (Angola), standing cereal crops, short grass on airports; coastal saltflats (north of Dar es Salaam); short grassland close to water, as around dams and stock farms; fallow fields, heavily cropped pasture; low coastal hills (South Africa); floodplains; vleis and surrounding sedges; herbage on heavy, muddy, trampled soil (Harrison et al. 1997). In pairs and all year in parties of 4–30. Entirely terrestrial. Shy, retiring and secretive. When foraging, hop-runs erratically, pecking soil surface for seeds. Sometimes flutters up to take flying termites. Adept at vertical take-off and landing. Flies high between feeding grounds. If alarmed, crouches flat against earth; takes flight only as a last resort; when flushed, almost from underfoot, small flock rises steeply, fast and rather jerkily, flies short distance, plummets back to ground and immediately disappears behind vegetation, whether thick or scant. Bird flushed from nest flies up and up in circles until almost lost to view. Regularly drinks at margins of rain puddles and lakes. Wanders widely and has been regarded as migratory (e.g. Ethiopia, NE Botswana), but no clear evidence of regular movements in southern Africa; where changes in abundance reported, probably a result of incidental nomadic dispersion and seasonal differences in conspicuousness (calling and flocking in family parties: R.J. Nuttall in Harrison et al. 1997). Food. Grass seeds, small insects, spiders. In captivity, seeds of Eragrostis tef and Panicum maximum, and variety of invertebrates. Breeding Habits. Solitary nester; monogamous, but in Kenya may be polygamous, single Y with 2 XX and 2 nests only 6 m apart (van Someren 1956); 2 pairs with nests 11 m apart (Zimbabwe, Vincent 1949); not obviously territorial. Displaying Y flies high then plunges to ground with clicking song. NEST: spherical or oval ball with flattened base, loosely made of dry grass blades and fine grass rootlets, with side entrance; withered grey grass blades outside; thinly or thickly lined with very delicate grass, fluffy seed-heads, and feathers. Situated on ground at base of tuft of grass or between 2 close-together tufts. Usually a small, bare area of ground in front of entrance (function: see Nuttall 1992).

389

390

ESTRILDIDAE

EGGS: 3–6 (av. 42 in one study, 47 in another). Laid at 1-day intervals. Pure white, almost spherical. SIZE: (n ¼ 115) 127–165  104–129 (144  111). LAYING DATES: Eritrea, June; Ethiopia, July; E Africa, Region C, Mar, June, Region D, Apr–July; Zambia, Jan– Mar, June; Zimbabwe, Dec–May, mainly Jan–Apr; South Africa, Transvaal (Barberspan, n ¼ 171), Nov–Sept, 81% in Jan–May and peak in Mar, (elsewhere, n ¼ 132), Sept, Dec– Jan, 59% in Jan–Feb; Free State, Oct–Feb; E Cape Prov. and KwaZulu-Natal, Nov–Apr, mainly Jan–Mar. INCUBATION: begins with 3rd or 4th egg laid; by Y and X in turns, sometimes both together; both sexes turn the eggs. Nest relief follows no consistent pattern. Incubating Y responds to silent approach of X by uttering lowintensity ‘burbling’ (Y’s high-intensity ‘burbling’, or song, has no territorial function but serves to strengthen pair bond: Nuttall 1992). Period 14–16 days. DEVELOPMENT AND CARE OF YOUNG: clutch hatches over period of 2–4 days. Young brooded by Y and X separately and together. Parents remove faecal sacs, sometimes

directly from nestlings’ cloaca, sometimes eating them, but nest soon becomes fouled with faeces. Development described by Nuttall (1992); eyes start to open on day 6, fully open by day 10; feathers on head are the last to emerge; fully feathered by day 17. Young utter begging calls, soft, barely audible wheezes, from 4 days; when disturbed by person, they threaten with harsh ‘tcheee tcheee’ calls from 6 days. Nestling period: 17–21 days (Brickell and Konigkramer 1997) or 18–19 days (Nuttall 1992); at 16 days brood scrambles out of nest and scatters into grass, returning to nest at night, scattering again on 17th day and flying a few m. BREEDING SUCCESS/SURVIVAL: nestlings taken by snakes (van Someren 1956) including Lamprophis fuliginosus (Nuttall 1992). Of 14 eggs laid in captivity, 8 hatched and 4 produced fledglings. Key References Nuttall, R.J. (1992, 1993), van Someren, V.G.L. (1956)., Traylor, M.A. (1963), Vincent, A.W. (1949).

Genus Paludipasser Neave

Endemic. Single species of tiny quailfinch. Somewhat cryptic, plumage of X like Ortygospiza but Y blackish with red face, rump and wings. Bill deep, with strongly ridged culmen; legs stout and muscular, lark-like long hind toe and long hind claw, only slightly curved; tail very short; breast muscles well developed, sternum exceptionally long. Exclusively ground feeding and nesting. Gregarious. Not at all closely related with Ortygospiza and, in fact, so distantly related as to form a sister clade to remainder of the family Estrildidae (R. B. Payne, pers. comm.).

Plate 21

Paludipasser locustella Neave. Locust-Finch. Astrild-caille a` gorge rouge.

(Opp. p. 330)

Paludipasser locustella Neave, 1909. Bull. Br. Orn. Club, 25, p. 25; Luansenshi R., near L. Bangweulu. Range and Status. Endemic resident and wanderer. Nigeria, 2 breeding pairs, Rockwater Fish Farm, Jos Plateau, Oct–Nov 2002, and flock of 10 in Dec (R. McGregor and J. Wilson, pers. comm.). Cameroon, records Boumba-Bek and Lobe´ke´ (Dowsett and DowsettLemaire 2000). Gabon, Le´koni and possibly between Gamba and Tchibanga (Sargeant 1993). Congo, Le´fini, and on Congo and Zaı¨re sides of Congo R. to east. Zaı¨re, uncommon, from Kotli east to Sudan border from opposite Bengengai (Sudan) to opposite Aloma Plateau (Sudan) though not yet found on Sudan side; Faradje, Garamba, Niangara, Kotili, Mauda. Kenya, 4 recent records at Mungatsi and Alupe. Tanzania, Ufipa Plateau and highˆ i. Angola, lands from south of Iringa town to L. Malaw uncommon: records from Malanje, Huı´la, Lunda Sul and Moxico. Zambia, widespread but rather local in Northern, Western and North-Western Provs; absent from drier country in S but extends to Lusaka, Choma, Luamala R. at

14
060 S, 27
080 E, Ndulumina in Kafue Nat. Park, Ntemwa, Mankoya at 14
500 S, 25
150 E, South Luete R., and Lundazi and Chipata; occurs in 35% of 300 Zambia squares. ˆ i, widespread at 900–1675 m south to Dzonze and Malaw in Rift Valley between Nkhotakota and Chintheche, also east of Rift near Namwera. Zimbabwe, extremely local, generally uncommon; Mashonaland Plateau from Rusape and Timaru at 1700 m to Harare and Umvukwe Range; in S, at Lalapanzi, Umvuma, Chatsworth and Fort Victoria. Botswana, twice on Shindi airstrip in Okavango Delta. Description. O. l. locustella Neave: south of 7
S. ADULT Y: centre of forehead to hindneck and sides of neck brownish black, pale brown feather fringes giving slightly streaked or spotted effect; mantle, scapulars, back and centre of rump and uppertail-coverts brownish black, each feather with small pointed subterminal white spot; sides of rump and uppertail-coverts bright red. Tail blackish, T6 with fine whitish fringe at tip. Sides of forehead, lores, supercilium, ear-coverts, cheeks and chin to upper breast red, deep scarlet on face, paler, more orange-red on breast. Rest of

390

ESTRILDIDAE

EGGS: 3–6 (av. 42 in one study, 47 in another). Laid at 1-day intervals. Pure white, almost spherical. SIZE: (n ¼ 115) 127–165  104–129 (144  111). LAYING DATES: Eritrea, June; Ethiopia, July; E Africa, Region C, Mar, June, Region D, Apr–July; Zambia, Jan– Mar, June; Zimbabwe, Dec–May, mainly Jan–Apr; South Africa, Transvaal (Barberspan, n ¼ 171), Nov–Sept, 81% in Jan–May and peak in Mar, (elsewhere, n ¼ 132), Sept, Dec– Jan, 59% in Jan–Feb; Free State, Oct–Feb; E Cape Prov. and KwaZulu-Natal, Nov–Apr, mainly Jan–Mar. INCUBATION: begins with 3rd or 4th egg laid; by Y and X in turns, sometimes both together; both sexes turn the eggs. Nest relief follows no consistent pattern. Incubating Y responds to silent approach of X by uttering lowintensity ‘burbling’ (Y’s high-intensity ‘burbling’, or song, has no territorial function but serves to strengthen pair bond: Nuttall 1992). Period 14–16 days. DEVELOPMENT AND CARE OF YOUNG: clutch hatches over period of 2–4 days. Young brooded by Y and X separately and together. Parents remove faecal sacs, sometimes

directly from nestlings’ cloaca, sometimes eating them, but nest soon becomes fouled with faeces. Development described by Nuttall (1992); eyes start to open on day 6, fully open by day 10; feathers on head are the last to emerge; fully feathered by day 17. Young utter begging calls, soft, barely audible wheezes, from 4 days; when disturbed by person, they threaten with harsh ‘tcheee tcheee’ calls from 6 days. Nestling period: 17–21 days (Brickell and Konigkramer 1997) or 18–19 days (Nuttall 1992); at 16 days brood scrambles out of nest and scatters into grass, returning to nest at night, scattering again on 17th day and flying a few m. BREEDING SUCCESS/SURVIVAL: nestlings taken by snakes (van Someren 1956) including Lamprophis fuliginosus (Nuttall 1992). Of 14 eggs laid in captivity, 8 hatched and 4 produced fledglings. Key References Nuttall, R.J. (1992, 1993), van Someren, V.G.L. (1956)., Traylor, M.A. (1963), Vincent, A.W. (1949).

Genus Paludipasser Neave

Endemic. Single species of tiny quailfinch. Somewhat cryptic, plumage of X like Ortygospiza but Y blackish with red face, rump and wings. Bill deep, with strongly ridged culmen; legs stout and muscular, lark-like long hind toe and long hind claw, only slightly curved; tail very short; breast muscles well developed, sternum exceptionally long. Exclusively ground feeding and nesting. Gregarious. Not at all closely related with Ortygospiza and, in fact, so distantly related as to form a sister clade to remainder of the family Estrildidae (R. B. Payne, pers. comm.).

Plate 21

Paludipasser locustella Neave. Locust-Finch. Astrild-caille a` gorge rouge.

(Opp. p. 330)

Paludipasser locustella Neave, 1909. Bull. Br. Orn. Club, 25, p. 25; Luansenshi R., near L. Bangweulu. Range and Status. Endemic resident and wanderer. Nigeria, 2 breeding pairs, Rockwater Fish Farm, Jos Plateau, Oct–Nov 2002, and flock of 10 in Dec (R. McGregor and J. Wilson, pers. comm.). Cameroon, records Boumba-Bek and Lobe´ke´ (Dowsett and DowsettLemaire 2000). Gabon, Le´koni and possibly between Gamba and Tchibanga (Sargeant 1993). Congo, Le´fini, and on Congo and Zaı¨re sides of Congo R. to east. Zaı¨re, uncommon, from Kotli east to Sudan border from opposite Bengengai (Sudan) to opposite Aloma Plateau (Sudan) though not yet found on Sudan side; Faradje, Garamba, Niangara, Kotili, Mauda. Kenya, 4 recent records at Mungatsi and Alupe. Tanzania, Ufipa Plateau and highˆ i. Angola, lands from south of Iringa town to L. Malaw uncommon: records from Malanje, Huı´la, Lunda Sul and Moxico. Zambia, widespread but rather local in Northern, Western and North-Western Provs; absent from drier country in S but extends to Lusaka, Choma, Luamala R. at

14
060 S, 27
080 E, Ndulumina in Kafue Nat. Park, Ntemwa, Mankoya at 14
500 S, 25
150 E, South Luete R., and Lundazi and Chipata; occurs in 35% of 300 Zambia squares. ˆ i, widespread at 900–1675 m south to Dzonze and Malaw in Rift Valley between Nkhotakota and Chintheche, also east of Rift near Namwera. Zimbabwe, extremely local, generally uncommon; Mashonaland Plateau from Rusape and Timaru at 1700 m to Harare and Umvukwe Range; in S, at Lalapanzi, Umvuma, Chatsworth and Fort Victoria. Botswana, twice on Shindi airstrip in Okavango Delta. Description. O. l. locustella Neave: south of 7
S. ADULT Y: centre of forehead to hindneck and sides of neck brownish black, pale brown feather fringes giving slightly streaked or spotted effect; mantle, scapulars, back and centre of rump and uppertail-coverts brownish black, each feather with small pointed subterminal white spot; sides of rump and uppertail-coverts bright red. Tail blackish, T6 with fine whitish fringe at tip. Sides of forehead, lores, supercilium, ear-coverts, cheeks and chin to upper breast red, deep scarlet on face, paler, more orange-red on breast. Rest of

390

ESTRILDIDAE

EGGS: 3–6 (av. 42 in one study, 47 in another). Laid at 1-day intervals. Pure white, almost spherical. SIZE: (n ¼ 115) 127–165  104–129 (144  111). LAYING DATES: Eritrea, June; Ethiopia, July; E Africa, Region C, Mar, June, Region D, Apr–July; Zambia, Jan– Mar, June; Zimbabwe, Dec–May, mainly Jan–Apr; South Africa, Transvaal (Barberspan, n ¼ 171), Nov–Sept, 81% in Jan–May and peak in Mar, (elsewhere, n ¼ 132), Sept, Dec– Jan, 59% in Jan–Feb; Free State, Oct–Feb; E Cape Prov. and KwaZulu-Natal, Nov–Apr, mainly Jan–Mar. INCUBATION: begins with 3rd or 4th egg laid; by Y and X in turns, sometimes both together; both sexes turn the eggs. Nest relief follows no consistent pattern. Incubating Y responds to silent approach of X by uttering lowintensity ‘burbling’ (Y’s high-intensity ‘burbling’, or song, has no territorial function but serves to strengthen pair bond: Nuttall 1992). Period 14–16 days. DEVELOPMENT AND CARE OF YOUNG: clutch hatches over period of 2–4 days. Young brooded by Y and X separately and together. Parents remove faecal sacs, sometimes

directly from nestlings’ cloaca, sometimes eating them, but nest soon becomes fouled with faeces. Development described by Nuttall (1992); eyes start to open on day 6, fully open by day 10; feathers on head are the last to emerge; fully feathered by day 17. Young utter begging calls, soft, barely audible wheezes, from 4 days; when disturbed by person, they threaten with harsh ‘tcheee tcheee’ calls from 6 days. Nestling period: 17–21 days (Brickell and Konigkramer 1997) or 18–19 days (Nuttall 1992); at 16 days brood scrambles out of nest and scatters into grass, returning to nest at night, scattering again on 17th day and flying a few m. BREEDING SUCCESS/SURVIVAL: nestlings taken by snakes (van Someren 1956) including Lamprophis fuliginosus (Nuttall 1992). Of 14 eggs laid in captivity, 8 hatched and 4 produced fledglings. Key References Nuttall, R.J. (1992, 1993), van Someren, V.G.L. (1956)., Traylor, M.A. (1963), Vincent, A.W. (1949).

Genus Paludipasser Neave

Endemic. Single species of tiny quailfinch. Somewhat cryptic, plumage of X like Ortygospiza but Y blackish with red face, rump and wings. Bill deep, with strongly ridged culmen; legs stout and muscular, lark-like long hind toe and long hind claw, only slightly curved; tail very short; breast muscles well developed, sternum exceptionally long. Exclusively ground feeding and nesting. Gregarious. Not at all closely related with Ortygospiza and, in fact, so distantly related as to form a sister clade to remainder of the family Estrildidae (R. B. Payne, pers. comm.).

Plate 21

Paludipasser locustella Neave. Locust-Finch. Astrild-caille a` gorge rouge.

(Opp. p. 330)

Paludipasser locustella Neave, 1909. Bull. Br. Orn. Club, 25, p. 25; Luansenshi R., near L. Bangweulu. Range and Status. Endemic resident and wanderer. Nigeria, 2 breeding pairs, Rockwater Fish Farm, Jos Plateau, Oct–Nov 2002, and flock of 10 in Dec (R. McGregor and J. Wilson, pers. comm.). Cameroon, records Boumba-Bek and Lobe´ke´ (Dowsett and DowsettLemaire 2000). Gabon, Le´koni and possibly between Gamba and Tchibanga (Sargeant 1993). Congo, Le´fini, and on Congo and Zaı¨re sides of Congo R. to east. Zaı¨re, uncommon, from Kotli east to Sudan border from opposite Bengengai (Sudan) to opposite Aloma Plateau (Sudan) though not yet found on Sudan side; Faradje, Garamba, Niangara, Kotili, Mauda. Kenya, 4 recent records at Mungatsi and Alupe. Tanzania, Ufipa Plateau and highˆ i. Angola, lands from south of Iringa town to L. Malaw uncommon: records from Malanje, Huı´la, Lunda Sul and Moxico. Zambia, widespread but rather local in Northern, Western and North-Western Provs; absent from drier country in S but extends to Lusaka, Choma, Luamala R. at

14
060 S, 27
080 E, Ndulumina in Kafue Nat. Park, Ntemwa, Mankoya at 14
500 S, 25
150 E, South Luete R., and Lundazi and Chipata; occurs in 35% of 300 Zambia squares. ˆ i, widespread at 900–1675 m south to Dzonze and Malaw in Rift Valley between Nkhotakota and Chintheche, also east of Rift near Namwera. Zimbabwe, extremely local, generally uncommon; Mashonaland Plateau from Rusape and Timaru at 1700 m to Harare and Umvukwe Range; in S, at Lalapanzi, Umvuma, Chatsworth and Fort Victoria. Botswana, twice on Shindi airstrip in Okavango Delta. Description. O. l. locustella Neave: south of 7
S. ADULT Y: centre of forehead to hindneck and sides of neck brownish black, pale brown feather fringes giving slightly streaked or spotted effect; mantle, scapulars, back and centre of rump and uppertail-coverts brownish black, each feather with small pointed subterminal white spot; sides of rump and uppertail-coverts bright red. Tail blackish, T6 with fine whitish fringe at tip. Sides of forehead, lores, supercilium, ear-coverts, cheeks and chin to upper breast red, deep scarlet on face, paler, more orange-red on breast. Rest of

Paludipasser locustella side of palate, nearly joining in front, palate between them bright red; at rear end of the line, another curved black line at right angles with it, near gape lobes; red of palate extends down throat; tongue with 3 raised red lobes overlapping edge, posterior pair lozenge-shaped, placed together on midline, single anterior lobe a little larger and heart-shaped (Irwin 1958). O. l. uelensis Chapin: W Congo to Central Nigeria and W Kenya. Lacks pale spots and bars on upperparts, flanks and tertials. Bill of Y dark above, red confined to cutting edge and lower mandible.

Paludipasser locustella

?

underparts dark sooty grey, lower flank feathers with narrow subterminal white bars and spots, undertail-coverts with narrow whitish fringes. Remiges dark brown, outer primaries narrowly edged golden buff, inner primaries and secondaries edged orange or orange-red, forming panel on closed wing, outer edges of tertials finely spotted with white; primary coverts and alula blackish brown, finely fringed golden buff, the latter with orangered outer webs; greater coverts blackish with broad orange-red outer edges and tips; median and lesser coverts orange, these and greater coverts all with small white shaft spot near tip. Upperwing-coverts form large orange panel on closed wing continuous with panel on remiges. Underwing-coverts and axillaries creamy white. Bill orange-scarlet with blackish culmen; eyes pale yellow; legs pale dull brown, yellowish or flesh brown. ADULT X: forehead to hindneck dark brown, grey-brown feather fringes forming pale narrow streaks; mantle, scapulars, back and centre of rump and uppertail-coverts dark brown, finely spotted with white; sides of rump and uppertail-coverts red. Tail as ad. Y. Whitish or buff spot on lores; supercilium, ear-coverts and cheeks dark brown. Chin to breast pale grey-buff with diffuse greyish mottling; sides of breast narrowly barred greyish and pale buff, rest of underparts sooty brown, flanks barred and undertailcoverts finely fringed with white. Upperwing feathers dark brown; remiges narrowly fringed golden buff to form buffy brown (not orange) panel on closed wing; primary coverts and alula finely fringed buff; greater coverts edged and tipped orange, median and lesser coverts broadly tipped orange; orange wing-covert panel thus less red, less intense than in ad. Y. Bare parts as ad. Y. SIZE: wing, Y (n ¼ 10) 42–47 (441), X (n ¼ 10) 42–48 (444); tail, Y (n ¼ 10) 23–27 (252), X (n ¼ 10) 24–27 (257); bill, Y (n ¼ 10) 95–105 (99), X (n ¼ 8) 95–10 (97); tarsus, Y (n ¼ 10) 14–15 (146), X (n ¼ 10) 14–155 (146). IMMATURE: juv. similar to ad. X, but lacks red or orange on sides of rump and wings; dark brown upperparts with few white spots, and streaked throughout with warm buffish brown; deeper tawny-buff below, throat and breast mottled and barred dark sepia, flanks barred dark brown and buff; edges and tips of tertials and upperwing-coverts tawny buff. Bill blackish; eyes brown. NESTLING: mouth at c. 6 days: 2 flat red lobes on each side protrude slightly from corner of mouth; curved black line on each

Field Characters. Length 9–10 cm. A tiny, short-tailed grassland bird; usually on ground and not seen until flushed, when easily identified by orange wing patches. Y bright red from face to breast, with red sides of rump and red bill, otherwise blackish; X has black face and white underparts with zebra barring on sides; white spots on upperparts in nominate race (S-central Africa). Quailfinches are same shape and found in same habitat, but in flight show uniformly dark wings. Immature mottled brown above; differs from young quailfinches in warm brown wings, blacker head and heavy barring on sides. Voice. Tape-recorded (86, 99, 104, B, GIB, LEM, STJ). Various brief calls, ‘tissep’, ‘chit’, ‘cheep’, ‘tsee-tset’, highpitched ‘see-see-see’, rattling ‘chichichichichit’, also downslurred ‘chyew’ or ‘tyee-tyee-tyee’. Flight call a hard ‘jee’ or down-slurred ‘jeea’ (sonagram in Penry 1986, where it is transcribed ‘drink drink’, even though it has no ringing quality). General Habits. Inhabits flat, open grassland with tufts of short, wiry grass interspersed with bare patches of waterlogged ground on sandveld rather than clayey soils (Zimbabwe); short-grass dambos and vleis running down through light Brachystegia woods with Parinari mobola and Syzygium guineense trees (Zimbabwe); low marshy spots that have dried out and been burnt (Uele, Zaı¨re); abandoned cultivation in highlands. Occurs in pairs and flocks of up to 10, once 16. Entirely terrestrial; forages on ground between tufts of grass or burned tussocks; never perches on vegetation. When disturbed, at first squats down motionless then rises suddenly and quickly, mounting steadily to some height and flying directly or erratically for 45–90 m before alighting; on ground moves away immediately, out of sight amongst grasses. Flock rises into air with faint whirr of wings and flies off in a body. Flight steady and direct. Rather silent. 5 birds once found roosting in a nest – probably a family. Food. Small grass seeds. Breeding Habits. Solitary nester. NEST: quite substantial ball of soft, fine dry grass, put together loosely but thickly, particularly in basal half, with circular entrance high on side; inside of shell made of short strips of grass stems, thought to be added when green, during early stages of incubation (Irwin 1958); lined with grass inflorescences, fine green grass, sometimes with feathers, once with vegetable down. Well hidden in dense, knee-high, fine wiry grass, tightly wedged between many stems but not attached to them, on ground between tufts of

391

392

ESTRILDIDAE

grass on wet, damp or dried-out soil. Nests in same area as nesting African Quailfinches Ortygospiza fuscocrissa but in even wetter places where grass is less even and more tufty. EGGS: 2–8, usually 4–7, av. (11 clutches) 60. Pure white. SIZE (n = 2): 122  99, 129  100. LAYING DATES: Nigeria, Oct (and fledglings late Oct: R. McGregor and J. Wilson, pers. comm.), Zambia, ˆ i, Jan–Feb; Jan, May (and gonads active Dec); Malaw Mozambique, Mar; Zimbabwe, Jan–May (15 clutches), mainly Jan (8 clutches).

INCUBATION: mainly by X, beginning 3–4 days before completion of larger clutch. DEVELOPMENT AND CARE OF YOUNG: young fed mainly by X. Both sexes incubate and care for young before and after they fledge (R. McGregor and J. Wilson, pers. comm.). Y and X roost in nest at night.

Key References Chapin, J.P. (1954), Irwin, M.P.S. (1958), Maclean, G.L. (1993), Vincent, A.W. (1949).

Genus Sporaeginthus Cabanis

One tiny weaver-finch, S. subflavus, in Africa and SW Arabia and a small one, S. formosa, in N India. YY yellow-olive with red bills, and banded sides of breast and flanks, XX like YY but duller; Y subflavus with red eyebrow and red rump, Y formosa without eyebrow and with green rump. Tail short or mid-length, well rounded. Forage on ground in or near rank grass, often by water. Formerly, generally placed with Oriental amandava in genus Amandava. A. amandava is similar in size and habitat (rank grass, reeds); nestlings have similar mouth patterns (Harrison 1962) and raise one or both wings when begging. However, subflavus and amandava have radically different plumage colours and patterns. Moreover, Sporaeginthus YY do not have non-breeding ‘eclipse’ plumage but amandava Y has; S. subflavus does not have structured song like that of amandava, and it does not hold grass stem in bill when displaying as amandava does. Molecular analysis shows that subflavus and amandava are quite distantly related and that, with formosa, they constitute a sister group of Ortygospiza (R. B. Payne, pers. comm.).

Plate 19 (Opp. p. 282)

Sporaeginthus subflavus (Vieillot). Zebra Waxbill; Goldbreast; Orange-breasted Waxbill. Bengali ze´bre ´. Fringilla subflava Vieillot, 1819. Nouv. Dict. Hist. Nat., nouv. ´ed., 30, p. 575; Senegal. Range and Status. Resident, sub-Saharan Africa and Tihamah region, Yemen. Absent from arid and forested regions; localized in much of W Africa; more widespread east of 25
E. Mauritania, 1 Rosso (Bull. Afr. Bird Club 7, 2000, 75). Senegal and Gambia, rare; recent records near RichardToll, near Dakar, in Parc Nat. du Delta du Saloum, south of Tambacounda, near Ke´dougou, in Banjul-Cape St Mary area and near Kaur and Sapu; in both countries formerly locally common and has become rare because of habitat destruction by rice cultivation. Guinea-Bissau, frequent Bula area (Rodwell 1996). Guinea, Wassou and Kolente´ areas (Demey 1995), Haut Niger Nat. Park (where frequent: Nikolaus 2000) and Macenta. Sierra Leone, frequent, widespread in E, northwest to Kilimi area. Liberia, uncommon in N Lofa county, not uncommon at Yekepa and Nimba; once flock on Cape Mount Peninsula. Mali, occurrence uncertain; said to be uncommon south of 15
N and common at Kara (Lamarche 1981) but not in definitive list (Lamarche 1993). Burkino Faso, only record a flock at Koubri, Ouagadougou (Thonnerieux et al. 1989).

Ivory Coast, only in NE, from Korhogo to Comoe´ Nat. Park. Ghana, only 4 localities: 2 at Bolgatanga, 20 at Sugu, 40 or more regularly at Denu, and numerous at Atsutsuare (Grimes 1987). Togo, locally not uncommon in flocks of 20–100 in N in Domaine Gravillou-Kandjo-Kara area and in S in Tokpli-Lome´ area (Cheke and Walsh 1996). Benin, not yet found. Nigeria, widespread and locally common, north to Sokoto, Kano, Potiskum and Maiduguri, southwest to Oyo, Iwo and Ibadan, along Niger valley from Jebba to Onitsha, on Jos Plateau (many localities), southeast to Mambilla Plateau (Yelwa); see also remarks on seasonality, below. Cameroon, known from c. 25 localities (M. Languy, pers. comm.). Chad, Central African Republic and Sudan, uncommon and very local, except for Bangui region, Central African Republic, where frequent (6 localities: Germain and Cornet 1994). Ethiopia, locally frequent. Rest of Africa, uncommon to frequent and locally common to abundant, as mapped. Estimated to be 7000 birds in Swaziland and 5000 in Sul do Save, Mozambique, and 10–100 in Lesotho. Apparently less common in Transkei and Lesotho than formerly,

392

ESTRILDIDAE

grass on wet, damp or dried-out soil. Nests in same area as nesting African Quailfinches Ortygospiza fuscocrissa but in even wetter places where grass is less even and more tufty. EGGS: 2–8, usually 4–7, av. (11 clutches) 60. Pure white. SIZE (n = 2): 122  99, 129  100. LAYING DATES: Nigeria, Oct (and fledglings late Oct: R. McGregor and J. Wilson, pers. comm.), Zambia, ˆ i, Jan–Feb; Jan, May (and gonads active Dec); Malaw Mozambique, Mar; Zimbabwe, Jan–May (15 clutches), mainly Jan (8 clutches).

INCUBATION: mainly by X, beginning 3–4 days before completion of larger clutch. DEVELOPMENT AND CARE OF YOUNG: young fed mainly by X. Both sexes incubate and care for young before and after they fledge (R. McGregor and J. Wilson, pers. comm.). Y and X roost in nest at night.

Key References Chapin, J.P. (1954), Irwin, M.P.S. (1958), Maclean, G.L. (1993), Vincent, A.W. (1949).

Genus Sporaeginthus Cabanis

One tiny weaver-finch, S. subflavus, in Africa and SW Arabia and a small one, S. formosa, in N India. YY yellow-olive with red bills, and banded sides of breast and flanks, XX like YY but duller; Y subflavus with red eyebrow and red rump, Y formosa without eyebrow and with green rump. Tail short or mid-length, well rounded. Forage on ground in or near rank grass, often by water. Formerly, generally placed with Oriental amandava in genus Amandava. A. amandava is similar in size and habitat (rank grass, reeds); nestlings have similar mouth patterns (Harrison 1962) and raise one or both wings when begging. However, subflavus and amandava have radically different plumage colours and patterns. Moreover, Sporaeginthus YY do not have non-breeding ‘eclipse’ plumage but amandava Y has; S. subflavus does not have structured song like that of amandava, and it does not hold grass stem in bill when displaying as amandava does. Molecular analysis shows that subflavus and amandava are quite distantly related and that, with formosa, they constitute a sister group of Ortygospiza (R. B. Payne, pers. comm.).

Plate 19 (Opp. p. 282)

Sporaeginthus subflavus (Vieillot). Zebra Waxbill; Goldbreast; Orange-breasted Waxbill. Bengali ze´bre ´. Fringilla subflava Vieillot, 1819. Nouv. Dict. Hist. Nat., nouv. ´ed., 30, p. 575; Senegal. Range and Status. Resident, sub-Saharan Africa and Tihamah region, Yemen. Absent from arid and forested regions; localized in much of W Africa; more widespread east of 25
E. Mauritania, 1 Rosso (Bull. Afr. Bird Club 7, 2000, 75). Senegal and Gambia, rare; recent records near RichardToll, near Dakar, in Parc Nat. du Delta du Saloum, south of Tambacounda, near Ke´dougou, in Banjul-Cape St Mary area and near Kaur and Sapu; in both countries formerly locally common and has become rare because of habitat destruction by rice cultivation. Guinea-Bissau, frequent Bula area (Rodwell 1996). Guinea, Wassou and Kolente´ areas (Demey 1995), Haut Niger Nat. Park (where frequent: Nikolaus 2000) and Macenta. Sierra Leone, frequent, widespread in E, northwest to Kilimi area. Liberia, uncommon in N Lofa county, not uncommon at Yekepa and Nimba; once flock on Cape Mount Peninsula. Mali, occurrence uncertain; said to be uncommon south of 15
N and common at Kara (Lamarche 1981) but not in definitive list (Lamarche 1993). Burkino Faso, only record a flock at Koubri, Ouagadougou (Thonnerieux et al. 1989).

Ivory Coast, only in NE, from Korhogo to Comoe´ Nat. Park. Ghana, only 4 localities: 2 at Bolgatanga, 20 at Sugu, 40 or more regularly at Denu, and numerous at Atsutsuare (Grimes 1987). Togo, locally not uncommon in flocks of 20–100 in N in Domaine Gravillou-Kandjo-Kara area and in S in Tokpli-Lome´ area (Cheke and Walsh 1996). Benin, not yet found. Nigeria, widespread and locally common, north to Sokoto, Kano, Potiskum and Maiduguri, southwest to Oyo, Iwo and Ibadan, along Niger valley from Jebba to Onitsha, on Jos Plateau (many localities), southeast to Mambilla Plateau (Yelwa); see also remarks on seasonality, below. Cameroon, known from c. 25 localities (M. Languy, pers. comm.). Chad, Central African Republic and Sudan, uncommon and very local, except for Bangui region, Central African Republic, where frequent (6 localities: Germain and Cornet 1994). Ethiopia, locally frequent. Rest of Africa, uncommon to frequent and locally common to abundant, as mapped. Estimated to be 7000 birds in Swaziland and 5000 in Sul do Save, Mozambique, and 10–100 in Lesotho. Apparently less common in Transkei and Lesotho than formerly,

392

ESTRILDIDAE

grass on wet, damp or dried-out soil. Nests in same area as nesting African Quailfinches Ortygospiza fuscocrissa but in even wetter places where grass is less even and more tufty. EGGS: 2–8, usually 4–7, av. (11 clutches) 60. Pure white. SIZE (n = 2): 122  99, 129  100. LAYING DATES: Nigeria, Oct (and fledglings late Oct: R. McGregor and J. Wilson, pers. comm.), Zambia, ˆ i, Jan–Feb; Jan, May (and gonads active Dec); Malaw Mozambique, Mar; Zimbabwe, Jan–May (15 clutches), mainly Jan (8 clutches).

INCUBATION: mainly by X, beginning 3–4 days before completion of larger clutch. DEVELOPMENT AND CARE OF YOUNG: young fed mainly by X. Both sexes incubate and care for young before and after they fledge (R. McGregor and J. Wilson, pers. comm.). Y and X roost in nest at night.

Key References Chapin, J.P. (1954), Irwin, M.P.S. (1958), Maclean, G.L. (1993), Vincent, A.W. (1949).

Genus Sporaeginthus Cabanis

One tiny weaver-finch, S. subflavus, in Africa and SW Arabia and a small one, S. formosa, in N India. YY yellow-olive with red bills, and banded sides of breast and flanks, XX like YY but duller; Y subflavus with red eyebrow and red rump, Y formosa without eyebrow and with green rump. Tail short or mid-length, well rounded. Forage on ground in or near rank grass, often by water. Formerly, generally placed with Oriental amandava in genus Amandava. A. amandava is similar in size and habitat (rank grass, reeds); nestlings have similar mouth patterns (Harrison 1962) and raise one or both wings when begging. However, subflavus and amandava have radically different plumage colours and patterns. Moreover, Sporaeginthus YY do not have non-breeding ‘eclipse’ plumage but amandava Y has; S. subflavus does not have structured song like that of amandava, and it does not hold grass stem in bill when displaying as amandava does. Molecular analysis shows that subflavus and amandava are quite distantly related and that, with formosa, they constitute a sister group of Ortygospiza (R. B. Payne, pers. comm.).

Plate 19 (Opp. p. 282)

Sporaeginthus subflavus (Vieillot). Zebra Waxbill; Goldbreast; Orange-breasted Waxbill. Bengali ze´bre ´. Fringilla subflava Vieillot, 1819. Nouv. Dict. Hist. Nat., nouv. ´ed., 30, p. 575; Senegal. Range and Status. Resident, sub-Saharan Africa and Tihamah region, Yemen. Absent from arid and forested regions; localized in much of W Africa; more widespread east of 25
E. Mauritania, 1 Rosso (Bull. Afr. Bird Club 7, 2000, 75). Senegal and Gambia, rare; recent records near RichardToll, near Dakar, in Parc Nat. du Delta du Saloum, south of Tambacounda, near Ke´dougou, in Banjul-Cape St Mary area and near Kaur and Sapu; in both countries formerly locally common and has become rare because of habitat destruction by rice cultivation. Guinea-Bissau, frequent Bula area (Rodwell 1996). Guinea, Wassou and Kolente´ areas (Demey 1995), Haut Niger Nat. Park (where frequent: Nikolaus 2000) and Macenta. Sierra Leone, frequent, widespread in E, northwest to Kilimi area. Liberia, uncommon in N Lofa county, not uncommon at Yekepa and Nimba; once flock on Cape Mount Peninsula. Mali, occurrence uncertain; said to be uncommon south of 15
N and common at Kara (Lamarche 1981) but not in definitive list (Lamarche 1993). Burkino Faso, only record a flock at Koubri, Ouagadougou (Thonnerieux et al. 1989).

Ivory Coast, only in NE, from Korhogo to Comoe´ Nat. Park. Ghana, only 4 localities: 2 at Bolgatanga, 20 at Sugu, 40 or more regularly at Denu, and numerous at Atsutsuare (Grimes 1987). Togo, locally not uncommon in flocks of 20–100 in N in Domaine Gravillou-Kandjo-Kara area and in S in Tokpli-Lome´ area (Cheke and Walsh 1996). Benin, not yet found. Nigeria, widespread and locally common, north to Sokoto, Kano, Potiskum and Maiduguri, southwest to Oyo, Iwo and Ibadan, along Niger valley from Jebba to Onitsha, on Jos Plateau (many localities), southeast to Mambilla Plateau (Yelwa); see also remarks on seasonality, below. Cameroon, known from c. 25 localities (M. Languy, pers. comm.). Chad, Central African Republic and Sudan, uncommon and very local, except for Bangui region, Central African Republic, where frequent (6 localities: Germain and Cornet 1994). Ethiopia, locally frequent. Rest of Africa, uncommon to frequent and locally common to abundant, as mapped. Estimated to be 7000 birds in Swaziland and 5000 in Sul do Save, Mozambique, and 10–100 in Lesotho. Apparently less common in Transkei and Lesotho than formerly,

Sporaeginthus subflavus

Sporaeginthus subflavus

? ?

tongue, and blackish crescent at front of floor, between rami of mandibles and partly under tip of tongue; tongue with blackish spots at sides and tip and concealed small black spot underneath (fig. p. 252 – Harrison 1962, Colahan 1982). (Descriptions vary somewhat; mouth pattern may vary individually, and changes with age. See also DEVELOPMENT AND CARE OF YOUNG below). S. s. clarkei (Shelley) (includes ‘niethammeri’): Gabon, S Zaı¨re, and Tanzania (except NW) to South Africa. Less brightly coloured than nominate race. Y has chin to belly golden yellow, often with orange-yellow band across breast; vent and undertailcoverts orange. Larger; wing, Y (n ¼ 10) 41–47 (447). WEIGHT: Y (n ¼ 287) 57–10.3 (73), X (n ¼ 192) 52–109 (76), imm. Y and X (n ¼ 113) 55–93 (70). In Zimbabwe, XX av. 80 in Mar–May and significantly less, 72, in Oct–Dec; in Mar–May, XX significantly heavier than YY. Dusk weights 07 g higher than dawn ones (Colahan 1982).

perhaps because of overgrazing of vleis; has become rare in parts of W Africa (see above). Density, Acacia savanna, Swaziland, 375 birds per 100 ha (Monadjem 2002). Description. S. s. subflavus (Vieillot): Senegal to E Zaı¨re (Kivu), Burundi, W Kenya and NW Tanzania. ADULT Y: forehead to back olive; rump and uppertail-coverts orange-red. Tail blackish brown, tips of T5–T6 fringed whitish. Superciliary stripe red. Lores, narrow ring around eye and mark behind it dark greybrown. Short, narrow buff stripe below eye. Ear-coverts, cheeks and sides of neck greenish olive, fore-cheeks washed orangeyellow. Chin and upper throat yellow or orange-yellow, grading to orange-red on centre of upper breast, bright orange on lower flanks and belly; sides of breast and upper flanks olive-green, barred pale orange; vent and undertail-coverts orange-red. Upperwing feathers greenish olive, tips of tertials finely fringed white. Underwing-coverts and axillaries yellowish buff. Bill red; eyes red; legs light brown or flesh. ADULT X: forehead to back and scapulars dark greyish olive above; rump and uppertailcoverts orange-red or rusty red. Sides of head greyish olive; no red superciliary stripe. Chin to centre of upper breast grey-buff; lower breast and belly orange-buff; sides of breast and flanks greenish olive barred pale buff; vent and undertail-coverts pale orange. Wings and tail as in ad. Y. SIZE (10 YY, 10XX ): wing, Y 42–45 (429), X 40–43 (421); tail, Y 29–35 (315), X 28–30 (296); bill, Y 85–95 (90), X 85–10 (92); tarsus, Y 115–125 (120), X 115–125 (119). WEIGHT: N Nigeria, 1 unsexed 71; Liberia, 1 X 4. IMMATURE: juv. browner than ad. X, less olive; lacks red on rump; buff below with tawny wash on breast, flanks and undertail-coverts; lacks barring on flanks. When newly fledged, bill dull red with culmen dark brown; eyes yellow, feet greyish buff (Chapin 1954). NESTLING: at hatching skin dark pink, crown with dense long white down, back, rump and thighs with shorter, sparse white down (A). Mouth pink, palate with 16 black spots: corner of mouth swollen (less than in other estrildids) and white, inner surfaces with pair of black maxillary and pair of black mandibular spots; palate with ring of 5 black spots, front one with white centre; small black spot on floor of mouth either side of base of

Field Characters. Length 85–10 cm. Tiny and shorttailed, with red bill and eye-stripe, yellow throat and zebra barring on sides; underparts solid orange (W Africa to W Kenya) or yellow with orange breast-patch (Tanzania to South Africa). Rump red, tail black. X lacks red eye-stripe but retains black lores; paler below, barring less distinct, orange confined to undertail-coverts. Immature dull brown above, distinguished from similar immature waxbills by combination of black bill, unbarred pale buff underparts, 2 pale wing-bars and orange-tinged brown rump. Voice. Tape-recorded (88, 99, 104, B, C, F, McVIC, MOY, PAY, STJ). Varied calls include high, thin, tuneless ‘sst’, sseet’ and ‘psit’; lower ‘chyewp’, ‘suet’ or ‘chep’; and ringing ‘sweet’, ‘see-doo’ or abbreviated ‘see-dp’. In flight group gives shrill repeated ‘see’. Principal vocalizations categorized by Colahan (1982) as follows: contact calls, uttered continuously at rate of c. 1–2 per s by birds moving about on ground, soft ‘chit, chit, chit’, or louder, higherpitched ‘cheet, cheet, cheet’ from large group; flight intention call, fairly loud ‘trip, trip, trip’, accelerating before take-off; take-off call, rapid burst of ‘trip’ notes; flight call, soft ‘tink’ at rate of c. 1 per s; alarm, loud,

393

396

ESTRILDIDAE

metallic ‘chink’ at rate of 1–2 per s; louder version of alarm used as threat; attack, harsh ‘cheerk’; and song, usually heard from breeding Y at dawn and dusk, a jumbled series of high-pitched notes interspersed with low ‘cheup’: ‘chip, chit, cheet, chink, cheup, chink, chink, chip, cheet . . . ’, sometimes continuing non-stop for several min. Lowintensity song also given by X and immature at any time of day. Begging call of young, wheezy, hissing ‘chee-cheechee . . . ’. Breeding Y also said to give long series of remarkably loud strident chirps (Goodwin 1982), but unclear if this is a type of song. For sonagrams of various calls see Payne and Payne (1994). General Habits. Inhabits rough montane grasslands, and at low elevations rank grass, tall grass in open savanna woodland interspersed with wet valleys, rank herbaceous vegetation bordering lowland forest, Hyparrhenia grassland, grass at edges of fallow cultivation and large stagnant pools, cassava crops, maize and rice fields, swamps, marshes, vleis, flood-plains and water meadows with grass layer c. 1 m tall, sedges, reedbeds and stands of reedmace Typha latifolia. Sometimes forages in burnt areas. Mainly below 700 m in west and 1600 m in east of range; up to 2000 m in Zimbabwe, in montane grassland, and 2400 m in Ethiopia. In pairs, family parties and flocks of up to c. 20, which coalesce outside breeding season into flocks of >50, occasionally >100. Tame and confiding. Forages only on ground, except for occasionally hawking small flying insects, from grass heads. Takes seeds from soil surface and from inflorescences bird can reach whilst standing on ground; picks small invertebrates from loose soil, sometimes making sideways swipes with bill. Climbs grass stems with ease; can take off from ground and alight on it almost vertically, between tall grass stems. Flight strong but uneven, bird dipping and rising slightly at irregular intervals; flock over open ground or low over reeds flies fast, swerving cohesively and synchronously (Colahan 1982). Continually on the move, small flocks making frequent short flights within or between stands of grass; when disturbed, foraging flock flies to nearest cover in small straggling groups. When moving along perch, flicks tail sideways and slightly upwards; constantly flicks wings. Drinks and bathes regularly at rain puddles, mainly in midafternoon. Drinks whilst standing on ground, or from perch on horizontal or vertical plant stem, bird reaching down to water. Shakes body with feathers ruffled, in conflict situations; commonly yawns and stretches wing and leg on same side; scratches head with foot over wing; preens when resting about midday; bouts of preening often interrupted by allopreening between members of pair or with neighbour; bird invites preening by putting bill between toes and exposing nape; allopreening confined to nape and sides of head; cleans bill by wiping it on perch; occasionally ‘ants’. Commonly dozes by day in shade, hunched, with eyes closed and bill pointing slightly up. In evening goes to roost gregariously in isolated clumps of tall, thick-stemmed grass (in Zimbabwe especially Hyparrhenia cymbaria) and in reedmace and reed beds; favours broken or leaning stems in dense grass or reed growth; sleeping birds huddle together, bills tucked into back feathers.

In southern Africa wanders locally, but essentially sedentary (Harrison et al. 1997). In N Togo, recorded only in Jan–July, ‘so may be intra-African migrant’ (Cheke and Walsh 1996). Around Zaria, Nigeria, common in dry season from start of Oct to mid May, but no records at all in wet season, June–Oct (Fry 1965). Not thought to be migratory in Nigeria, however (Elgood et al. 1973); rather, moves locally, and probably widely overlooked in breeding season, when not flocking; when level of Niger R. at Onitsha falls in Apr–May, fertile alluvium quickly becomes covered with herbage, attracting granivores including many Zebra Waxbills in May–June; species not seen at Onitsha at other times (Serle 1957). Regarded as a migrant in SW Gabon, where locally common in Nov–Feb, a few occurring in Oct and Mar–Apr (Sargeant 1993). Flocks appear on Congo coast (Kouilou Basin) in Nov around pools and grass soaked by rains. In Kenya, local and generally uncommon, but in L. Victoria basin can be common to abundant in rice fields (Zimmerman et al. 1996). Food. Small ripe and unripe grass seeds; some soft shoots and a few insects. Seeds of Setaria sphacelata, Rhynchelytrum repens, Digitaria milanjiana, Panicum novenmere and Hyparrhenia cymbaria (Zimbabwe). Alate termites. Nestlings fed on small seeds, and small insects including beetle larvae, termite workers and ants. Eats av. 226 g of mixed seed daily (in captivity). Breeding Habits. Solitary nester, but can appear to be colonial when several pairs take over adjacent nests in old colony of colonial Ploceus weaver; monogamous. Territorial, Y defending immediate area around nest from incursion by conspecifics. Builds own nest, but more commonly uses old ball nest of weavers and grass-warblers. Of 440 old nests of Red Bishops Euplectes orix examined in Zimbabwe, 318 (72%) were used by Zebra Waxbills; other pairs built 45 of their own nests (Colahan 1982). Also uses insignificant numbers of nests of E. ardens, E. macrourus, E. albonotatus, E. hordeaceus, E. capensis, Ploceus xanthops, P. velatus, Quelea quelea, Paludipasser locustella, Cisticola tinniens, C. erythrops and Prinia subflava. 3 eggs once found in same nest as 2 eggs of E. ardens. 2 courtship displays: (1) Y hops slowly on ground, back and forth next to X, his body upright and bill pointing slightly downwards, tail bent towards her, feathers of flanks and belly fluffed out, rest of plumage sleeked, especially on crown, giving head triangular shape. (2) Y stands next to X, facing same way, and bows deeply, often several times in quick succession. NEST: spherical, barrel-shaped or cylindrical, rather frail, loosely and untidily built (occasionally neatly, looking almost woven) of dry grass stems, leaves and culms; with side or end entrance (B). Built upon foundation of coarser plant material; nest lined with fine grass inflorescences (in Dangila, NW Ethiopia, of dwarf millet) and a few feathers, often of guineafowl; lining grass stems may project from entrance to form a simple hood (Colahan 1982), or entrance can be in form of a tube 7–10 cm long made of grass heads directed outwards (van Someren 1956). Nest quite substantial, but less so when built in dense clump of supporting grass species, e.g. Hemarthria altissima (Zimbabwe). SIZE (n ¼ 11): ext. height 103, ext. width

Sporaeginthus subflavus

96  131, entrance diam. 37, hood length 46. Sited in bush 1 (rarely up to 2) m high (in Zimbabwe often Acacia rehmanniana with grass growing through it), in thickly matted grass or dense grass tussock of e.g. Cymbopogon excavatus; sometimes on ground; nest foundation lodged on clump of grass blades or in forks of one or more branches; height above ground (n ¼ 28 nests) 23–141 (713) cm, nests in bushes being twice as high up as those in grass clumps. Nest built by Y; X may help with lining. One nest consisted of: (platform) 71 grass blades, 3 culms, 40 inflorescences of 7 grass spp., (outer shell) 88 blades, 5 culms, 145 inflorescences, (inner shell) 160 inflorescences, and (lining) 30 feathers and some bits of animal fur and plant down. Structure of ‘adopted’ nest (old nest of another species), which is always a more-or-less spherical nest in good condition, is generally not altered except for addition of 10-mm-thick base (in E. orix nests, which are large) and lining, but if entrance hole has become closed by distortion bird makes new one in side of nest. Lining can be thick, or insubstantial; of fine grasses, and feathers of e.g. dove Streptopelia senegalensis, guineafowl Numida meleagris or domestic fowl, collected from ground or evidently from weaver nests already used by Zebra Waxbills. Nests of weaver Ploceus velatus, with bottom entrance, have second entrance made by waxbill in top. Added base in one E. orix nest consisted of 32 blades, 4 culms and 35 inflorescences, and lining consisted of 138 inflorescences and 56 feathers (Colahan 1982). Pair searches for weaver nest whilst flying over or moving through vegetation. When new nest found, pair alights a few m away, Y approaches cautiously, flies to it,

clings to entrance and peers cautiously inside. If nest is empty, X also inspects it, outside and inside. Addition of lining takes 2–5 days. EGGS: 3–9, usually 4–6, av. (n ¼ 110) 50. Eggs laid at night or early morning on successive days. At first pinkish white, smooth, not glossy; as incubation proceeds become off-white and glossy. SIZE: (n ¼ 211 eggs from 51 clutches, Zimbabwe) 118–153  93–110 (136  102). WEIGHT: (n ¼ 25) av. 077 g. LAYING DATES: Senegal, Richard-Toll, Jan–Feb; Nigeria, Nov (and accompanied young, Jan); Central African Republic, (active gonads, Nov); Sudan, June–Sept; Ethiopia, Dec; NE Zaı¨re, (oviduct egg, Oct; fledgling Jan); E Africa: Region B, Oct–Jan, Mar, June–July, Region C, May–June, Region D, Feb, Apr–June, Aug–Sept, Dec; Angola, Apr (and breeding condition May); SE Zaı¨re, Apr– ˆ i, Mar– May; Zambia, Mar–May (mainly Apr–May); Malaw Aug; Zimbabwe, Dec–July, mainly Feb (81 clutches), Mar (286), Apr (115) and May (35); Botswana (carrying food, May); Transvaal, Jan–July (mainly Feb–Apr), KwaZuluNatal, Jan–May. INCUBATION: starts with 2nd or 3rd egg. By X and Y equally by day; by X at night. Clutch covered for >99% of time. Y adds feathers and grass inflorescences to nest during incubation period (and first few days of nestling period). Period: 13–14 days. Clutches of 5 and 6 eggs take 33–38 h to hatch. DEVELOPMENT AND CARE OF YOUNG: parents remove eggshells and some faecal sacs, but nest soon becomes fouled. Young fed by X and Y, by regurgitation. Growth data tabulated by Colahan (1982). Remex quills break through on days 5–6, rectrices on days 6–7; feathers emerge from days 9–10 (remiges, rectrices), 11 (ventral tract) and 13–14 (head); eyes open from day 6 to day 9. Mouth of fledgling and juv.: palate pinkish white with 5 black spots, front one with pale centre, front of palate with 2 large black chevrons; corner of mouth swollen, pinkish or greyish and black, maxilla with 2 opalescent whitish or bluish papillae, mandible with 1 blue papilla; floor of mouth with row of 4 black spots (Chapin 1954, Harrison 1962, Colahan 1982, Payne and Payne 1994, R.B. Payne, pers. comm.). Period: (n ¼ 7 broods) 17–19 days. Young leave nest by scrabbling out together, fluttering off in different directions to find cover. Young out of nest dependent for 2 weeks; they return to roost in nest at night, with parents, for up to 16 days. BREEDING SUCCESS/SURVIVAL: in N tropics the only host of brood parasitic Jambandu Indigobird Vidua raricola. 62% of eggs hatch; 30% of nestlings fledge; so overall success rate is 18% (Colahan 1982).

Key References L.L. (1994).

Colahan, B.D. (1982), Payne, R.B. and Payne,

397

398

ESTRILDIDAE

Genus Amandava Blyth

Single Oriental species, with red bill, black tail and black wings with white spots, Y with white-spotted red body and black undertail-coverts, X with fawn body and pale undertail-coverts. Affinities controversial; molecular evidence of quite close relationship with Sporaeginthus, q.v. Superficially resembles white-spotted pink African waxbills Lagonosticta spp., particularly L. senegala, but amandava feathers have 1 white spot, on shaft, Lagonosticta ones paired white spots, on vanes. Old captive Amandava whose reds have become browns, can bear strong resemblance to Lonchura (Spermestes) spp., especially Oriental L. leucosticta (Harrison 1962). Able to rotate foot outwards and grasp one or more vertical stems on either side of body. Singing posture (body upright, plumage sleeked, bill horizontal) like Lonchura and unlike Estrilda, Lagonosticta and Uraeginthus; song behaviour and aspects of sexual display also like Lonchura (Harrison 1962).

Plate 19

Amandava amandava (Linnaeus). Red Avadavat. Bengali rouge.

(Opp. p. 282)

Fringilla amandava Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 180; eastern India. Range and Status. Central Pakistan to SE China and Vietnam, south to Java, Bali, Lombok, Flores, Sumba and Timor. Introduced to or feral in Singapore, Japan, Philippines, Hawaii, Fiji, Puerto Rico, N Italy, Spain (4 areas), Saudi Arabia (Riyadh), Israel (Eilat), N Egypt and Re´union; escapes in Iran, Bahrein, Oman, Lebanon, Israel and elsewhere. Introduced in Nile Delta, Egypt, perhaps near Alexandria, before 1861. Old records at Giza (where formerly large flocks in Zoological Gardens), Inchas, Bilbeis, Luxor, Faiyum, Abu Rauwash and El Mansura. In 1970s, frequent near Ismailiya and Asher Talaaf; presently widespread and locally common in W, E and S parts of Delta and near Suez, where c. 100 since 1980; common along Nile just south of Cairo; frequent on borders of L. Qarun; recent records at El Saff, between Mazghuna and El Wasta, at Beni Suef (Goodman and Meininger 1989) and on Nile in S Egypt, between Nag Hammadi and Kom Ombo; 60 on Crocodile I., Luxor, in 1997 (Bull. Afr. Bird Club 1998, 71, and 2001, 149). Description. A. a. amandava (Linnaeus): only race in Africa. ADULT Y (breeding): forehead to hindneck, sides of neck, mantle and back olive-brown with crimson-red feather tips, largely masking brown bases on forehead; scapulars olive-brown with broad crimson-red tips, each with small white subterminal spot. Rump feathers and uppertail-coverts brighter scarlet-red with blackish bases, each with small subterminal white or pinkish white spot. Tail black, upperside with slight violet gloss; T5 and T6 with white spot at tip of outer web (small on T5). Lores, narrow ring around eye and triangular mark behind eye black, bordered below eye by fine whitish stripe; supercilium, cheeks and ear-coverts crimson-red with hidden brown feather bases. Chin to upper breast and flanks bright crimson-red with blackish brown feather bases; breast and flank feathers each with white spot near tip, small on breast, large and circular on flanks; redtinged sooty brown lower breast merges into sooty black belly, thighs and undertail-coverts. Upperwing feathers dark olivebrown, with white spots on tips of tertials and greater, median and lesser coverts. Underwing-coverts and axillaries pale buff. Bill red, base or whole of culmen ridge dusky; eyes orange-red or red; legs flesh brown. ADULT Y (non-breeding): like ad. X, but

Amandava amandava

rump and scapulars with white spots; sometimes a few scattered red feathers below. ADULT X: forehead to back olive-brown; rump feathers and uppertail-coverts crimson with brown bases. Tail blackish brown, upperside slightly glossy. Lores, narrow ring around eye and mark behind eye black, bordered below by white stripe. Cheeks and ear-coverts greyish brown, merging with olivebrown sides of neck and with buffy white chin; throat and upper breast to flanks light olive-grey; rest of underparts rich ochreous buff. Upperwing feathers dark olive-brown, with small white spots at tips of tertials, greater and median coverts; underwingcoverts and axillaries pale ochreous buff. SIZE (15 YY, 12 XX): wing, Y 47–51 (490), X 46–50 (476); tail, Y 35–39 (370), X 33– 39 (363); bill, Y 105–120 (110), X 98–115 (107); tarsus, Y 132–146 (140), X 133–140 (140). IMMATURE: juv. fawn-brown above including rump and uppertail-coverts, buffish white below; lacks white spots and black eye

398

ESTRILDIDAE

Genus Amandava Blyth

Single Oriental species, with red bill, black tail and black wings with white spots, Y with white-spotted red body and black undertail-coverts, X with fawn body and pale undertail-coverts. Affinities controversial; molecular evidence of quite close relationship with Sporaeginthus, q.v. Superficially resembles white-spotted pink African waxbills Lagonosticta spp., particularly L. senegala, but amandava feathers have 1 white spot, on shaft, Lagonosticta ones paired white spots, on vanes. Old captive Amandava whose reds have become browns, can bear strong resemblance to Lonchura (Spermestes) spp., especially Oriental L. leucosticta (Harrison 1962). Able to rotate foot outwards and grasp one or more vertical stems on either side of body. Singing posture (body upright, plumage sleeked, bill horizontal) like Lonchura and unlike Estrilda, Lagonosticta and Uraeginthus; song behaviour and aspects of sexual display also like Lonchura (Harrison 1962).

Plate 19

Amandava amandava (Linnaeus). Red Avadavat. Bengali rouge.

(Opp. p. 282)

Fringilla amandava Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 180; eastern India. Range and Status. Central Pakistan to SE China and Vietnam, south to Java, Bali, Lombok, Flores, Sumba and Timor. Introduced to or feral in Singapore, Japan, Philippines, Hawaii, Fiji, Puerto Rico, N Italy, Spain (4 areas), Saudi Arabia (Riyadh), Israel (Eilat), N Egypt and Re´union; escapes in Iran, Bahrein, Oman, Lebanon, Israel and elsewhere. Introduced in Nile Delta, Egypt, perhaps near Alexandria, before 1861. Old records at Giza (where formerly large flocks in Zoological Gardens), Inchas, Bilbeis, Luxor, Faiyum, Abu Rauwash and El Mansura. In 1970s, frequent near Ismailiya and Asher Talaaf; presently widespread and locally common in W, E and S parts of Delta and near Suez, where c. 100 since 1980; common along Nile just south of Cairo; frequent on borders of L. Qarun; recent records at El Saff, between Mazghuna and El Wasta, at Beni Suef (Goodman and Meininger 1989) and on Nile in S Egypt, between Nag Hammadi and Kom Ombo; 60 on Crocodile I., Luxor, in 1997 (Bull. Afr. Bird Club 1998, 71, and 2001, 149). Description. A. a. amandava (Linnaeus): only race in Africa. ADULT Y (breeding): forehead to hindneck, sides of neck, mantle and back olive-brown with crimson-red feather tips, largely masking brown bases on forehead; scapulars olive-brown with broad crimson-red tips, each with small white subterminal spot. Rump feathers and uppertail-coverts brighter scarlet-red with blackish bases, each with small subterminal white or pinkish white spot. Tail black, upperside with slight violet gloss; T5 and T6 with white spot at tip of outer web (small on T5). Lores, narrow ring around eye and triangular mark behind eye black, bordered below eye by fine whitish stripe; supercilium, cheeks and ear-coverts crimson-red with hidden brown feather bases. Chin to upper breast and flanks bright crimson-red with blackish brown feather bases; breast and flank feathers each with white spot near tip, small on breast, large and circular on flanks; redtinged sooty brown lower breast merges into sooty black belly, thighs and undertail-coverts. Upperwing feathers dark olivebrown, with white spots on tips of tertials and greater, median and lesser coverts. Underwing-coverts and axillaries pale buff. Bill red, base or whole of culmen ridge dusky; eyes orange-red or red; legs flesh brown. ADULT Y (non-breeding): like ad. X, but

Amandava amandava

rump and scapulars with white spots; sometimes a few scattered red feathers below. ADULT X: forehead to back olive-brown; rump feathers and uppertail-coverts crimson with brown bases. Tail blackish brown, upperside slightly glossy. Lores, narrow ring around eye and mark behind eye black, bordered below by white stripe. Cheeks and ear-coverts greyish brown, merging with olivebrown sides of neck and with buffy white chin; throat and upper breast to flanks light olive-grey; rest of underparts rich ochreous buff. Upperwing feathers dark olive-brown, with small white spots at tips of tertials, greater and median coverts; underwingcoverts and axillaries pale ochreous buff. SIZE (15 YY, 12 XX): wing, Y 47–51 (490), X 46–50 (476); tail, Y 35–39 (370), X 33– 39 (363); bill, Y 105–120 (110), X 98–115 (107); tarsus, Y 132–146 (140), X 133–140 (140). IMMATURE: juv. fawn-brown above including rump and uppertail-coverts, buffish white below; lacks white spots and black eye

Amandava amandava patch. Tertials fringed buff; greater and median coverts broadly tipped buff to give 2 prominent wingbars. Bill dusky, lower mandible with pink base; eyes dull brown. NESTLING: dark-skinned, crown and back with loose brownish down; mouth markings like those of Zebra Waxbill Sporaeginthus subflavus nestling (q.v.), but paired not single black mark at front of palate, and 3 not 2 black mandibular marks near corner of mouth.

Field Characters. Length 10 cm. Confined to N Egypt, where breeding Y unmistakable (no firefinches in Egypt), bright crimson with white spots, dark wings and tail. X and non-breeding Y brown with red bill and rump, white spots on dark wing. Immature nondescript, like, small, neat X House Sparrow Passer domesticus, but wing darker than rest of body, with 2 narrow white bars and white tips to tertials. Voice. Tape-recorded (B). Short, sweet song, variable and descending. Alternative song, a rapid descending trill (Goodwin 1982). Call notes high and shrill but with sweet tone, louder in aggression or alarm. Nest call, rapid ‘tehteh-teh-teh’ (Goodwin 1982). General Habits. Inhabits emergent vegetation, mainly Phragmites reed beds, also reedmace Typha, rushes, sedges, rank grass, tamarisks, overgrown ditches; irrigated crops and gardens; generally over or near standing water. Occurs in pairs and small flocks of 5–10 and up to c. 30; formerly large flocks near Cairo. Forms small wandering flocks from late Nov, which congregrate for roosting. Tame and confiding. Forages on bare ground and fallen grassheads, amongst herbaceous vegetation, and particularly in reeds along ditches and in tall waterside grasses; also in farmland. Hops on ground; agile in reeds, holding 1 or more vertical shoots in each foot with legs splayed out; makes short flights through reeds to clump a few m away, clings to stems and leaves, and works its way rapidly up to seeding head to feed there. Picks insects from vegetation and takes termites in short flights. Rests often during day, birds clumping in thick vegetation, 2–4 or more perching in row, flank to flank; clumping often happens after bathing and preening; self-preening and allopreening common; bird being preened exposes neck by putting bill between feet or tilting head back. When excited, swings tail from side to side in U-shaped path. When disturbed, foraging flock rises in tight pack and settles in nearby trees or in distant reeds. Flight swift and direct; often calls in flight. Roosts communally, in reeds and reedmace beds, birds settling noisily then sleeping flank to flank. Sedentary and locally nomadic in Egypt.

in all directions from nest; he sings from favoured perch, often top of reed; sometimes sings in flight; often sings on ground, next to crouched X, turning his head from side to side. In pair-bonding display, perched bird picks up thin seed-head, stands with ruffled plumage in front of mate, bows slowly to one side, raises head, then bows to other side; at lowest point of bow, turns head stiffly to one side, bill pointing down. NEST: quite strong, slightly flattened ball of coarse and fine grass stems, flowering heads and leaf strips up to 80 cm long, with central side entrance, often resting on platform, av. 227 long, made of grass and twigs; lined with fine grass flower-heads, plant down and feathers. Not elaborately woven; sometimes spider web bound into outer surface of nest; entrance may be somewhat tubular; one entrance diam. 38. SIZE: ext. av. width 146  126, height 109. Sited 500,000.

Spermestes cucullatus

?

?

?

species in the continent. Occurs in Grande Comore, Moheli, Anjouan and Mayotte (Comoro Is). Introduced to Puerto Rico. Occurs up to 700 m in Liberia, 2000 m in ˆ i (Mt Zomba), nearly 2200 m in Sudan, E Africa and Malaw E Zaı¨re (but only 1900 m in Itombwe) and 1200 (occasionally 1500) m in Zimbabwe. Ranges north to 15
N in W Senegal (Thie`s District),
14 N in Mali (Boucle du Baoule´ and Sansanding north of Se´go), Comoe´ Nat. Park in Ivory Coast and Ouagadougou in Burkina Faso. Further east, markedly less common north than south of 10
N; just reaches Niger in ‘W’ Nat. Park; north in Nigeria to Kano; in NE Cameroon/W Chad common in Chari drainage north to 12
N (Maga, Logone Birni), in E Chad common north to Bahr Sara and Bahr Azoum; in Sudan nominate cucullatus west of Nile to 8
300 or 9
N, in Ethiopia (and along E Sudan border) scutatus in lowlands and highlands north to central Gondar Prov. and east nearly to Somali border. In Kenya, cucullatus west of Rift Valley, north to 2
N (common) and 4
N (uncommon, local), scutatus east to Mt Kenya, Thika and Machakos, with a coastal plains population north to Lamu and Bura. Avoids dense forest, as in large parts of Liberia, Gabon, Congo and Zaı¨re, as shown; avoids arid areas, e.g. central and W Tanzania. Almost certainly widespread in E Central African Republic, Moxico Prov. (Angola) and N Mozambique (question marks in map; in Moxico Prov., known only from Calunda, L. Calundo, L. Cameia and Luacano in NE – Dean 2000). Locally common on islands of Bioko, Prı´ncipe, Sa˜o Tome´, Pemba, Zanzibar and Mafia; a recent arrival on Annobn, where breeding (del Val 2001). Gabon, locally frequent to abundant, probably more widespread than shown. Angola, cucullatus in Cabinda, Zaı¨re, Uı´ge, N Cuanza Norte, scutatus from Bengo, S Cuanza Norte, Malanje, Cuanza Sul and Bie´ to coastal Benguela and Namibe, W and N Huı´la and NE Moxico

Description. S. c. cucullatus (Swainson): Senegal to Sudan, south to L. Victoria basin and N Cuanza Norte, Angola; Bioko, Prı´ncipe and Sa˜o Tome´. ADULT Y: Head to chin, throat and centre of upper breast brownish black with dark green or bronzy gloss (greenish on top of head, more bronzy or violet on cheeks and breast). Hindneck, sides of neck, mantle and back greybrown; rump and uppertail-coverts barred greyish white and blackish; scapulars dark glossy green. Tail black. Sides of upper breast greyish white, barred blackish brown; patch on side of lower breast black, glossed dark green; centre of lower breast and belly white; flanks, thighs and undertail-coverts barred blackish and greyish white. Flight feathers blackish brown, primaries narrowly edged whitish brown, bases of outer webs barred whitish, secondaries edged grey-brown; rest of upperwing feathers dull dark brown, lesser coverts with some glossy green tips. Underwing-coverts and axillaries white. Upper mandible black, lower pale blue-grey; eyes dark brown; legs dark grey, greenish grey or blackish. Sexes alike. Intergrades with scutatus. SIZE (10 YY, 10 XX): wing, Y 48–50 (489), X 46–51 (480); tail, Y 28–31 (294), X 27–30 (289); bill, Y 11–12 (112), X 105–12 (110); tarsus, Y 12–13 (123), X 12–13 (124). WEIGHT: N Sierra Leone (n ¼ 552), weights decline in dry season to min. of c. 83 g (XX) in Apr–May and 87 g (YY) in Apr–Aug, and are max. of c. 106 (XX) and 101 (YY) in Sept (Thompson and Tye 1995); SW Nigeria (n ¼ 102), av. weight varies seasonally from 8–85 g in Aug to 12–13 g in Apr–May (Akinpelu 1994a); N Nigeria, unsexed (n ¼ 52, Oct–June) 79–96 (85), N Ghana, unsexed (n ¼ 54) 70–105 (93), W Kenya, unsexed (n ¼ 97) 77–118 (92). IMMATURE: juv. has head to throat and centre of upper breast dark grey-brown, tinged tawny; mantle to scapulars and uppertail-coverts grey-brown; sides of breast and flanks deep tawnybuff, becoming paler on belly and undertail-coverts. Wings and tail as ad., but tertials narrowly fringed buff and tail tinged brown. Bill black. NESTLING: at 2 days skin pink, darker areas on crown and back, wing with black line along it; short, fine, white down on supraorbital, occipital, humeral, femoral and spinal tracts. Palate and tongue yellowish, palate with 2 concentric black crescents, floor of mouth with 2 less distinct ones; tongue encircled by prominent black ring (fig. p. 252); tip of bill black, base off-white; gape whitish, not very swollen and without papillae. Tarsus white, claws with black tips; gape white; eyes closed, eyelids black. S. c. scutatus (Heuglin) (includes ‘tessellatus’): Ethiopia and adjacent E Sudan, Kenya (Rift Valley and to east), Tanzania, S Zaı¨re (S Kasai and Katanga) and Angola (except NW) to Botswana and South Africa. Like nominate race but lacks greenglossed patch on side of breast. WEIGHT: central Kenya, unsexed (n ¼ 25) 92–11 (100); Kenya coast, unsexed (n ¼ 6) 74–85 (81); Zimbabwe, Y (n ¼ 4) 82–96 (90), X (n ¼ ?) 81–95 (87), unsexed ad. (n ¼ 106) 84–122 (99), imm. (n ¼ 50) 84–106 (95), ad. weight varies daily from c. 95 at 09h00 to c. 103 at 13h00, and seasonally from av. 98 in Apr–May to av. 101 in Sept ˆ i, unsexed (n ¼ 49) 70–103 (86); (Woodall 1975); Malaw Mozambique, unsexed (n ¼ 41) 70–110 (91).

Field Characters. Length 85–10 cm. Small flocks of this tiny, plump, short-tailed bird are a common and familiar sight throughout much of Africa, feeding on the ground and making short flights on whirring wings when flushed. Dark head, grey-brown upperparts, pale rump barred

Spermestes cucullatus brown, black tail; underparts white, with dark bars on flanks (coalescing in nominate race to form smudgy bottlegreen patch at sides of breast); bottle green patch on shoulder visible in good light. Bill stout, 2-tone, blue-grey with dark upper mandible. Black on head more restricted than in other mannikins, black crown and face grading into warm brown nape and hindneck; dark bib is rounded (white of lower breast continues onto sides of neck). Immature uniform buffy brown with pale belly; told from other mannikins by barred undertail-coverts. Voice. Tape-recorded (9, 86, 88, 99, 104, B, C, F). Calls of 2 main types: high-pitched and shrill, buzzy ‘dzeep’, ‘bzheet’ or ‘peejee’, or rolled ‘drrreep’; or sparrow-like chirping, ‘cheew’, ‘chrup’, ‘cha-cha’, ‘chee-jata’, ‘chukka’. In flight, ‘tsrii, tsrii, tsri’. Continuous dry chatter given by group on branch. Alarm, ‘tsek’, ‘cheep’ or ‘chuck’; fighting call a prolonged ‘chhiiii . . . ’. Roosting and nest-ceremony call, series of very short phrases of 2–8 syllables, rising then falling, ‘chi chi chi chi chu chuu’ (Woodall 1975). Song rather deep, slow and measured, in 2 parts, ‘chuckalajuvajuvajuvajuvajuva . . . ’. Quiet courtship song of 8 syllables, ‘chi, chu, chi, chu, cheri-hit, chu’, lasting 1–2 s, repeated several times in 1–2 min (Woodall 1975). General Habits. Inhabits long grass, rank vegetation along streams and rivers in woodlands, thornveldt, roadsides, forest clearings, grassland, farmland especially with cereal cultivation, edges of villages, fallow ground, suburban and rural gardens and parks. Availability of water a more important determinant than vegetal structure (Woodall 1975). Pairs off for nesting, otherwise gregarious, year-round; in flocks usually of up to c. 15, often 20–50. Birds in flock forage and fly close together in orderly, structured manner; they take off and alight synchronously and abruptly; flight generally short and whirring. Brisk, hopping locomotion on ground, where often frantic short-lived scuffles between neighbours. Tame. Often mixes with other estrildids, particularly mannikins and waxbills, when feeding; moves forward by hopping and making flights of c. 1 m. One or more birds alight on thin Panicum maximum stem causing it to bend, when they edge along sideways to seeding head and nibble and husk seeds; eats seeds of Setaria mainly on ground but sometimes at heads that have become matted together. Forages on bare ground near bushy or grassy cover, on short-grass sward (playing fields, airstrips), and amongst grasses and herbs c. 1 m tall. Bird pulls stalk towards it with bill, then holds it underfoot and pecks at seed head. On dry, pulverized earth, feeds by hopping and searching; when with indigobirds Vidua which are feeding by ‘jump-kicking’, often makes a dash at one which retreats, leaving mannikin to search powdery soil for minute seeds of e.g. grass Eleusine coracina where the indigobird has just kicked; this appears to be a standard commensal feeding practice by Bronze Mannikins (Fry 1975). Commonly ‘flycatches’ for termites, flying out short distance from small tree, catching insect in air and returning to perch with it; if bird fails to catch insect at first attempt it hovers briefly and catches it at second or third attempt; returning to perch, bird moves termite

around in bill using tongue to remove wings. Perches on stalk behind flower in Eucalyptus tree, leans forward, inserts bill into middle of inflorescence, apparently taking nectar (Woodall 1975); hand-reared nestlings sucked liquid feed from small paintbrush (Jewitt 1989). When alarmed, bird calls, cranes neck and flicks wings and tail. In aggression or defence, raises one wing; nestling soliciting food does the same (Morris 1957). Flock of c. 6 birds commonly rests in shrub during day, tightly huddled together on branch; a late arrival forces its way into middle of row. They may spread tails, put heads together, and give continuous dry chatter whilst bobbing (Haagner 1964). Flock has strong individual dominance or ‘pecking’ order; when small flocks (family parties?) combine, it leads to much scuffling and fighting, with feathers flying and sometimes a bird almost killed (Woodall 1975). Small flocks of XX and juvs described as chasing lone Y around garden, displacing him from perch, with frantic activity at dusk when going to roost, birds fighting and piling on top of each other (Reid 2000); appears to be unusually robust, physically and socially, even when very young (Jewitt 1989). Bathes in water once or twice daily, then preens in set pattern: shakes feathers, wipes bill on branch, preens, shakes feathers, then preens several times using preen gland. Sometimes allopreens: 1 bird in huddle on branch solicits by crouching, then neighbour preens its head and throat (and, uncommonly, breast, belly, rump and tail); bird being preened bends its head back or sideways and closes eyes; allopreening then reciprocated. Outside breeding season, roosts gregariously in own nests; flock gathers near nest, one bird enters, shuffles around for a few min., then calls remainder in; uses same roosting nest for months on end (Woodall 1975). On cold nights several birds huddle in nest to keep warm, which benefits dominant individuals particularly (Calf et al. 1998). Roosts either in old breeding nest, which can be relined and maintained for roosting throughout non-breeding season, or in purpose-built roosting nest; these are constructed communally (breeding nests built by pair), and are thinner and much less robust than breeding nests; 3 were made of 210, 360 and 400 pieces of grass; they are sometimes pulled apart and the material used for new nest. Moult follows usual passerine pattern but is very protracted. In Sierra Leone, mainly in dry season, from Sept to Apr or June (ads) or Nov to about June (juvs); primary moult of juvs may be incomplete (Thompson and Tye 1995). In Nigeria, moults in May–Oct, between major and minor breeding seasons; duration estimated at 183 days (Akinpelu 1994a). In Zimbabwe, from Mar to Sept (ads) or June to Oct (juvs); av. replacement times for single primary feather, ad. (n ¼ 12) 19 days, juv. (n ¼ 11) 165 days (Woodall 1975). Resident, mainly sedentary, but may wander locally (at least 9 km, Zimbabwe); numerous at Koubri, Burkina Faso, in Apr but absent in other months (Thonne´rieux et al. 1989); seasonal up to 2000 m in S Sudan. Food. Mainly grass seeds; in Zimbabwe, of Eleusine indica, Sporobolus pyramidalis, Panicum maximum, Setaria homonyma, S. anceps, S. pallidefusca, Echinocholoa pyramidalis, Brachiaria

403

404

ESTRILDIDAE

nigropedata, Paspalum urvillei, Digitaria gazensis, Rhynchelytrum repens, R. setifolium and Hyperthelia dissoluta. Nectar (see above); small bits of lettuce leaves; and filamentous alga Spirogyra (Zimbabwe: Woodall 1975). In Sierra Leone, seeds of additional grasses Panicum laxum, Sorghum, Anadelphia, Andropogon, Pennisetum, Eragrostis, Axonopus and Paspalum spp. In Gabon, mainly Paspalum. Spirogyra forms 5% of dietary items in Sierra Leone, where other components (n ¼ 305 items) are rice Oryza sativa grains 30%, Sorghum seeds 14%, other grass seeds 48%, Eugenia uniflora (Myrtaceae) fruit mesocarp 1% (Thompson 1989). Eats many termites (e.g. Macrotermes bellicosus) and a few other insects (hunting wasps, weevils, flies, ants including Oecophylla longinoda); takes only one-third as many insects as eaten by S. bicolor, and does not give insects to young nestlings. Ad. eats 21–24 g of food per day. Breeding Habits. Solitary or semi-colonial nester; monogamous; territorial, territory extending c. 1 m around nest. Solitary nests sometimes as little as 1 m apart in adjacent trees, forming colony of up to 6 nests. 3rd bird often tries to enter pair’s nest, but is vigorously driven off. Fighting rivals may occasionally fall to ground. Aggression an integral part of pre-courtship behaviour, breaking flock up into pairs (Morris 1957). Courting Y sings, perched at angle of c. 45
, with head and tail turned towards X, bill wide open and pointing down, tongue protruding, and crown, back and underpart feathers raised; he bobs up and down on bent legs, and may also rapidly open and close bill and quiver tongue. Occasionally Y courts whilst carrying grass stem in bill, but soon drops it. X, in similar oblique posture, quivers tail. Y starts to build nest; X soon takes over, twisting and turning stems into place, Y continuing to bring material. Bird collecting material stands on grass tuft, selects a stem, and jerks it up or to side until it comes away. Bird’s flight whilst carrying stem is noisy. When building nest, Y, carrying grass stalk, perches adjacently for a few s, X in nest makes scooping movements and repeatedly utters nest-ceremony calls which become more insistent when Y enters nest; calls are followed by rapid opening and closing of bill or by open-mouth display with tongue quivering (Morris 1957, Woodall 1975). Later, X in nest gives nest-ceremony call, Y enters and they copulate. If Y not ready to copulate, he vaults back and forth over perched X, with quick displacement preen between each jump (Morris 1957). Commonly 2–3 broods per season and sometimes 4, in same nest. NEST: bulky, untidy ball with side entrance; made of green grass stems, leaves and flowering heads, the entrance partially covered by feathery grass heads projecting over it or forming slight tunnel. In Kenya almost always made of grass Rhynchelytrum repens; in Zimbabwe uses grasses including Thalictrum sp. and most of the grass species whose seeds are eaten (see Food), also needles of Pinus patula and stems of wild asparagus. Often distinct outer layer of long grasses Hyparrhenia dichroa and Hyperthelia dissoluta (Zimbabwe). 4 nests were made of 540, 610, 640 and 800 pieces of grass (twice as many as in roosting nests: Woodall 1975). SIZE (n ¼ 4): height 140–180 (163), width 110–130 (125), depth 130–170 (155), entrance height 38–40 (39), entrance width 40–45 (42). Sited in great variety of

situations, mainly in vegetation, from 06 to 12 m above ground (av. of 8 nests, 55 m); placed amongst fine twigs or thick foliage or cluster of thorns, towards end of branch. Often uses Eucalyptus trees (W and E Africa) and fir trees (southern Africa), also man-made structures, e.g. security ˆ i); in Zaı¨re, nests between lights at Chileka Airport (Malaw palm frond bases, in forks of Acacia, Citrus and Mangifera trees, in eaves of grass-thatched houses, holes in walls, and within creepers or bunches of plantain fruits. In Liberia, less dependent on thatched roofs than formerly (Gatter 1997). Often sites nest near stinging hymenopterans’ nest, e.g. wasps Belonogaster griseus or Ropalidia sp. (Kenya, Zimbabwe), or vicious tree ants Oecophylla smaragdina (Nigeria) and O. longinoda (South Africa); once only 40 cm from polistine wasp nest (Togo), once ‘only inches’ away. Occasionally uses old weaver or estrildine nests (e.g. Ploceus xanthopterus, P. velatus, P. cucullatus, P. nigricollis, Anaplectes rubriceps, Euplectes orix, Uraeginthus angolensis, Estrilda astrild), and once old nest of swift Apus affinis (Togo). ˆ i, where av. 475), or 3–4 EGGS: 2–8, mainly 5 (Malaw (Gabon) or 4–5 (Kenya). Laid daily. Pure white; long oval or blunt ovoid, shape varying greatly. SIZE: (n ¼ 28, Zimbabwe) 131–161  101–108 (148  104). WEIGHT: (n ¼ 28) 06–09 (08) g. LAYING DATES: Senegal, Sept–Oct; Gambia, Apr, Sept– Dec; Liberia, Aug–Oct (–Nov) (and imms all months, most often Oct–Apr); Burkina Faso, (nest-building July–Aug); Ghana, (nesting, Mar–July; dependent young May, June, Aug, Oct–Jan); Togo, Aug, Oct, (and building Apr–Sept; hatchlings July; young in nest Aug and Oct); Nigeria, Apr–Dec, mainly June–Sept); Sudan, July, Sept; Ethiopia, Aug–Oct; E Africa, Region A, June, Sept, Nov, Region B, Feb–Dec, mainly Apr–June, Region C, Dec, Jan, Mar, May, June, Region D, all months, mainly Apr–June; Gabon, Oct–Feb, Apr, June–Aug; Angola, Jan, Apr; Zambia, Oct– ˆ i, Dec–May, mainly Feb; May, mainly Jan (–Mar); Malaw Zimbabwe, Aug–May, mainly Dec–Mar (80% of 154 clutches); Botswana, Sept–Mar; South Africa, Aug–Apr. INCUBATION: by day mainly by X, at night by both Y and X. Daytime incubation starts with last or penultimate egg laid. Period: 14–16 days, usually 14 (Woodall 1975), or 12 days (Restall 1996). Clutch takes 1–2 days to hatch. DEVELOPMENT AND CARE OF YOUNG: when begging, nestling moves pied tongue from side to side and up and down. Skin darkens and wing quills just protruding by 7 days; at 11 days, quills long and prominent, feathers beginning to protrude from them, and eyes 75% open; 14 days, half feathered, but quills still prominent; 30 days, bill uniformly dark, white gape swelling now small, head uniformly brown (Woodall 1975). From 20 to 75 days after young fledges, mandible changes from black to grey; from 20 to 35 days, gape changes from black to yellow; in 5th–8th week head starts to change from brown to glossy black, taking 80–100 days; bird fully ad. after 130 days (Scogins 1988). Parents feed young by regurgitating partially softened seed, apportioning small quantity into mouth of each chick; several rounds may be made before parent’s crop is emptied. Feeding visits by both parents occur at regular intervals of c. 25 min in morning, and from mid-afternoon until dusk. Parent remains at nest for long time but does not remove faeces, so nest rapidly becomes fouled;

Spermestes bicolor however, most nestling faeces fall through loose structure of nest when dried out (van Someren 1956). Period: 15–17 days (Kenya); in Zimbabwe, 17±1 days; young birds may actually leave nest at 14 days, 2–3 days before they can fly. If disturbed prematurely in nest, they scatter in all directions, up to 50 m away; parents entice each one back by calling to move it gradually up to top of tree, parent flies from treetop back to nest, nestling flutters down after it and may have to be led up to another treetop before finally returning to nest (Woodall 1975). Out of nest, young keep with ads; they nibble at twigs, leaves and grass stalks, and are fed regularly by ads at c. 30 min intervals, clustering around parent with mouths wide open, sometimes raising one wing; young occasionally led back to nest and fed inside. BREEDING SUCCESS/SURVIVAL: eggs taken by climbing mice Dendromus sp. (Kenya) and snake Thrasops flavigularis

405

(Gabon), and nestlings and fledglings by African Goshawk Accipiter tachiro, Wahlberg’s Eagle Hieraaetus wahlbergi, coucals Centropus spp., Woodland Kingfisher Halcyon senegalensis, drongos Dicrurus, Pied Crow Corvus albus; ads taken by domestic cats, Fiscal Shrike Lanius collaris, Redthighed Sparrowhawk A. erythropus and Little Sparrowhawk A. minullus. Occasionally parasitized by Pin-tailed Whydah Vidua macroura. Max. longevity (recapture data) 4 years; 35% of ads at start of breeding season survive to following year; av. ad. life expectancy, 125 months (Woodall 1975).

Key References Akinpelu, A.I. (1994a, b), Brosset, A. and Erard, C. (1986), Clement, P. et al. (1993), Goodwin, D. (1982), Morris, D. (1957), Restall, R. (1996), van Someren, V.G.L. (1956), Thompson, H.S. (1989), Thompson, H.S. and Tye, A. (1995), Wolters and Clancey (1974), Woodall, P.F. (1975).

Spermestes bicolor (Fraser). Black-and-white Mannikin; Red-backed Mannikin. Capucin bicolore.

Plate 24

Amadina bicolor Fraser, 1843. Proc. Zool. Soc. London, 1842, p. 145; Cape Palmas, Liberia.

(Opp. p. 347)

Range and Status. Endemic resident, wanderer and altitudinal migrant. Common to abundant in W African, equatorial and Congo Basin lowland rainforest zone where red on map, and probably almost throughout, although positively known at rather few locations in that part of forest zone shown pink. Patchily distributed, locally frequent, in mesic woodland savannas distant from main forest block, in E and SE Africa. Senegal, record of 10 in Casamance garden, 1977, may refer to S. cucullatus, although there are old records from W Guinea-Bissau. Mali, uncommon to locally frequent resident, 12 localities near Siby, Bougouni, Se´lingue´, Sikasso and Bamako. Common from Sierra Leone to Ghana; in Ivory Coast, mainly south of 7
N, but twice in Comoe´ Nat. Park (Salewski and Go ¨ken 2001). Togo, not uncommon, known from at least 18 localities, several c. 7
300 N, north to Djodji and Klabe´ Ape´game´ at 7
400 N. Benin, uncommon in SW, east to Noyau Central. Nigeria, common in SW north to Ile-Ife, frequent to common in SE north to Obudu and Mambilla Plateaux; in N, old records from Zungeru, and occasional locally on Jos Plateau. Cameroon, common north to Ngoume´; north of forest only in Tchabal Mbabo hills; on Mt Kupe´ frequent and widespread in cultivated areas at 350–900 m (Bowden 2001). Central African Republic, abundant in SW. Zaı¨re, north to Sudan border but not to E half of Central African Republic border. Sudan, common breeding visitor in Imatong and Didinga Mts, up to 2400 m; once Maridi. Ethiopia, frequent in SW, east to Rift Valley. Uganda, widespread, north to West Nile, Bunyoro, Acholi and Moroto; up to 2150 m in Ruwenzori Mts, Zaı¨re/Uganda. Kenya, poensis widespread, locally common up to 2000 m, from Mt Elgon and Saiwa Nat. Park to L. Victoria basin, Lolgorien and NW Mara Game Res.; nigriceps local in central highlands to Mathews Range, Mt Endau, Taita Hills and coastal lowlands from

Spermestes bicolor

?

?

Pink: see text. ?

Lamu to Tanzania border. Somalia, rare, flocks of up to 20, lower Jubba valley, Fanoole, Jilib and Boni Forest. South of equator, probably occurs in central Congo and S Uganda, and likely to be far more widespread than shown in S Zaı¨re, but from Angola and Tanzania to South Africa distribution patchy, as mapped. Density of 4–8 birds per ha (NE Gabon). Probably >10,000 birds in Sul do Save, Mozambique. Description. S. b. bicolor (Fraser): Guinea Bissau to S Nigeria, intergrading with poensis in SW Cameroon. ADULT Y: head, neck,

Spermestes bicolor however, most nestling faeces fall through loose structure of nest when dried out (van Someren 1956). Period: 15–17 days (Kenya); in Zimbabwe, 17±1 days; young birds may actually leave nest at 14 days, 2–3 days before they can fly. If disturbed prematurely in nest, they scatter in all directions, up to 50 m away; parents entice each one back by calling to move it gradually up to top of tree, parent flies from treetop back to nest, nestling flutters down after it and may have to be led up to another treetop before finally returning to nest (Woodall 1975). Out of nest, young keep with ads; they nibble at twigs, leaves and grass stalks, and are fed regularly by ads at c. 30 min intervals, clustering around parent with mouths wide open, sometimes raising one wing; young occasionally led back to nest and fed inside. BREEDING SUCCESS/SURVIVAL: eggs taken by climbing mice Dendromus sp. (Kenya) and snake Thrasops flavigularis

405

(Gabon), and nestlings and fledglings by African Goshawk Accipiter tachiro, Wahlberg’s Eagle Hieraaetus wahlbergi, coucals Centropus spp., Woodland Kingfisher Halcyon senegalensis, drongos Dicrurus, Pied Crow Corvus albus; ads taken by domestic cats, Fiscal Shrike Lanius collaris, Redthighed Sparrowhawk A. erythropus and Little Sparrowhawk A. minullus. Occasionally parasitized by Pin-tailed Whydah Vidua macroura. Max. longevity (recapture data) 4 years; 35% of ads at start of breeding season survive to following year; av. ad. life expectancy, 125 months (Woodall 1975).

Key References Akinpelu, A.I. (1994a, b), Brosset, A. and Erard, C. (1986), Clement, P. et al. (1993), Goodwin, D. (1982), Morris, D. (1957), Restall, R. (1996), van Someren, V.G.L. (1956), Thompson, H.S. (1989), Thompson, H.S. and Tye, A. (1995), Wolters and Clancey (1974), Woodall, P.F. (1975).

Spermestes bicolor (Fraser). Black-and-white Mannikin; Red-backed Mannikin. Capucin bicolore.

Plate 24

Amadina bicolor Fraser, 1843. Proc. Zool. Soc. London, 1842, p. 145; Cape Palmas, Liberia.

(Opp. p. 347)

Range and Status. Endemic resident, wanderer and altitudinal migrant. Common to abundant in W African, equatorial and Congo Basin lowland rainforest zone where red on map, and probably almost throughout, although positively known at rather few locations in that part of forest zone shown pink. Patchily distributed, locally frequent, in mesic woodland savannas distant from main forest block, in E and SE Africa. Senegal, record of 10 in Casamance garden, 1977, may refer to S. cucullatus, although there are old records from W Guinea-Bissau. Mali, uncommon to locally frequent resident, 12 localities near Siby, Bougouni, Se´lingue´, Sikasso and Bamako. Common from Sierra Leone to Ghana; in Ivory Coast, mainly south of 7
N, but twice in Comoe´ Nat. Park (Salewski and Go ¨ken 2001). Togo, not uncommon, known from at least 18 localities, several c. 7
300 N, north to Djodji and Klabe´ Ape´game´ at 7
400 N. Benin, uncommon in SW, east to Noyau Central. Nigeria, common in SW north to Ile-Ife, frequent to common in SE north to Obudu and Mambilla Plateaux; in N, old records from Zungeru, and occasional locally on Jos Plateau. Cameroon, common north to Ngoume´; north of forest only in Tchabal Mbabo hills; on Mt Kupe´ frequent and widespread in cultivated areas at 350–900 m (Bowden 2001). Central African Republic, abundant in SW. Zaı¨re, north to Sudan border but not to E half of Central African Republic border. Sudan, common breeding visitor in Imatong and Didinga Mts, up to 2400 m; once Maridi. Ethiopia, frequent in SW, east to Rift Valley. Uganda, widespread, north to West Nile, Bunyoro, Acholi and Moroto; up to 2150 m in Ruwenzori Mts, Zaı¨re/Uganda. Kenya, poensis widespread, locally common up to 2000 m, from Mt Elgon and Saiwa Nat. Park to L. Victoria basin, Lolgorien and NW Mara Game Res.; nigriceps local in central highlands to Mathews Range, Mt Endau, Taita Hills and coastal lowlands from

Spermestes bicolor

?

?

Pink: see text. ?

Lamu to Tanzania border. Somalia, rare, flocks of up to 20, lower Jubba valley, Fanoole, Jilib and Boni Forest. South of equator, probably occurs in central Congo and S Uganda, and likely to be far more widespread than shown in S Zaı¨re, but from Angola and Tanzania to South Africa distribution patchy, as mapped. Density of 4–8 birds per ha (NE Gabon). Probably >10,000 birds in Sul do Save, Mozambique. Description. S. b. bicolor (Fraser): Guinea Bissau to S Nigeria, intergrading with poensis in SW Cameroon. ADULT Y: head, neck,

406

ESTRILDIDAE

upperparts, andchintoupperbreast black with darkgreen gloss. Tail black,uppersidefaintlyglossy.Sidesoflowerbreastandflanksbarred black and white, the lateral row of feathers black with broad white fringes;restofunderpartswhite.Upperwingblack,tertialsandwingcovertsglossy;tertialswithafewsmallwhitespotsatbaseofouterweb, median and lesser coverts with dark green sheen like upperparts. Underwing-coverts and axillaries white. Bill light bluish grey; eyes dark brown, outer rim reddish; legs grey-brown or dark grey. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 48–51 (498), X 47–52 (493); tail, Y 28–33 (308), X 28–33 (303); bill, Y 12–13 (126), X 12–13 (125);tarsus,Y13–14(133),X125–14(131). WEIGHT:Sierra Leone (n ¼ 660), weights decline in dry season to min. of c. 90–95 in Mar–June, and are max. of c. 100 (YY) and 110 (XX) in Sept–Oct (Thompson and Tye 1995); Liberia, Y (n ¼ 6) 83–98 (93), X (n ¼ 7) 81–105 (95); Nigeria, av. weight varies seasonally from 8–9 g in Aug to 13–14 g in May (Akinpelu 1994a). IMMATURE: juv. dull, dark brown above, including sides of head and neck; underparts, including chin and throat, pale tawny-buff. Tail and flight feathers blackish brown; rest of upperwing dark brown, tertials narrowly fringed tawny-buff. NESTLING: skin of gape whitish and little swollen; buccal markings very like those of L. cucullata, palate with double horseshoe of black, tongue ringed, and inside of mandible with double crescent; ground colour of palate yellow, and of tongue flesh-coloured (Chapin 1954) (fig. p. 252). S. b. poensis (Fraser) (includes ‘stigmatophorus’): Bioko; S Cameroon to N Angola, Zaı¨re (south to Kasai), S Sudan, SW Ethiopia, Uganda, Kenya (west of Rift) and NW Tanzania (south to Gombe). Differs from nominate race in having narrow white barring on outer webs of flight feathers and tertials, forming large barred patch on closed wing; white spots and bars on rump feathers and uppertail-coverts; more than one row of white scaled feathers along sides of lower breast and flanks. Gloss slightly less intense, less green (more violet). Juvenile as nominate race, but wings barred. WEIGHT: Cameroon, Y (n ¼ 12) 8–11 (100); W Kenya, unsexed (n ¼ 8) 9–105 (94). S. p. nigriceps (Cassin) (includes ‘minor’): S Somalia, Kenya (east ˆ i and Zambia (except of Rift), Tanzania (except NW), Malaw where replaced by woltersi) to South Africa. Like poensis, but mantle, back, scapulars, tips and outer edges of tertials and greater coverts, and median and lesser coverts rufous-brown (not black). Juvenile paler, more tawny brown above than poensis. WEIGHT: Kenya, unsexed (n ¼ 42) 83–114 (95); Zimbabwe, Y (n ¼ 4) 84–93 (91), X (n ¼ 6) 80–98 (86), unsexed (n ¼ 121) 73–113 (92). S. p. woltersi Schouteden: SE Zaı¨re (W Katanga) to NW Zambia (N Mwinilunga). Like nigriceps but mantle duller and browner. TAXONOMIC NOTE: black-backed populations (bicolor, poensis) and the red-backed nigriceps were treated as separate species (S. bicolor, S. nigriceps) by Sibley and Monroe (1990), supposing them to be parapatric. However, poensis changes clinally from blackbacked in W to brown-backed in E, and the poorly known woltersi, intermediate in shade, may prove to link poensis and nigriceps clinally too. For that reason Hall and Moreau (1970) and Dowsett and Dowsett-Lemaire (1993) recognized only one species, as do White (1963), Goodwin (1982), Clement et al. (1993), Restall (1996), Zimmerman et al. (1996), Stevenson and Fanshawe (2002) and ourselves.

Field Characters. Length 85–10 cm. A much neaterlooking bird than Bronze Mannikin S. cucullatus, sharply black and white, with red back in E and S. Head, breast and upperparts entirely black in nominate race (Senegal to Cameroon), underparts white except for black bars on flanks; bill pale silver-blue. Other races have fine white barring on flight feathers and rump; white bars are so narrow that rump appears more black than white, whereas in Bronze Mannikin bars are broader and rump looks pale.

Lower border of black breast straight, not rounded. Immature less uniform than Bronze Mannikin, with sooty grey-brown or reddish brown upperparts and pale underparts; flanks buffy but throat and breast whitish. Voice. Tape-recorded (99, 104, B, C, F, ERA, GREG, GUT, PAY). Soft contact calls (given with bill closed: Chapin 1954), ‘tip’, ‘tyip’, ‘sip’ or ‘seep’. Louder calls from flock feeding in grass, high, shrill ‘pee’, ‘pew’, more sibilant ‘psew’ and longer, rather squeaky ‘psseew’, and a sweeter, piping ‘tyoo’; from perched group, long-drawn, downslurred ‘peeeya’, high-pitched but quite sweet. In flight, ‘pyip’ or ‘pyeep’; alarm short and harsh. Song said to be short and sweet (Bannerman 1949), apparently consisting of contact notes run together (Clement et al. 1993). Feeding call of nestlings, ‘cheep, cheep . . . ’; fledged young beg with shrill ‘ssi, ssi, ssi’ (Ginn et al. 1989). General Habits. Inhabits tall, rank grass in forest clearings and undergrowth; emergent, woody and grassy vegetation on riversides and riverbanks; wooded marshland, secondary growth near water, cultivated land, rice fields and native gardens. In Gabon, Paspalum in clearings; waste and fallow ground, recently abandoned plantations with young parasol trees and solanaceous plants; less associated with people than Bronze Mannikin. In southern Africa occurs in moist, broad-leaved woodland, evergreen and coastal forests, open dune forest and open bushveld. Usually in flock of 50 localities); up to 1000 m in Nimba and Wologizi Mts. Ivory Coast, distributed patchily from Soubre to Danane, Sipilou, Abengourou, Bouake´ and Yapo Forest (in clearings); several hundred resident in Dabou ricefields (Demey and Fishpool 1991). Ghana, uncommon (Kade, Cape Coast, Kumasi, Asukawkaw, Akwatia, regular Tafo). Togo, uncommon, coastal areas, in W around Badou, and north to Ahoue´-houe´. Benin, frequent in Be´te´rou area. Nigeria: common near Serti (several flocks of up to 20, Sept–Jan, Hall 1977a); frequent in Gashaka-Gumti Nat. Park (foothills of Chappal Hendu, Chappal Shirgu, Chappal Tale and Chappal Yumti, Sept–Dec, Green 1990); otherwise rare, records only in Shaffini Swamp Forest (L. Kainji), Epe (Lagos), Tayu, Panshanu (Jos

Spermestes fringilloides In Sierra Leone, moult very protracted, mainly in dry season, Jan–May; primary moult of juvs may be incomplete (Thompson and Tye 1995); in Nigeria, moults in May– Oct, between major and minor breeding seasons; duration c. 145 days (Akinpelu 1994a). Mainly sedentary, but breeding visitor to mountains in S Sudan, moving to low ground from Apr to July. Food. Mainly small grass seeds. In Sierra Leone, grass seeds compose 57% of diet (mainly Sorghum, 14%, and in Nov–Dec rice Oryza sativa, 9%, also Panicum laxum, Anadelphia tenuiflora, Andropogon guyana, Pennisetum, Brachiaria stigmatisata, Eleusine indica, Eragrostis aspera, Axonopus compressus and Paspalum); also fibrous mesocarp of fruits of Eugenia uniflora (Myrtaceae, 6% of diet) and oil palm Elaeis guineensis (Palmae, 16%) (Thompson 1989). In Nigeria, seeds of grasses Panicum, Digitaria, Sporobolus and Chloris; seeds of all of these fed to nestlings, and also seeds of Urochloa and Eragrostis (grasses) and Portulaca (Portulacaceae) (Akinpelu 1994b). In Kenya, principally seeds of Panicum maximum and Setaria sp. In captivity, South Africa, eats seeds of grasses P. maximum (ripe and unripe), Eleusine indica, Echinochloa pyramidalis, Digitaria sanguinalis, D. gazensis, Hyparrhenia hirta, Rhynchelytrum repens, Setaris homonyma, S. pallidefusca, Poa trivialis, Phalaris minor, Chloris guyana and rice. Petals of Schotia brachypetala (South Africa), nectar (Brickell and Konigkramer 1997), grass sap, and filamentous alga Spirogyra. Eats small invertebrates (12% of dietary items in Sierra Leone) including termites Macrotermes bellicosus, ants Oecophylla longinoda, beetles and flies; these and caterpillars and spiders are fed to nestlings. Breeding Habits. Solitary nester; monogamous. In Zimbabwe a nest with 6 young in Harpephyllum caffrum tree fell to ground; observer replaced it 1 m from original site; nest was attended by 3 YY and 2 XX, 1 or more of which

407

tore it to pieces during next 2 days; the young, all with full crops, fell to ground again and died (Howells 1990). NEST: thick-walled ball, densely made of coarse dry grass stems, dry leaves and palm fibres, which project around side entrance and can partially close it. Usually unlined. Sited in twiggy or leafy fork 1–12 m, usually 3–4 m, up in shrub or small tree; often in wild asparagus (Kenya); sometimes in house roof. Built by both parents. EGGS: 3–5 (Gabon); mainly 4–5 (Kenya); 2–7, usually 4– 6, av. 48 (southern Africa). White. SIZE: (n ¼ 5, Cameroon) 142–157  100–104 (146  103); (n ¼ 27, southern Africa) 134–160  97–111 (148  103). LAYING DATES: Guinea, (nests, Mar, June); Sierra Leone, Sept–Nov, coinciding with maturation of rice and availability of alga Spirogyra (Thompson 1989); Ghana, Mar–Aug; Nigeria, Apr–Dec; breeding distinctly bimodal, mainly in rains in Apr–Aug, also in Oct–Dec, very few records in Sept (Akinpelu 1994b); Gabon, Nov–Mar (in one year, Nov–May); Sudan, Sept–Oct; E Africa, breeds mainly in long rains: Region B, Apr–Aug, Oct–Nov, Region C, Feb, Region D, Oct and Dec–Aug, mainly Apr– June, Region E, May, July, Oct–Dec; Zambia, Jan–May; ˆ i, Dec–Apr; Zimbabwe, Nov–May, mainly Mar– Malaw Apr; Transvaal, Aug–Apr, KwaZulu-Natal, Sept–Apr. INCUBATION: parents relieve each other infrequently. Period 12 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: fresh asparagus often taken into nest when young have hatched (Kenya: van Someren 1956). Droppings not removed, but they soon dry out and crumble away; after some days young excrete at nest entrance. Nestling period 14 days (Kenya) or 17 days (in captivity, South Africa). Key References Akinpelu, A.I. (1994a, b), Clement, P. et al. (1993), Goodwin, D. (1982), Restall, R. (1996), van Someren, V.G.L. (1956), Thompson, H.S. (1989), Thompson, H.S. and Tye, A. (1995).

Spermestes fringilloides (Lafresnaye). Magpie Mannikin. Capucin pie.

Plate 24

Plocus fringilloides Lafresnaye, 1835. Mag. Zool. [Paris], 5, cl. 2, pl. 48; India, emended to Liberia by Hartlaub, 1857, Syst. Orn. Westafrika, p. 147.

(Opp. p. 347)

Range and Status. Endemic resident and wanderer; range patchy and highly localized. Senegal, rare; records from Niokolo Koba Nat. Park (Badi, 1955), Ke´dougou (1966), Se´dhiou (1988) and Basse-Casamance Nat. Park (1973; flock of 8 in 1988). Gambia, common to abundant 100 years ago, when thousands caught (by inference, annually) near Bathurst by bird traders (Bannerman 1949); since 1918 rare: Bakau, 1974, and Bijilo, 1989. Mali, uncommon, Mandingo Mts and Bamako, and 4 localities south of 12
N including Yanfolila and Kolondie´ba. Guinea, old records from W, near Sierra Leone border, but not found there by Demey (1995); frequent, Macenta area. Sierra Leone, common at Rotifunk and Njala (Harkrider 1993), rare at Bo; occurs in Little Scarcies valley, and irregularly in flocks near Freetown; uncommon, farmbush in Gola

Forest. Liberia, frequent to common throughout (>50 localities); up to 1000 m in Nimba and Wologizi Mts. Ivory Coast, distributed patchily from Soubre to Danane, Sipilou, Abengourou, Bouake´ and Yapo Forest (in clearings); several hundred resident in Dabou ricefields (Demey and Fishpool 1991). Ghana, uncommon (Kade, Cape Coast, Kumasi, Asukawkaw, Akwatia, regular Tafo). Togo, uncommon, coastal areas, in W around Badou, and north to Ahoue´-houe´. Benin, frequent in Be´te´rou area. Nigeria: common near Serti (several flocks of up to 20, Sept–Jan, Hall 1977a); frequent in Gashaka-Gumti Nat. Park (foothills of Chappal Hendu, Chappal Shirgu, Chappal Tale and Chappal Yumti, Sept–Dec, Green 1990); otherwise rare, records only in Shaffini Swamp Forest (L. Kainji), Epe (Lagos), Tayu, Panshanu (Jos

408

ESTRILDIDAE

Spermestes fringilloides

Plateau) and Uyo (Calabar, in 1954). Cameroon, uncommon to frequent, Kumba area (Serle 1950) and Yaounde´ area (Hall and Moreau 1970, Quantrill and Quantrill 1998); records near Mbalmayo, Nkolbisson, Ndibi, Eseka and Nguilili (R. Quantrill, pers. comm.). Gabon, common sporadically in Ivindo Basin, rare in Lope´ area, frequent but local in Gamba, Oyem, Moukalaba and Libreville areas. Congo, old record from Ouesso. Central African Republic, frequent in Lobaye and Bangui areas (6 localities, mainly Oct–Jan: Germain and Cornet 1994); record, S Bamingui-Bangoran Nat. Park (Hall and Moreau 1970). Zaı¨re, frequent to locally common, c. 35 localities as mapped; numerous around Kananga (Luluabourg); very common in Itombwe Highlands, at 800–1000 m (Bionga, Kamituga, Kitutu, Mulungu). Sudan, known from Bengengai and Maridi, and fairly common in Imatong Mts in Oct–May. Uganda, frequent in Bwamba lowlands; records in NE and near Kampala. Kenya, 1991 sight record at Alupe in W; Kitovu Forest (Taveta) in SE; formerly in Ganda Forest (Shimoni), and appears to have declined considerably in last 60 years. Ethiopia, 3, Didessa, 1233 m, July 1970, 1, same place, Sept 1995, and pair building nest, Kamashi (9
290 N, 35
490 E, 1485 m), June 2001 (J.S. Ash, pers. comm.). Rwanda, single record (Stevenson and Fanshawe 2002). Tanzania, c. 200 birds in Moshi area (N.E. Baker, pers. comm.); locally common in Amani, E Usambaras; moderately widespread in S and SE, as mapped (20–30 localities); uncommon in Zanzibar, mainly in NE (Potowa, Moga, Chutama: Pakenham 1979). South of about 10
S, very closely associated with Bindura bamboo Oxytenanthera abyssinica, except on Mozambique coastal plain. Distribution of bamboo is patchy and the plant, on whose seeds Magpie Mannikins ˆ i, flowering depend, flowers every 18–20 years; in Malaw cycle is of 30 years (F.A. McClure in Restall 1996). (Elsewhere in Africa any association with bamboo either does

not hold good or has been overlooked: Jackson 1972. In W Africa and on Mozambique coast associates with rice cultivation). Angola, rare nomad: scattered localities in W Malanje; also W Huı´la (Fazenda Mahita, Humpata) and S Cunene (Chitado). Zambia, very local, generally uncommon; c. 40 localities in 14% of Zambia squares (R.J. Dowsett, pers. comm.); was common in Copperbelt in early 1990s when ˆ i, sparse and bamboos were flowering (Beel 2000). Malaw local; Songwe R. mouth south to Karonga, Nkhotakota, Ntchisi, Mankhamba, Mitongwe, Zomba, Thondwe, ˆi Michiru, Blantyre, Thyolo, and above 1600 m on Malaw Hill. Mozambique, widespread in W Nampula, Zambe´zia and Sofala south to Beira, abundant in bamboo patches in Ile and Gurue districts (Vincent 1936); uncommon in Sul do Save, perhaps not more than 1000 birds, ‘being depleted by trapping for the cage bird trade’ (Parker 1999). Zimbabwe, range closely coincident with that of Bindura bamboo (mapped by Jackson 1972; Bindura bamboo poorly studied and range in other countries not well known; putative E African range mapped in Restall 1996). Local and erratic; mainly in N Mashonaland and NE, from Musengedzi R. gorge south through Sipolilo, Umvukwes and Shamva to Harare; in SE, Macheke R., Honde Valley and Stapleford south to Penhalonga, Umtali, Vumba area and Cashel, below 1700 m. Magpie Mannikins appeared in Salisbury (Harare) in 1967 and for some years depended on cultivated Bindura bamboo (see below). In SE, semi-nomadic and sporadic; becomes established in an area for a few years, then disappears, presumably when bamboo seeds exhausted (Irwin 1981). Namibia, once, Okavango River. Botswana, 3 at Kasane, Chobe R., Mar 1994. Transvaal, vagrant: small flock at Mataffin in 1976– 1984, old records at Sibasa and near Komatipoort. KwaZulu-Natal, rare, any month, records from at least 10 localities on or near coast: Pinetown, Malvern (small flocks), Berea and Mandini (breeding), Reunion Rocks, Durban, Kwa Mashu Sewage Farm (twice), Umpambinyoni Valley, Umgeni R., and St Lucia. Description. ADULT Y: head, neck, upper mantle, chin and throat black, glossed greenish blue. Lower mantle pale brown with tawny tinge, feathers with fine pointed buff spots at tips; scapulars and back darker brown; rump and uppertail-coverts black, glossed violet. Tail black, upperside glossed violet, T6 narrowly edged white. Sides of breast to lower rear sides of neck white, forming partial collar; pointed patch on side of lower breast glossy violet-black; flanks tawny buff, boldly blotched black and white; rest of underparts white or creamy white. Upperwing feathers glossy dark brown, inner median coverts with creamy white shafts. Underwing-coverts and axillaries creamy white. Bill black or dark grey above, bluish grey or pale silvery grey below; eyes dark brown or red-brown; legs dark grey to blackish. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 58–63 (608), X 59–63 (602); tail, Y 34–38 (360), X 32–35 (335); bill, Y 17–18 (176), X 16–175 (169); tarsus, Y 145–16 (155), X (145– 16 (153). WEIGHT: Liberia, Y (n ¼ 5) 15–177 (160), X (n ¼ 3) 15– 158 (153); Cameroon, Y (n ¼ 5) 13–18 (164), 1 imm. 13. IMMATURE: juv. dull brown above, darker, slightly glossy, on top of head and on rump and uppertail-coverts; cheeks and earcoverts paler, greyer brown; underparts pale buff, patches at side of breast dark brown; upperwing dull brown, tertials tipped pale buff, greater and median coverts tipped buff to form 2 poorly

Spermestes fringilloides marked wing-bars, inner median coverts with pale buff shafts; tail blackish brown. NESTLING: skin pink, gape pale grey, scarcely swollen. Bill white, with black tip; after 2 weeks, all black. At 3 days, eyes still closed, eyelids black. Palate with fine black horseshoe-shaped mark encircling internal nares and a broad dusky horseshoeshaped area outside it with intervening skin yellow; lining of throat dusky but tongue yellowish with black spot on each side, and beneath it, inside mandible, a double black crescent (Fig. p. 252). Down greyish white, sparse, only 3 mm long, on inner and outer supraorbital, occipital, spinal, humeral and femoral tracts (Serle 1950). TAXONOMIC NOTE: divided into 2 races, nominate and ‘pica’, by Clancey (1986); ‘pica’ has mantle less dark, smaller black marks on sides of breast, flank feathers less rusty and with smaller black marks; tail av. 1–2 mm shorter than tail of most nominate populations (but in NE Zaı¨re, tail 2–3 mm longer than in other nominate populations); ‘pica’ bill slightly less robust. Nominate birds range to north and ‘pica’ to south of straight line between Benguela (Angola) and Garissa (Kenya), almost the same line that divides the main taxa of S. bicolor and of S. cucullatus. Race ‘pica’ recognized by Restall (1996) but in our view larger samples need to be studied before it can be validated. Readily hybridizes in captivity with S. bicolor nigriceps (Restall 1996).

Field Characters. Length 11–13 cm. Largest mannikin; wide-ranging but much less common than Black-andwhite Mannikin S. bicolor, with which often confused. Long, heavy bill and short tail suggest ploceine weaver. Head solid black, bib slightly rounded, separated from black breast patch by wedge of white reaching onto side of neck. Back and wings warmer and darker brown than Bronze Mannikin S. cucullatus, with buffy streaks and dark mottling, and rump black, unbarred. White below with buffy wash on undertail-coverts, black and chestnut bars on flanks; upper mandible blackish, lower blue-grey. Immature brown above, darker on crown and face, buff below, brightest on flanks, told from other mannikins by large size, heavy bill and black rump, from Bronze by unbarred undertail-coverts. Voice. Tape-recorded (99, 104, C, CART, GUT, McVIC). Loud piping contact calls from flock, descending ‘pee-yoo’ and rising ‘whee’, often in combination, ‘pyoo-wee’ or ‘pyoo-wee-yoo’. In flight a shorter, high ‘chee’. Song variously reported as a repeated short note similar to the contact note (Clement et al. 1993), a ‘single stereotyped strophe’ (Goodwin 1982), or as soft and bubbling (Maclean 1993). Alarm, thin ‘cheep’ or ‘tsek’; begging call of young, ‘t-t-ttey-tey’. General Habits. Inhabits wet and drying wet-rice fields, dry-soil-rice patches in forest, millet Pennisetum fields (Gabon), savanna and woodland mosaic, newly cleared savanna with relict trees, rank grass in abandoned plantations, thickets near cultivation, farmbush, farmsteads, villages and gardens; forest edges; follows logging roads into forest clearings and large treefall gaps, where sometimes nests (Liberia). Hyparrhenia grassland in Combretum/Terminalia savanna (Ethiopia). Seldom far from water. In southern Africa, inhabits Bindura bamboo Oxytenanthera abyssinica thickets along rivers and sometimes reeds and sedges; margins of lowland evergreen

forest; occasionally Acacia woodland (Zimbabwe); coastal bush and rank vegetation near water (KwaZulu-Natal). Generally in pairs or flocks of c. 5, sometimes 10–15, uncommonly 20–40, rarely up to 500 (Zimbabwe). Flock may comprise 1 or more families, e.g. 4 ads and 6 juvs. Keeps in flock year-round, even when breeding (Vincent 1936). Flock feeds on ground, but perhaps more commonly on seeds in standing herbs, grasses, especially rice in W Africa and Bindura bamboo inflorescences in southern Africa. In Feb 1967 ‘indigenous bamboos in my garden seeded in profusion and subsequently died . . . had not happened for 20 years . . . In the same month a large flock of 30–40 . . . appeared and, I think, fed exclusively on these seeds . . . mannikins remained here in strength for 2 years and must have found other food . . . by the end of 1970 there were very few left’ (D. Searson in Jackson 1972). Shy; flies to thick cover when disturbed. Agile when feeding; alights upon grass stem, and quickly works its way up to seeding head. Bird’s weight may make stem bend down, when bird continues feeding, hanging upside down. Stands on broken or fallen stem and pecks seeds from grass head. Forages alongside either Bronze Mannikins or Black-andwhite Mannikins S. bicolor; occasionally all 3 species feed together. 2 Magpie, 8 Black-and-white and many Bronze Mannikins foraged at edge of small, shallow pool; 1 Magpie and 1 Black-and-white perched on broken reeds 5 cm above surface covered with green alga Spirogyra, and ate strands up to 15 cm long, ‘fishing it out and eating it non-stop’ for 20 min (Hall 1981). Extracts sap from grass stems in aviary (Brickell et al. 1980), by nibbling. Bathes frequently, up to 7 times daily in showery weather or when running water at hand (Brickell et al. 1990). A large colony at Rotifunk, Sierra Leone, roosted in mangoes and citrus trees in a garden (Kemp 1905); often roosts in reedbeds; builds nests specifically for roosting. Sedentary; in W Africa, nomadic in south of range. Food. Small seeds of grasses, cereals (rice, millet) and herbs; seeds of Bindura bamboo, which measure 125– 160  20–25 mm. Eats grass sap and Spirogyra algae (see above); in captivity, large quantities of watercress and spinach, seeds of grasses Panicum aequinerve, P. deustum, P. maximum, P. laevifolium, Sorghum halapense, S. versicolor, Poa annua, Sporobolus fimbriatus and Eragrostis curvula, and buds, petals, leaves and seeds of composites Schkuhria pinnata and Bidens pilosa (Brickell 1981), and dying leaves of Ficus vogelii (Brickell and Konigkramer 1997). Also winged termites, small ants and (in captivity) mealworms; taste for animal food varies individually. Nestlings fed on vegetable matter and a few insects. Breeding Habits. Nests solitarily or in small colonies, sometimes with Bronze Mannikins, e.g. colony in suburban almond tree, Cameroon, of 3 Magpie Mannikin nests with eggs or young and one Bronze Mannikin nest with eggs, with another Magpie Mannikin nest with young in adjacent tree (Serle 1950). Colonies in captivity, South Africa, each comprised 3–4 nests built together as one large structure (Brickell et al. 1980). Evidently monogamous. Displaying Y, holding straw in bill, flies to X, then to nest site. Singing Y perches rather upright, body close to perch, head slightly

409

412

ESTRILDIDAE

downturned, bill open (Restall 1966), head turning from side to side through small arc; in courtship he angles body at c. 45
and bobs up and down by bending legs. NEST: rough, untidy, loose ball, made of dry (sometimes green) grass stems and bents 50–150 long and at times a few leaves; walls up to 25 mm thick; side entrance without porch or with one up to 15–35 long and 20–50 diam.; entrance may be placed above middle of nest side, and in South Africa nearly always faces west. Lined with fine seeding grass heads and bamboo flowers; once with grass heads Eragrostis curvula, seeding reed heads Phragmites australis, and leaves of ironwood Vepris lanceolata. Sometimes oval, when entrance at one end. SIZE: length 125– 200, from entrance to rear 110–130, height 120–130, int. diam. 65; also (averages of 8 aviary nests), length 210, width 100, height 70, entrance height 38 and width 51. 6 aviary nests comprised 335–370 (355) pieces of material and 52–83 (68) feathers (Brickell et al. 1980). Sited 2–5 m (usually 4–5 m) above ground in tall shrub or sapling, bamboo, palm, citrus, Euphorbia, Acacia burkei tree or exotic conifer. 10 nests in an aviary were all in colonies in Hypericum revolutum bushes, at av. height above ground of 20 m. Y brings material to nest where X remains and works it in; nest built in 7–10 days. Nests made only for roosting are not known to differ from those for breeding. ˆ i, EGGS: 2–9 (southern Africa, usually 4–6), 2–5 (Malaw 6 clutches av. 33), 6 (Nigeria, Zaı¨re). Laid daily. Immaculate white; ovate, smooth, not glossy. SIZE: (Cameroon, n ¼ 7) 156–170  111–124 (162  119), (Zaı¨re, Itombwe, n ¼ 2) 156  112, 156  114, (southern Africa, n ¼ 25) 146– 178  109–123 (162  115); av. in collection of Nat.

Hist. Mus. (London) 153  113; av. of 20 aviary eggs, South Africa, 16  12. LAYING DATES: Guinea, (nest-building July–Aug); Liberia, (nest-building and nestlings Nov–Feb); Ghana, Apr, (and juvs July, Oct); Togo, (juvs, display, Jan; nestbuilding, Feb); Nigeria, (nest-building Shaffini, Feb, and Serti, Nov); Cameroon, July–Sept (mainly Sept); Gabon, (gonads max. size Nov–Dec); Zaı¨re, Avakubi, Oct–Nov; appears to nest all year in Ituri and Itombwe; Uganda, ˆ i, Jan–May, July, Oct; Apr–Sept; Zambia, Feb, Mar; Malaw Zimbabwe, Dec–Jan, Mar–June (10 out of 14 clutches in Dec and Mar–Apr); Mozambique, Oct–Mar; KwaZuluNatal, Apr. INCUBATION: on leaving nest incubating bird pulls feathers across inside of entrance to block it. Period: (n ¼ 12) 14–16 (15) days. DEVELOPMENT AND CARE OF YOUNG: nestlings period: 21– 26 days (Brickell et al. 1980); or 19–22 days. Fledglings beg with wings stretched out, mouths wide open, giving shrill ‘ssi, ssi, ssi . . . ’ calls. Full ad. plumage attained in 49– 55 days; earlier, young YY carry grass; they breed soon afterwards (Brickell et al. 1980). BREEDING SUCCESS/SURVIVAL: nestlings taken by cobra Dendroaspis jamesoni and hawk Polyboroides typus (Zaı¨re). Of 15 aviary clutches of 4–6 eggs, 3–4 nestlings survived (Brickell et al. 1980). Key References Bannerman, D.A. (1949), Brickell, N. (1981, 1986), Brickell, N. et al. (1980), Clancey, P.A. (1986), Clement, P. et al. (1993), Goodwin, D. (1982), Irwin, M.P.S. (1981), Jackson, H.D. (1972), Maclean, G.L. (1993), Restall, R. (1996), Vincent, J. (1936).

Genus Euodice Reichenbach

Pallid mannikins of arid country, with silver bills, white underparts, and longer, more pointed tails than Spermestes and most Lonchura spp., with which, on genetic evidence, quite distantly related (R.B. Payne, pers. comm.). 2 species, one African and SW Arabian, one E Arabian and Indian, forming a superspecies.

Plate 24

Euodice cantans (Gmelin). African Silverbill. Capucin bec d’argent.

(Opp. p. 347)

Loxia cantans Gmelin, 1789. Syst. Nat., 1 (2), p. 859; Africa (restricted to Dakar, Senegal, by W.L. Sclater and C. Mackworth-Praed, 1918, Ibis, p. 440). Forms a superspecies with E. malabarica (Israel to Bangladesh; record S Sinai, Bull. Afr. Bird Club 2001, 149). NOMENCLATURAL NOTE:

commonly called Lonchura malabarica.

Range and Status. Africa and SE Arabia; resident, and wide-ranging wanderer or partial migrant near N and S borders of range, along S Sahara and in E Africa. Algeria, 1952 record at Tamanrasset, SW Hoggar Massif; 2 at Amsel (20 km south of Tamanrasset); and breeding

colony of c. 60 birds established at Abalessa (22
550 N, 4
520 E, 65 km west of Tamanrasset) in about 1996 (Belbachir 2000). Mauritania, frequent and widespread in S, north to 17
N in wet season; frequent around Nouakchott in 1974 to 1980 but not since; present in

412

ESTRILDIDAE

downturned, bill open (Restall 1966), head turning from side to side through small arc; in courtship he angles body at c. 45
and bobs up and down by bending legs. NEST: rough, untidy, loose ball, made of dry (sometimes green) grass stems and bents 50–150 long and at times a few leaves; walls up to 25 mm thick; side entrance without porch or with one up to 15–35 long and 20–50 diam.; entrance may be placed above middle of nest side, and in South Africa nearly always faces west. Lined with fine seeding grass heads and bamboo flowers; once with grass heads Eragrostis curvula, seeding reed heads Phragmites australis, and leaves of ironwood Vepris lanceolata. Sometimes oval, when entrance at one end. SIZE: length 125– 200, from entrance to rear 110–130, height 120–130, int. diam. 65; also (averages of 8 aviary nests), length 210, width 100, height 70, entrance height 38 and width 51. 6 aviary nests comprised 335–370 (355) pieces of material and 52–83 (68) feathers (Brickell et al. 1980). Sited 2–5 m (usually 4–5 m) above ground in tall shrub or sapling, bamboo, palm, citrus, Euphorbia, Acacia burkei tree or exotic conifer. 10 nests in an aviary were all in colonies in Hypericum revolutum bushes, at av. height above ground of 20 m. Y brings material to nest where X remains and works it in; nest built in 7–10 days. Nests made only for roosting are not known to differ from those for breeding. ˆ i, EGGS: 2–9 (southern Africa, usually 4–6), 2–5 (Malaw 6 clutches av. 33), 6 (Nigeria, Zaı¨re). Laid daily. Immaculate white; ovate, smooth, not glossy. SIZE: (Cameroon, n ¼ 7) 156–170  111–124 (162  119), (Zaı¨re, Itombwe, n ¼ 2) 156  112, 156  114, (southern Africa, n ¼ 25) 146– 178  109–123 (162  115); av. in collection of Nat.

Hist. Mus. (London) 153  113; av. of 20 aviary eggs, South Africa, 16  12. LAYING DATES: Guinea, (nest-building July–Aug); Liberia, (nest-building and nestlings Nov–Feb); Ghana, Apr, (and juvs July, Oct); Togo, (juvs, display, Jan; nestbuilding, Feb); Nigeria, (nest-building Shaffini, Feb, and Serti, Nov); Cameroon, July–Sept (mainly Sept); Gabon, (gonads max. size Nov–Dec); Zaı¨re, Avakubi, Oct–Nov; appears to nest all year in Ituri and Itombwe; Uganda, ˆ i, Jan–May, July, Oct; Apr–Sept; Zambia, Feb, Mar; Malaw Zimbabwe, Dec–Jan, Mar–June (10 out of 14 clutches in Dec and Mar–Apr); Mozambique, Oct–Mar; KwaZuluNatal, Apr. INCUBATION: on leaving nest incubating bird pulls feathers across inside of entrance to block it. Period: (n ¼ 12) 14–16 (15) days. DEVELOPMENT AND CARE OF YOUNG: nestlings period: 21– 26 days (Brickell et al. 1980); or 19–22 days. Fledglings beg with wings stretched out, mouths wide open, giving shrill ‘ssi, ssi, ssi . . . ’ calls. Full ad. plumage attained in 49– 55 days; earlier, young YY carry grass; they breed soon afterwards (Brickell et al. 1980). BREEDING SUCCESS/SURVIVAL: nestlings taken by cobra Dendroaspis jamesoni and hawk Polyboroides typus (Zaı¨re). Of 15 aviary clutches of 4–6 eggs, 3–4 nestlings survived (Brickell et al. 1980). Key References Bannerman, D.A. (1949), Brickell, N. (1981, 1986), Brickell, N. et al. (1980), Clancey, P.A. (1986), Clement, P. et al. (1993), Goodwin, D. (1982), Irwin, M.P.S. (1981), Jackson, H.D. (1972), Maclean, G.L. (1993), Restall, R. (1996), Vincent, J. (1936).

Genus Euodice Reichenbach

Pallid mannikins of arid country, with silver bills, white underparts, and longer, more pointed tails than Spermestes and most Lonchura spp., with which, on genetic evidence, quite distantly related (R.B. Payne, pers. comm.). 2 species, one African and SW Arabian, one E Arabian and Indian, forming a superspecies.

Plate 24

Euodice cantans (Gmelin). African Silverbill. Capucin bec d’argent.

(Opp. p. 347)

Loxia cantans Gmelin, 1789. Syst. Nat., 1 (2), p. 859; Africa (restricted to Dakar, Senegal, by W.L. Sclater and C. Mackworth-Praed, 1918, Ibis, p. 440). Forms a superspecies with E. malabarica (Israel to Bangladesh; record S Sinai, Bull. Afr. Bird Club 2001, 149). NOMENCLATURAL NOTE:

commonly called Lonchura malabarica.

Range and Status. Africa and SE Arabia; resident, and wide-ranging wanderer or partial migrant near N and S borders of range, along S Sahara and in E Africa. Algeria, 1952 record at Tamanrasset, SW Hoggar Massif; 2 at Amsel (20 km south of Tamanrasset); and breeding

colony of c. 60 birds established at Abalessa (22
550 N, 4
520 E, 65 km west of Tamanrasset) in about 1996 (Belbachir 2000). Mauritania, frequent and widespread in S, north to 17
N in wet season; frequent around Nouakchott in 1974 to 1980 but not since; present in

412

ESTRILDIDAE

downturned, bill open (Restall 1966), head turning from side to side through small arc; in courtship he angles body at c. 45
and bobs up and down by bending legs. NEST: rough, untidy, loose ball, made of dry (sometimes green) grass stems and bents 50–150 long and at times a few leaves; walls up to 25 mm thick; side entrance without porch or with one up to 15–35 long and 20–50 diam.; entrance may be placed above middle of nest side, and in South Africa nearly always faces west. Lined with fine seeding grass heads and bamboo flowers; once with grass heads Eragrostis curvula, seeding reed heads Phragmites australis, and leaves of ironwood Vepris lanceolata. Sometimes oval, when entrance at one end. SIZE: length 125– 200, from entrance to rear 110–130, height 120–130, int. diam. 65; also (averages of 8 aviary nests), length 210, width 100, height 70, entrance height 38 and width 51. 6 aviary nests comprised 335–370 (355) pieces of material and 52–83 (68) feathers (Brickell et al. 1980). Sited 2–5 m (usually 4–5 m) above ground in tall shrub or sapling, bamboo, palm, citrus, Euphorbia, Acacia burkei tree or exotic conifer. 10 nests in an aviary were all in colonies in Hypericum revolutum bushes, at av. height above ground of 20 m. Y brings material to nest where X remains and works it in; nest built in 7–10 days. Nests made only for roosting are not known to differ from those for breeding. ˆ i, EGGS: 2–9 (southern Africa, usually 4–6), 2–5 (Malaw 6 clutches av. 33), 6 (Nigeria, Zaı¨re). Laid daily. Immaculate white; ovate, smooth, not glossy. SIZE: (Cameroon, n ¼ 7) 156–170  111–124 (162  119), (Zaı¨re, Itombwe, n ¼ 2) 156  112, 156  114, (southern Africa, n ¼ 25) 146– 178  109–123 (162  115); av. in collection of Nat.

Hist. Mus. (London) 153  113; av. of 20 aviary eggs, South Africa, 16  12. LAYING DATES: Guinea, (nest-building July–Aug); Liberia, (nest-building and nestlings Nov–Feb); Ghana, Apr, (and juvs July, Oct); Togo, (juvs, display, Jan; nestbuilding, Feb); Nigeria, (nest-building Shaffini, Feb, and Serti, Nov); Cameroon, July–Sept (mainly Sept); Gabon, (gonads max. size Nov–Dec); Zaı¨re, Avakubi, Oct–Nov; appears to nest all year in Ituri and Itombwe; Uganda, ˆ i, Jan–May, July, Oct; Apr–Sept; Zambia, Feb, Mar; Malaw Zimbabwe, Dec–Jan, Mar–June (10 out of 14 clutches in Dec and Mar–Apr); Mozambique, Oct–Mar; KwaZuluNatal, Apr. INCUBATION: on leaving nest incubating bird pulls feathers across inside of entrance to block it. Period: (n ¼ 12) 14–16 (15) days. DEVELOPMENT AND CARE OF YOUNG: nestlings period: 21– 26 days (Brickell et al. 1980); or 19–22 days. Fledglings beg with wings stretched out, mouths wide open, giving shrill ‘ssi, ssi, ssi . . . ’ calls. Full ad. plumage attained in 49– 55 days; earlier, young YY carry grass; they breed soon afterwards (Brickell et al. 1980). BREEDING SUCCESS/SURVIVAL: nestlings taken by cobra Dendroaspis jamesoni and hawk Polyboroides typus (Zaı¨re). Of 15 aviary clutches of 4–6 eggs, 3–4 nestlings survived (Brickell et al. 1980). Key References Bannerman, D.A. (1949), Brickell, N. (1981, 1986), Brickell, N. et al. (1980), Clancey, P.A. (1986), Clement, P. et al. (1993), Goodwin, D. (1982), Irwin, M.P.S. (1981), Jackson, H.D. (1972), Maclean, G.L. (1993), Restall, R. (1996), Vincent, J. (1936).

Genus Euodice Reichenbach

Pallid mannikins of arid country, with silver bills, white underparts, and longer, more pointed tails than Spermestes and most Lonchura spp., with which, on genetic evidence, quite distantly related (R.B. Payne, pers. comm.). 2 species, one African and SW Arabian, one E Arabian and Indian, forming a superspecies.

Plate 24

Euodice cantans (Gmelin). African Silverbill. Capucin bec d’argent.

(Opp. p. 347)

Loxia cantans Gmelin, 1789. Syst. Nat., 1 (2), p. 859; Africa (restricted to Dakar, Senegal, by W.L. Sclater and C. Mackworth-Praed, 1918, Ibis, p. 440). Forms a superspecies with E. malabarica (Israel to Bangladesh; record S Sinai, Bull. Afr. Bird Club 2001, 149). NOMENCLATURAL NOTE:

commonly called Lonchura malabarica.

Range and Status. Africa and SE Arabia; resident, and wide-ranging wanderer or partial migrant near N and S borders of range, along S Sahara and in E Africa. Algeria, 1952 record at Tamanrasset, SW Hoggar Massif; 2 at Amsel (20 km south of Tamanrasset); and breeding

colony of c. 60 birds established at Abalessa (22
550 N, 4
520 E, 65 km west of Tamanrasset) in about 1996 (Belbachir 2000). Mauritania, frequent and widespread in S, north to 17
N in wet season; frequent around Nouakchott in 1974 to 1980 but not since; present in

Euodice cantans

Euodice cantans

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?

?

?

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?

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?

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central massifs north to 20
N (16 localities in Assaba, Tagant and Adrar); once Atar. Senegal, common north of 15
N, uncommon further south, once in Casamance, absent from SE, except for record east of Gambia (Sauvage and Rodwell 1998). Gambia, frequent on Kotu beaches; not uncommon locally at a few locations inland, mainly north of river, east to Kaur; once Bansang. Mali, common and widespread, north to 17
N; south to Boucle du Baoule´ and in Niger valley to Guinea border, mainly in dry season (July–Sept). Burkina Faso, frequent resident in N Yatenga, uncommon in gardens of Ouagadougou and nearby (Gonse´, Koubri); record in S, near Ghana/Togo border (Hall and Moreau 1970); rare, Arli-Pendjari Nat. Park, across Benin border. Niger, frequent in Niger floodplain, common resident in ‘W’ Nat. Park and occurs around Ayorou, Tillabe´ri and Niamey; in S, regular and not infrequent south of 15
N; further north, occurs between Cheri and Maine and south of Nguigmi; further north still (Zone 5: Giraudoux et al. 1988) resident and locally numerous; in NW, numerous between Arlit and Akokan; in Aı¨r Massif, widespread and sometimes numerous (Aoude´ras, Belkusi, Bagzans Mts, Kori Tessouba, Tazole´; absent from Te´ne´re´ except around Bilma oasis, where common, perhaps introduced long ago by Tuareg caravans). Ghana, uncommon in far N: Tumu, Mole, Bolgatanga (where regular), Bongo Hills, Bawku. Togo, uncommon in far N, 6 localities south to Naboulgou (10
090 N). Nigeria, common resident in N and NE, south to Kainji Lake Nat. Park, Zaria (frequent), Jos Plateau and Yankari Game Res. (uncommon); well established population around Lagos, probably originated from escapes. Cameroon, common in N, south to Benue Plain and (once) Tello on Adamawa Plateau. Chad, common throughout sahelian zone, much less so in soudanian zone; north to Ouadi Rime-Ouadi Achim Faunal Res., Biltine and Ennedi regions and occasionally to Faya-Largeau.

Central African Republic, uncommon, N Vakaga Pre´fecture. Sudan, north of 8
N, common and widespread as mapped, locally very common, at least in dry season; up to 2300 m on Jebel Marra; orientalis uncommon in SE, west to Maridi. Egypt/Sudan, only in Gebel Elba: known from Wadi Rabdeit, Gebel Shellal, Wadi Aideib. Eritrea, frequent to locally abundant throughout, except for Dancalia and offshore islands; at 50–2500 m (Zinner 2001), mainly below 1800 m. Ethiopia: in border country in NW; throughout arid Adal-Esa, between Djibouti and Rift Valley; frequent in SW and (possibly a separate population) in SE. Djibouti, throughout, uncommon but locally frequent (e.g. Le He´ron, Djibouti City). Somalia, uncommon resident in NW and Webi Shabeelle valley; also in upper Jubba valley, northwest of Luuq; elsewhere rare. Uganda, Kidepo Valley Nat. Park. Kenya, mainly at 400– 1300 m but up to 1600 m in SW; locally common along Sudan border, west and east of L. Turkana and south of it to equator; uncommon in far NE; frequent in S from Loita, Kedong, L. Magadi and Olorgesailie to Tsavo East. Tanzania, as mapped, south to Njombe valley in S Dodoma (N.E. Baker, pers. comm.). Coastal populations at Lamu (Kenya) and Dar es Salaam (Tanzania) may have originated as escapes. Description. E. c. cantans (Gmelin): Senegal to central Sudan; intergrades with orientalis in E Sudan and Eritrea. ADULT Y: forehead and crown buffy brown with dark shaft marks and pale buff fringes, giving dark-spotted or scaly effect; mottling becomes progressively more diffuse on buffy brown nape and hindneck. Mantle, scapulars and back buffy brown, feathers with faint darker cross-barring; rump and uppertail-coverts black. Tail black, rather long, well graduated, central feathers narrowly pointed and elongated. Lores, supercilium, cheeks and ear-coverts buff, spotted dark brown. Chin and upper throat buff, spotted or barred tawny brown; breast and flanks pale buff with diffuse tawny brown barring, plainer on centre of breast; lower breast, belly, thighs and undertail-coverts buffy white, almost unbarred. Flight feathers dark brown, outer webs of primaries and outer secondaries blackish, outer webs of inner secondaries barred buff; tertials buffy brown, barred darker brown, each with blackish subterminal bar and buffy white tip; primary coverts and alula blackish brown; greater coverts barred dark brown and buff, outer feathers edged blackish; median and lesser coverts barred dark brown and blackish, outer medians tinged blackish. Underwingcoverts and axillaries pale buff. Bill pale silvery blue-grey, sometimes with culmen or entire upper mandible dark grey; eyes brown to blackish; legs pinkish grey to flesh. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 53–55 (542), X 52–56 (543); tail, Y 36–49 (422), X 34–43 (395); bill, Y 11–12 (117), X 11–12 (116); tarsus, Y 125–14 (134), X 125–14 (134). WEIGHT: (N Nigeria, unsexed, n ¼ 14) 98–120 (108); (Chad, unsexed, n ¼ 36) 10–14 (120). IMMATURE: juv. differs from ad. in having crown diffusely streaked, no sharp spotting on head; lacks barring on back and wing feathers. NESTLING: skin dark; waxy, yellow gape swellings; inside mouth pale, with single black circle on palate and around tongue. E. c. orientalis Lorenz and Hellmayr (includes ‘inornata’): Sudan to Ethiopia, Somalia, Kenya and N Tanzania. Slightly browner above than nominate race, more strongly barred; slightly whiter (less buff) below, chin and upper throat more sparsely marked with tawny brown. WEIGHT: (Kenya, unsexed, n ¼ 51) 85–127 (107). TAXONOMIC NOTE: now widely accepted to be a full species separate from E. malabarica (Kakizawa and Watada 1985, Sibley and Monroe 1990, Restall 1996, R.B. Payne pers. comm.).

413

414

ESTRILDIDAE

Field Characters. Length 10–115 cm. A pale dry-country finch with silver-grey bill, black graduated tail with pointed central feathers; differs from Grey-headed Silverbill Odontospiza griseicapilla in black rump and flight feathers, buffy brown head and body with speckling on crown but not on face. Black-rumped Waxbill Estrilda troglodytes has red bill and eye-stripe. Voice. Tape-recorded (104, B, CHA, GUT, HAM, HAZ, MOR). Song a continuous low sweet warbling, notes almost pure but with a slight burr. Other songs higher-pitched: 2 long-drawn trills, different in tone, and a more complex 3-part type, starting with a series of rapidly repeated descending notes, followed by slurred double notes, accelerating and descending in pitch, ending with a ‘small interrogative-sounding phrase of about 4 notes’ (Goodwin 1982). Flight calls from group, excited ‘chip’ or ‘chyup’ and fast ‘cheet-cheet-cheet’. General Habits. Inhabits sub-desert steppe and villages, wadis with shrubs, sparse grass, Acacia raddiana, Tamarix aphylla and Calotropis procera trees, sorghum and millet crops (N Chad, S Algeria), rocky savanna with scattered trees, short-grass floodplains, edges of marshes and low waterside bushes, farmland, homesteads, gardens, scattered thorn shrubs in eroded areas, and sandy country with Borassus palms (10
N, Nigeria). In pairs or family parties of c. 6 birds and flocks of 12–30, rarely 40; once, 300–500 near Hargeysa, N Somalia, and even larger flock of migrants in Mali; also, hundreds around water holes at end of dry season, Far North Prov., Cameroon (Scholte et al. 1999). Pairs and small parties can be inconspicuous, rather secretive and inactive, and easily overlooked. Occurs in flocks all year. Generally tame. Forages on ground, birds keeping close together; flock in flight less close, sometimes straggling; readily perches in shrubs, small flock huddling together, perched in a row, for long spells. Feeds also in grasses, clinging to stem and pecking seeds from inflorescences. Roosts at night communally. Mainly resident. In Mali, migratory flocks of 50–1000 birds occur in wet season; a dry season visitor to Missira, Boucle du Baoule´ (de Bie and Morgan 1989); may move southward in N Ghana for dry season; mainly dry season visitor in N Sudan; resident in SE Sudan, ranging west to Maridi in dry season (Nikolaus 1989); wanders in E Africa (and in SW Arabia); onset of rains prompts migration to drier regions (Restall 1996). Food. Seeds, mainly of grasses, taken from ground and herbage. A few small invertebrates, including aphids. In captivity, eats seeds of Chloris guayana, Echinochloa crusgalli, Panicum maximum and Stellaria. Captive birds feed nestlings on seeds and small insects (Brickell and Konigkramer 1997) or wholly on seeds even when invertebrates readily available (Restall 1996). Breeding Habits. Solitary nester (but once c. 12 nests close together: Belbachir 2000); monogamous or perhaps polygamous (see below). Courting Y grasps grass stem at 1 end, hops or flies to be near X, sleeks feathers down, stands

upright with tail pointing straight down, jerks head upward a few times; he leans forward, twisting tail a little towards X, fluffs flank and belly feathers out, then usually drops straw and starts singing, stretching neck and fluffing neck feathers out somewhat, accompanied by bobbing movement of body as legs are bent and stretched (Restall 1996). Successful copulation followed by bill-fencing and allopreening. In ‘undirected advertisement song display’, Y starts with side to side twisting movement of trunk, fluffs out flank feathers little or not at all, and either grasps straw or sings, sways and bobs on legs. NEST: globular bundle of fine, dry, yellow grass stems, inflorescences and leaves and soft, pliable herb stems, with entrance at side, sometimes pointing obliquely upward and occasionally with small porch. In one nest, stems c. 37 cm long, mixed with numerous flaky seeds and a few feathers. Another nest made of entangled Tamarix aphylla twigs. Lined with soft fibres and occasionally feathers. Placed near end of branch of small thorn tree or 1–15 m up in shrub in thicket; sometimes in date palm (Sudan); often around human habitation, in hedge, creeper, thick bush, thatched roof or rolled-up sunblind; once in a street lamp. Nest twice sited near wasps’ nest. Built by both Y and X; Y collects all nest material. Rarely, captive X brings grass bent to nest under construction and drops it there (Restall 1996). Regularly adopts old weaver nests. Of 39 nests in Nigeria, 2 were made by African Silverbills and 37 were in old nests of Ploceus cucullatus, P. velatus and P. heuglini (Mundy and Cook 1974); of 25 nests in Kenya, one was made by Silverbills and all others were in old nests of Ploceus spekei, P. intermedius, P. rubiginosus and P. vitellinus (van Someren 1956). In contrast, of 25 nests in Senegal, 24 made by Silverbills and only 1 in old Ploceus nest (Morel and Morel 1962). Commonly uses nests of P. galbula (Oman). Pair once examined old nest of Scaly-fronted Weaver Sporopipes frontalis. When Silverbills take over old weaver nest they line it copiously with feathers (of e.g. hen or Helmeted Guineafowl Numida meleagris). EGGS: 4 –7 (–8 in captivity), usually 3–5. Good evidence that 2 XX may lay in same nest (van Someren 1956, Mundy and Cook 1974). In 11 nests, Nigeria, only 1–2 eggs laid; ‘perhaps the XX had forgotten their own nests and continued to lay in other occupied nests’ (Mundy and Cook 1974); or, YY may be polygamous (van Someren 1956; 2 eggs laid on one day in each of 4 nest: Mundy and Cook 1974). Clutch often added to deserted clutch of original owners of nest taken over by African Silverbills; ‘in addition, a nest now occupied by Euodice sometimes held the eggs of Cut-throat Finch Amadina fasciata . . . a very complicated situation’ (van Someren 1956). Oval, smooth, white, dull and glossless when just laid but shiny and porcelain-like when well incubated. SIZE: Nigeria (n ¼ 3) 142–153  108–119 (148  114) and (n ¼ 9) 120–180  105–145 (155  110). WEIGHT: av. (n ¼ 11) 10 g. LAYING DATES: Mauritania, Oct–May, mainly Oct–Apr; Senegal, Sept–May (50 clutches), mainly Nov–Jan (60%), 12% in Oct and Feb; Gambia, (fledglings, Oct); Mali, breeds ‘practically all year’ (Lamarche 1993); Burkina Faso, (nests being built, Jan); Ghana, Apr (and copulation Oct); Algeria, Hoggar, Aug; Niger, ‘W’ Nat. Park, Jan; Nigeria,

Padda oryzivora Sept–May; Chad, Oct–Apr, and in Waza. Nat. Park Aug– Nov (–Feb); Sudan, cantans, Aug–Mar (in Darfur, Oct– Mar), orientalis Dec; Eritrea, Oct, Feb–Mar (and nestlings Nov and Mar); Ethiopia, Apr; Somalia, in N, May, June (and nestlings Dec), in S, June; E Africa, Region D, Mar– Aug, mainly June (27% of 51 records) and July (51%). INCUBATION: by X; Y may relieve her when she leaves to forage, but may not actually brood; both parents in nest at night. Parents sometimes continue to bring feathers to nest during incubation and nestling periods, nest becoming very cosy. Period: 11–14 days. DEVELOPMENT AND CARE OF YOUNG: for several days after young hatch nest is kept clean, then ring of excreta

415

slowly builds up. Young fed by both parents, mainly during c. 2 h periods in morning and evening, but not for much of the intervening time (Mundy and Cook 1974). Chick hatches at 075 g, fledges at weight of ad. Period: 16– 18 (–21) days. BREEDING SUCCESS/SURVIVAL: nests in old weaver nests subject to ravages of African Harrier Hawk Polyboroides typus (Nigeria). Key References Baptista, L. and Horblit, H.M. (1990), Brickell, N. and Konigkramer, T. (1997), Clement, P. et al. (1993), Goodwin, D. (1982), Mundy, P.J. and Cook, A.W. (1974), Restall, R. (1996), van Someren, V.G.L. (1956).

Genus Padda Reichenbach

2 estrildid finches or ‘sparrows’, closely allied with Lonchura and often considered congeneric (e.g. by Goodwin 1982), but differing in having sharply-defined white cheek patches, nestling palate pattern, and shape of vomer. Further, one species P. oryzivora, is large and the only ‘Lonchura’ with red bill and pink in plumage. Biology like Lonchura. One species native to Java and Bali but kept as a cagebird worldwide and feral in many parts of Asia, the other restricted to Lesser Sunda Is.

Padda oryzivora (Linnaeus). Java Sparrow. Padda de Java.

Plate 24

Loxia oryzivora Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 173; Asia and Ethiopia [ ¼ Java].

(Opp. p. 347)

Range and Status. Native to Java and Bali; introduced in India, Sri Lanka, Thailand, Malaysia, Vietnam, E China, Taiwan, Philippines, Hawaii, Fiji, Mauritius, Mascarene Is, Comoro Is, Zanzibar and Pemba. Resident in and around larger towns on Zanzibar and Pemba Is, Tanzania. Introduced about 1957; but no further introductions in past 50 years, so population clearly well established (N.E. Baker, pers. comm.). Common in Zanzibar Town; large flocks appear in rural areas, e.g. Mkokotoni, at rice harvest time; >50 resident around Bwawani swamp. In Pemba, resident in Chake-Chake town. Description. ADULT Y: forehead to hindneck, lores and below eye glossy black; mantle, sides of neck, scapulars and back pale bluish grey; rump and uppertail-coverts black. Tail black. Cheeks and ear-coverts white, encircled by black cap, narrow black border behind and below, and narrow black line adjacent to lower mandible. Chin and upper throat black; lower throat and breast bluish grey, concolorous with neck and mantle; flanks, thighs and belly pinkish cinnamon; vent and undertail-coverts buffy white. Primaries, primary coverts and alula blackish brown, narrowly edged grey; secondaries, tertials, greater, median and lesser coverts bluish grey. Underwing-coverts and axillaries pale pinkish buff. Bill deep rose-pink with whitish tip and broad whitish line along commissure; eyes reddish brown to deep red; narrow eye-ring pale pink; legs pale fleshy pink. Sexes alike. SIZE (10 YY, 6 XX): wing, Y 69–75 (704), X 69–72 (702); tail, Y 42–51 (445), X 43– 46 (450); bill, Y 175–19 (182), X 17–18 (174); tarsus, Y 185– 195 (189), X 175–185 (178).

Padda oryzivora

IMMATURE: juv. dull buffy brown above, darker and greyer on top of head; sides of head, neck and underparts pale buff, whiter

Padda oryzivora Sept–May; Chad, Oct–Apr, and in Waza. Nat. Park Aug– Nov (–Feb); Sudan, cantans, Aug–Mar (in Darfur, Oct– Mar), orientalis Dec; Eritrea, Oct, Feb–Mar (and nestlings Nov and Mar); Ethiopia, Apr; Somalia, in N, May, June (and nestlings Dec), in S, June; E Africa, Region D, Mar– Aug, mainly June (27% of 51 records) and July (51%). INCUBATION: by X; Y may relieve her when she leaves to forage, but may not actually brood; both parents in nest at night. Parents sometimes continue to bring feathers to nest during incubation and nestling periods, nest becoming very cosy. Period: 11–14 days. DEVELOPMENT AND CARE OF YOUNG: for several days after young hatch nest is kept clean, then ring of excreta

415

slowly builds up. Young fed by both parents, mainly during c. 2 h periods in morning and evening, but not for much of the intervening time (Mundy and Cook 1974). Chick hatches at 075 g, fledges at weight of ad. Period: 16– 18 (–21) days. BREEDING SUCCESS/SURVIVAL: nests in old weaver nests subject to ravages of African Harrier Hawk Polyboroides typus (Nigeria). Key References Baptista, L. and Horblit, H.M. (1990), Brickell, N. and Konigkramer, T. (1997), Clement, P. et al. (1993), Goodwin, D. (1982), Mundy, P.J. and Cook, A.W. (1974), Restall, R. (1996), van Someren, V.G.L. (1956).

Genus Padda Reichenbach

2 estrildid finches or ‘sparrows’, closely allied with Lonchura and often considered congeneric (e.g. by Goodwin 1982), but differing in having sharply-defined white cheek patches, nestling palate pattern, and shape of vomer. Further, one species P. oryzivora, is large and the only ‘Lonchura’ with red bill and pink in plumage. Biology like Lonchura. One species native to Java and Bali but kept as a cagebird worldwide and feral in many parts of Asia, the other restricted to Lesser Sunda Is.

Padda oryzivora (Linnaeus). Java Sparrow. Padda de Java.

Plate 24

Loxia oryzivora Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 173; Asia and Ethiopia [ ¼ Java].

(Opp. p. 347)

Range and Status. Native to Java and Bali; introduced in India, Sri Lanka, Thailand, Malaysia, Vietnam, E China, Taiwan, Philippines, Hawaii, Fiji, Mauritius, Mascarene Is, Comoro Is, Zanzibar and Pemba. Resident in and around larger towns on Zanzibar and Pemba Is, Tanzania. Introduced about 1957; but no further introductions in past 50 years, so population clearly well established (N.E. Baker, pers. comm.). Common in Zanzibar Town; large flocks appear in rural areas, e.g. Mkokotoni, at rice harvest time; >50 resident around Bwawani swamp. In Pemba, resident in Chake-Chake town. Description. ADULT Y: forehead to hindneck, lores and below eye glossy black; mantle, sides of neck, scapulars and back pale bluish grey; rump and uppertail-coverts black. Tail black. Cheeks and ear-coverts white, encircled by black cap, narrow black border behind and below, and narrow black line adjacent to lower mandible. Chin and upper throat black; lower throat and breast bluish grey, concolorous with neck and mantle; flanks, thighs and belly pinkish cinnamon; vent and undertail-coverts buffy white. Primaries, primary coverts and alula blackish brown, narrowly edged grey; secondaries, tertials, greater, median and lesser coverts bluish grey. Underwing-coverts and axillaries pale pinkish buff. Bill deep rose-pink with whitish tip and broad whitish line along commissure; eyes reddish brown to deep red; narrow eye-ring pale pink; legs pale fleshy pink. Sexes alike. SIZE (10 YY, 6 XX): wing, Y 69–75 (704), X 69–72 (702); tail, Y 42–51 (445), X 43– 46 (450); bill, Y 175–19 (182), X 17–18 (174); tarsus, Y 185– 195 (189), X 175–185 (178).

Padda oryzivora

IMMATURE: juv. dull buffy brown above, darker and greyer on top of head; sides of head, neck and underparts pale buff, whiter

Padda oryzivora Sept–May; Chad, Oct–Apr, and in Waza. Nat. Park Aug– Nov (–Feb); Sudan, cantans, Aug–Mar (in Darfur, Oct– Mar), orientalis Dec; Eritrea, Oct, Feb–Mar (and nestlings Nov and Mar); Ethiopia, Apr; Somalia, in N, May, June (and nestlings Dec), in S, June; E Africa, Region D, Mar– Aug, mainly June (27% of 51 records) and July (51%). INCUBATION: by X; Y may relieve her when she leaves to forage, but may not actually brood; both parents in nest at night. Parents sometimes continue to bring feathers to nest during incubation and nestling periods, nest becoming very cosy. Period: 11–14 days. DEVELOPMENT AND CARE OF YOUNG: for several days after young hatch nest is kept clean, then ring of excreta

415

slowly builds up. Young fed by both parents, mainly during c. 2 h periods in morning and evening, but not for much of the intervening time (Mundy and Cook 1974). Chick hatches at 075 g, fledges at weight of ad. Period: 16– 18 (–21) days. BREEDING SUCCESS/SURVIVAL: nests in old weaver nests subject to ravages of African Harrier Hawk Polyboroides typus (Nigeria). Key References Baptista, L. and Horblit, H.M. (1990), Brickell, N. and Konigkramer, T. (1997), Clement, P. et al. (1993), Goodwin, D. (1982), Mundy, P.J. and Cook, A.W. (1974), Restall, R. (1996), van Someren, V.G.L. (1956).

Genus Padda Reichenbach

2 estrildid finches or ‘sparrows’, closely allied with Lonchura and often considered congeneric (e.g. by Goodwin 1982), but differing in having sharply-defined white cheek patches, nestling palate pattern, and shape of vomer. Further, one species P. oryzivora, is large and the only ‘Lonchura’ with red bill and pink in plumage. Biology like Lonchura. One species native to Java and Bali but kept as a cagebird worldwide and feral in many parts of Asia, the other restricted to Lesser Sunda Is.

Padda oryzivora (Linnaeus). Java Sparrow. Padda de Java.

Plate 24

Loxia oryzivora Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 173; Asia and Ethiopia [ ¼ Java].

(Opp. p. 347)

Range and Status. Native to Java and Bali; introduced in India, Sri Lanka, Thailand, Malaysia, Vietnam, E China, Taiwan, Philippines, Hawaii, Fiji, Mauritius, Mascarene Is, Comoro Is, Zanzibar and Pemba. Resident in and around larger towns on Zanzibar and Pemba Is, Tanzania. Introduced about 1957; but no further introductions in past 50 years, so population clearly well established (N.E. Baker, pers. comm.). Common in Zanzibar Town; large flocks appear in rural areas, e.g. Mkokotoni, at rice harvest time; >50 resident around Bwawani swamp. In Pemba, resident in Chake-Chake town. Description. ADULT Y: forehead to hindneck, lores and below eye glossy black; mantle, sides of neck, scapulars and back pale bluish grey; rump and uppertail-coverts black. Tail black. Cheeks and ear-coverts white, encircled by black cap, narrow black border behind and below, and narrow black line adjacent to lower mandible. Chin and upper throat black; lower throat and breast bluish grey, concolorous with neck and mantle; flanks, thighs and belly pinkish cinnamon; vent and undertail-coverts buffy white. Primaries, primary coverts and alula blackish brown, narrowly edged grey; secondaries, tertials, greater, median and lesser coverts bluish grey. Underwing-coverts and axillaries pale pinkish buff. Bill deep rose-pink with whitish tip and broad whitish line along commissure; eyes reddish brown to deep red; narrow eye-ring pale pink; legs pale fleshy pink. Sexes alike. SIZE (10 YY, 6 XX): wing, Y 69–75 (704), X 69–72 (702); tail, Y 42–51 (445), X 43– 46 (450); bill, Y 175–19 (182), X 17–18 (174); tarsus, Y 185– 195 (189), X 175–185 (178).

Padda oryzivora

IMMATURE: juv. dull buffy brown above, darker and greyer on top of head; sides of head, neck and underparts pale buff, whiter

416

VIDUIDAE

on chin, washed greyish on breast, more tawny on flanks and belly. Upperwing dark brown, tertials and coverts tipped buffy brown; tail blackish brown. Bill blackish at first with pale pink base; legs dark flesh. NESTLING: pinkish with dusky down; bill blackish with creamy white gape and gape tubercles; mouth pink with crescentic black mark on palate.

Field Characters. Length 15–16 cm. A plump grey finch with prominent white cheek patch and stout pink bill; larger than waxbills, size of a House Sparrow Passer domesticus. Confined to Zanzibar and Pemba, where cannot be confused with any other bird. Immature drab, with dark bill. Voice. Not tape-recorded. Clicking and chirruping contact calls, softer and more liquid than House Sparrow, often given by flock in flight. Song starts with a few bell-like notes, accelerates into clucking trill and ends with whistling ‘weee’; for further variations see Goodwin (1982). General Habits. Inhabits suburbs, gardens, markets and similar places in town centres, small-holdings, cultivated

land, paddies, sugarcane and maize fields, grassland, open woodland with grass and scrub, and mangroves. Occurs in pairs when breeding; otherwise strongly gregarious. Forages in large flocks, on ground amongst young cereals and herbs; perches freely in shrubs, trees and palms; roosts in large flocks in dense vegetation. Food. Seeds, mainly of grasses and cereals (rice, millet, maize); some small fruits and insects. Breeding Habits. Not studied in Africa. Displaying Y adopts peculiar, bent-over posture and jumps up and down on perch (Goodwin 1982). In Java, nests in cavities under roofs of buildings or high up in trees and palms; nest roughly globular, made of grass bents. In captivity, nest often not roofed (depending on shape of nest cavity); lined with coconut fibres and fine grass, or with feathers; some nests very slight. 3–8 (usually 4–6) white eggs. LAYING DATES: Pemba, (nest-building Apr). Key Reference Goodwin, D. (1982).

Family VIDUIDAE: whydahs, indigobirds, Cuckoo-Finch

Obligate brood parasites. Bill short, plumage sexually dimorphic; Y whydahs and indigobirds, Vidua, have breeding plumage black, often glossy, often with white or buff below, X streaked brown; in non-breeding season YY moult into Xlike plumage; Cuckoo-Finch Anomalospiza imberbis Y acquires bright plumage by wear rather than moult; X streaked brown. Outer primary short and narrow. Adult skull with unpneumatized frontal areas (10–30% of dorsal area persisting unpneumatized for at least 3 years in Vidua: Chapin 1954, Payne 1973a). P10 vestigial. Juvenile plumage distinct from ad. plumage. Anomalospiza and Vidua share several feather tract characteristics (Vidua: Morlion 1971, 1980): 2 rows of upper greater secondary coverts, 8 upper median secondary coverts, 4 upper tertiary coverts, 9 under greater primary coverts, 1 row of ocular feathers, 8 longitudinal rows of feathers on crown, and 3 rictal vibrissae. (Some of these characters also occur in Estrildidae.) Anomalospiza feather tract characters are quite unlike those of ploceids. Viduidae are most closely related to estrildid finches Estrildidae, several of which are brood hosts of whydahs and indigobirds (Sibley and Ahlquist 1990, Groth 1998). Both Vidua and estrildid nestlings and fledglings are fed as they crouch, twist their heads around and upside down when begging, hold mouth open and wave head from side to side, behaviours that display the open mouth and palate. Those parts in Vidua are coloured and marked much as in estrildid host nestlings. Foster parent inserts its bill into mouth of young and regurgitates into the crop (Nicolai 1964, 1974, Kunkel 1969; nestlings of euplectines and canaries stretch neck up, beg without twisting head; parent drops food into open mouth). A. imberbis are fed by cisticoline warblers and the young beg like host young, not like Vidua young. Viduidae differ from ad. estrildids in lack of nestbuilding and parental behaviour, in lack of social allopreening or perching in contact with each other, and (most species) in presence of distinct breeding and non-breeding plumages. Horny palate is unlike palate of most ploceids (in which lateral ridges extend behind and do not converge or converge at hind edge: Sushkin 1927); lateral ridges converge just past halfway back, obscure the medial ridge (absent in Anomalospiza) and are displaced inward by pair of large fossae on posterior surface of palate. Anomalospiza and Vidua share the horny palate conformation of raised surface anteriorly, medially displaced lateral ridges, and bilateral fossae. Molecular genetics indicate that Anomalospiza and Vidua are each others’ closest relatives (Payne 1997a, Sorenson and Payne 1998, 2001, 2002). Breeding biology information includes ‘clutch’ or ‘set’ of eggs. X lays in one nest or more than one, not all in one nest (as ‘clutch’ implies). A ‘set’ for a brood-parasite is the number of eggs that develop in series in the ovary and are laid on successive days: sets are determined by inspection of ovary (post-ovulatory follicles and growing follicles: Payne 1977a). Endemic; 20 species in 2 genera.

416

VIDUIDAE

on chin, washed greyish on breast, more tawny on flanks and belly. Upperwing dark brown, tertials and coverts tipped buffy brown; tail blackish brown. Bill blackish at first with pale pink base; legs dark flesh. NESTLING: pinkish with dusky down; bill blackish with creamy white gape and gape tubercles; mouth pink with crescentic black mark on palate.

Field Characters. Length 15–16 cm. A plump grey finch with prominent white cheek patch and stout pink bill; larger than waxbills, size of a House Sparrow Passer domesticus. Confined to Zanzibar and Pemba, where cannot be confused with any other bird. Immature drab, with dark bill. Voice. Not tape-recorded. Clicking and chirruping contact calls, softer and more liquid than House Sparrow, often given by flock in flight. Song starts with a few bell-like notes, accelerates into clucking trill and ends with whistling ‘weee’; for further variations see Goodwin (1982). General Habits. Inhabits suburbs, gardens, markets and similar places in town centres, small-holdings, cultivated

land, paddies, sugarcane and maize fields, grassland, open woodland with grass and scrub, and mangroves. Occurs in pairs when breeding; otherwise strongly gregarious. Forages in large flocks, on ground amongst young cereals and herbs; perches freely in shrubs, trees and palms; roosts in large flocks in dense vegetation. Food. Seeds, mainly of grasses and cereals (rice, millet, maize); some small fruits and insects. Breeding Habits. Not studied in Africa. Displaying Y adopts peculiar, bent-over posture and jumps up and down on perch (Goodwin 1982). In Java, nests in cavities under roofs of buildings or high up in trees and palms; nest roughly globular, made of grass bents. In captivity, nest often not roofed (depending on shape of nest cavity); lined with coconut fibres and fine grass, or with feathers; some nests very slight. 3–8 (usually 4–6) white eggs. LAYING DATES: Pemba, (nest-building Apr). Key Reference Goodwin, D. (1982).

Family VIDUIDAE: whydahs, indigobirds, Cuckoo-Finch

Obligate brood parasites. Bill short, plumage sexually dimorphic; Y whydahs and indigobirds, Vidua, have breeding plumage black, often glossy, often with white or buff below, X streaked brown; in non-breeding season YY moult into Xlike plumage; Cuckoo-Finch Anomalospiza imberbis Y acquires bright plumage by wear rather than moult; X streaked brown. Outer primary short and narrow. Adult skull with unpneumatized frontal areas (10–30% of dorsal area persisting unpneumatized for at least 3 years in Vidua: Chapin 1954, Payne 1973a). P10 vestigial. Juvenile plumage distinct from ad. plumage. Anomalospiza and Vidua share several feather tract characteristics (Vidua: Morlion 1971, 1980): 2 rows of upper greater secondary coverts, 8 upper median secondary coverts, 4 upper tertiary coverts, 9 under greater primary coverts, 1 row of ocular feathers, 8 longitudinal rows of feathers on crown, and 3 rictal vibrissae. (Some of these characters also occur in Estrildidae.) Anomalospiza feather tract characters are quite unlike those of ploceids. Viduidae are most closely related to estrildid finches Estrildidae, several of which are brood hosts of whydahs and indigobirds (Sibley and Ahlquist 1990, Groth 1998). Both Vidua and estrildid nestlings and fledglings are fed as they crouch, twist their heads around and upside down when begging, hold mouth open and wave head from side to side, behaviours that display the open mouth and palate. Those parts in Vidua are coloured and marked much as in estrildid host nestlings. Foster parent inserts its bill into mouth of young and regurgitates into the crop (Nicolai 1964, 1974, Kunkel 1969; nestlings of euplectines and canaries stretch neck up, beg without twisting head; parent drops food into open mouth). A. imberbis are fed by cisticoline warblers and the young beg like host young, not like Vidua young. Viduidae differ from ad. estrildids in lack of nestbuilding and parental behaviour, in lack of social allopreening or perching in contact with each other, and (most species) in presence of distinct breeding and non-breeding plumages. Horny palate is unlike palate of most ploceids (in which lateral ridges extend behind and do not converge or converge at hind edge: Sushkin 1927); lateral ridges converge just past halfway back, obscure the medial ridge (absent in Anomalospiza) and are displaced inward by pair of large fossae on posterior surface of palate. Anomalospiza and Vidua share the horny palate conformation of raised surface anteriorly, medially displaced lateral ridges, and bilateral fossae. Molecular genetics indicate that Anomalospiza and Vidua are each others’ closest relatives (Payne 1997a, Sorenson and Payne 1998, 2001, 2002). Breeding biology information includes ‘clutch’ or ‘set’ of eggs. X lays in one nest or more than one, not all in one nest (as ‘clutch’ implies). A ‘set’ for a brood-parasite is the number of eggs that develop in series in the ovary and are laid on successive days: sets are determined by inspection of ovary (post-ovulatory follicles and growing follicles: Payne 1977a). Endemic; 20 species in 2 genera.

Anomalospiza imberbis

417

Genus Anomalospiza Shelley

Single species of endemic, thick-billed weaver-like finch, brood parasitic on Prinia and Cisticola warblers of open grasslands. Bill short, deep, robust, powerful, laterally compressed, culmen ridged and nearly straight, cutting edge of upper mandible bent at sharp angle near base; appears to be effective in cracking and crushing seeds. Wing rounded, P10 slender and same length as coverts, P9 and P8 longest and subequal in length, P7 and P6 next longest. Tail short and slightly rounded, 12 rectrices. Sternum with forked spina interna, with base formed so that internal corner of each coracoid fits into a small socket (rather than in a groove), as in buffalo-weavers Bubalornis and Dinemellia (though much smaller in Anomalospiza) but not in other thick-billed weavers such as Amblyospiza. Bill and palate distinctive: jaw bent down at frontal-nasal-maxillary hinge at angle of c. 110
relative to jugal. Jaw has laterally compressed expansion and ventral protuberance of jugal bone. Inside upper jaw thickened, reducing buccal volume to size of a large seed within the palate groove. Lower mandible has a thick, flat rostral flange, expanded ventrally at its posterior margin. Caudal angle of palatine narrow, relatively smaller than in the larger Vidua species. Rostral pes of pterygoid expanded. Palate with medial groove; sides of mandibles dilated inward to form 2 horny pads, leaving only a narrow medial groove into which the small tongue or a seed can fit. On either side of palate near gape is a hard circular hollow in which the horny mandible pads fit when bill is closed. The horny, bony palate is adapted to break or crush hard seeds. Relationships to other finches uncertain in the past. Originally described in 1868 as a canary, it was considered a canary by Reichenow (1904). It was next considered a weaver Ploceidae on the basis of wing shape (with short outer primary like Euplectes and Quelea) and its sexually dimorphic plumage (Shelley 1905, Delacour 1943); it was ‘clearly not closely related to the Viduinae’ (Friedmann 1960). Sibley and Monroe (1990) questioned a ploceid relationship and Chapin (1917) and Bannerman (1949) thought it closely related to Vidua. In fact, morphology and molecular genetics now indicate a close relationship with Vidua (Sorenson and Payne 1998, 2001, 2002, Lahti and Payne 2003).

Anomalospiza imberbis (Cabanis). Cuckoo-Finch; Parasitic Weaver. Tisserin-coucou.

Plate 25

Crithagra imberbis Cabanis, 1868. J. Orn., 16, p. 412; Zanzibar.

(Opp. p. 394)

Range and Status. Endemic, resident and wanderer. Generally uncommon or scarce, especially in W Africa; locally common in E and S with seasonal movements. Gambia (Banjul). Mali (Kati, Se´gou, Niger Delta). Guinea (Koundara, Ayama plains, Mt Nimba). Sierra Leone (Bo, Sefadu, Bruman). Liberia, local and rare as mapped (Gatter 1997). Ivory Coast (Mt Tonkoui, Maroue´ Nat. Park). Ghana (in N at Sugu, in S at Abokobi on Accra Plains). Togo (Namoundjoga, Kara, Te´te´tou). Benin, 1, CotonouLome´ road (Claffey 2003). Nigeria (Zaria, Jos Plateau, Gongola R.). Cameroon (Waza, Poli, Ngambe´, Tibati, Foumban-Tibati). S Sudan (floodplains south of 6
N and Ethiopian border at 9
N). N Zaı¨re (Uele). Ethiopia (L. Tana and Dangila in W highlands, L. Zwai, and south of Addis Ababa). In E Africa, local and uncommon below 2000 m, locally resident and a seasonal wanderer breeding after rains. Uganda (Murchison Falls Nat. Park to Soroti, Jinja, Moroto). Kenya (upper Turkwell R. area, near Mt Elgon, L. Victoria basin in Siaya and S Nyanza, Mara, Laikipia Plateau, Nairobi and Thika; Voi and lower Tana R. and along coast near Mombasa). Tanzania (Serengeti Nat. Park, Arusha, Dodoma, Iringa region and Ufipa Plateau; Zanzibar, Pemba). Zaı¨re (lower Congo near Kinshasa, Uele, Kasai, Katanga and E Kivu). Angola (L. Carumbo in Lunda Norte, Chimporo in S Huı´la, L. Dilolo in Moxico). Zambia. Zimbabwe (grasslands with scattered bushes on central plateau above 900 m, most records near Harare and north of 18
S, occasional in upper Zambezi

Anomalospiza imberbis

grasslands; in Hwange Nat. Park large flocks in dry season). Namibia (Caprivi strip). Botswana (upper Okavango and

Anomalospiza imberbis

417

Genus Anomalospiza Shelley

Single species of endemic, thick-billed weaver-like finch, brood parasitic on Prinia and Cisticola warblers of open grasslands. Bill short, deep, robust, powerful, laterally compressed, culmen ridged and nearly straight, cutting edge of upper mandible bent at sharp angle near base; appears to be effective in cracking and crushing seeds. Wing rounded, P10 slender and same length as coverts, P9 and P8 longest and subequal in length, P7 and P6 next longest. Tail short and slightly rounded, 12 rectrices. Sternum with forked spina interna, with base formed so that internal corner of each coracoid fits into a small socket (rather than in a groove), as in buffalo-weavers Bubalornis and Dinemellia (though much smaller in Anomalospiza) but not in other thick-billed weavers such as Amblyospiza. Bill and palate distinctive: jaw bent down at frontal-nasal-maxillary hinge at angle of c. 110
relative to jugal. Jaw has laterally compressed expansion and ventral protuberance of jugal bone. Inside upper jaw thickened, reducing buccal volume to size of a large seed within the palate groove. Lower mandible has a thick, flat rostral flange, expanded ventrally at its posterior margin. Caudal angle of palatine narrow, relatively smaller than in the larger Vidua species. Rostral pes of pterygoid expanded. Palate with medial groove; sides of mandibles dilated inward to form 2 horny pads, leaving only a narrow medial groove into which the small tongue or a seed can fit. On either side of palate near gape is a hard circular hollow in which the horny mandible pads fit when bill is closed. The horny, bony palate is adapted to break or crush hard seeds. Relationships to other finches uncertain in the past. Originally described in 1868 as a canary, it was considered a canary by Reichenow (1904). It was next considered a weaver Ploceidae on the basis of wing shape (with short outer primary like Euplectes and Quelea) and its sexually dimorphic plumage (Shelley 1905, Delacour 1943); it was ‘clearly not closely related to the Viduinae’ (Friedmann 1960). Sibley and Monroe (1990) questioned a ploceid relationship and Chapin (1917) and Bannerman (1949) thought it closely related to Vidua. In fact, morphology and molecular genetics now indicate a close relationship with Vidua (Sorenson and Payne 1998, 2001, 2002, Lahti and Payne 2003).

Anomalospiza imberbis (Cabanis). Cuckoo-Finch; Parasitic Weaver. Tisserin-coucou.

Plate 25

Crithagra imberbis Cabanis, 1868. J. Orn., 16, p. 412; Zanzibar.

(Opp. p. 394)

Range and Status. Endemic, resident and wanderer. Generally uncommon or scarce, especially in W Africa; locally common in E and S with seasonal movements. Gambia (Banjul). Mali (Kati, Se´gou, Niger Delta). Guinea (Koundara, Ayama plains, Mt Nimba). Sierra Leone (Bo, Sefadu, Bruman). Liberia, local and rare as mapped (Gatter 1997). Ivory Coast (Mt Tonkoui, Maroue´ Nat. Park). Ghana (in N at Sugu, in S at Abokobi on Accra Plains). Togo (Namoundjoga, Kara, Te´te´tou). Benin, 1, CotonouLome´ road (Claffey 2003). Nigeria (Zaria, Jos Plateau, Gongola R.). Cameroon (Waza, Poli, Ngambe´, Tibati, Foumban-Tibati). S Sudan (floodplains south of 6
N and Ethiopian border at 9
N). N Zaı¨re (Uele). Ethiopia (L. Tana and Dangila in W highlands, L. Zwai, and south of Addis Ababa). In E Africa, local and uncommon below 2000 m, locally resident and a seasonal wanderer breeding after rains. Uganda (Murchison Falls Nat. Park to Soroti, Jinja, Moroto). Kenya (upper Turkwell R. area, near Mt Elgon, L. Victoria basin in Siaya and S Nyanza, Mara, Laikipia Plateau, Nairobi and Thika; Voi and lower Tana R. and along coast near Mombasa). Tanzania (Serengeti Nat. Park, Arusha, Dodoma, Iringa region and Ufipa Plateau; Zanzibar, Pemba). Zaı¨re (lower Congo near Kinshasa, Uele, Kasai, Katanga and E Kivu). Angola (L. Carumbo in Lunda Norte, Chimporo in S Huı´la, L. Dilolo in Moxico). Zambia. Zimbabwe (grasslands with scattered bushes on central plateau above 900 m, most records near Harare and north of 18
S, occasional in upper Zambezi

Anomalospiza imberbis

grasslands; in Hwange Nat. Park large flocks in dry season). Namibia (Caprivi strip). Botswana (upper Okavango and

418

VIDUIDAE

ˆ i (mainly at 500–1800 m, once in Chobe grasslands). Malaw Shire lowlands). Mozambique (Tete, Lower Zambezi and Beira areas in coastal and wet grasslands in N, once in S at Boane). South Africa (Transvaal and cereal lands of KwaZulu-Natal). Description. A. i. imberbis (Cabanis): Ethiopia, E Africa, S Zaı¨re (Kasai, Katanga) and S-central Africa to South Africa. ADULT Y (fresh plumage): forehead and face yellowish, back olive greenish, streaked, wings and tail blackish with outer edges of primaries and both margins of tail feathers buff to yellow, central tail feathers narrow and pointed; below dull yellow from chin to undertail-coverts, breast in some birds with grey wash; underwingcoverts yellow; in worn plumage, upperparts brighter, paler, more distinctly streaked black; below bright yellow. Seasonal change in plumage occurs without moult and is due to wear, which on nape and back removes dark grey feather margins and reveals black feather shafts and yellow vanes, creating black streaks and yellow vanes for breeding season. Feathers become pointed with wear. Bill black, lower mandible with pale base (breeding season) or uniform light brown (non-breeding season); eyes blackish brown; feet pinkish grey, pale horn grey, light brown or slate-grey. ADULT X: crown and back buffy brown broadly streaked blackish, pale crown-stripe and superciliary stripe, wing brown, greater primary coverts dark brown, tail dark brown edged buffy, central rectrices pointed as in Y; face dull yellow to whitish with cheeks brown; below buffy white with variable streaking on breast and flanks (throat whitish, buffier breast in fresh plumage, dark streaks more evident in worn plumage); underwing-coverts and inner webs of primaries buff, undertail-coverts buff. Bill dark brown on culmen merging to pale pinkish horn on cutting edge of both mandibles; eyes blackish brown; feet pale greyish horn to horn. SIZE: wing, Y (n ¼ 56) 64–74 (696), X (n ¼ 30) 63–71 (670); tail, Y (n ¼ 56) 35–47 (387), X (n ¼ 31) 33–41 (375); bill, Y (n ¼ 53) 113–143 (123), X (n ¼ 29) 101–130 (119); tarsus, Y (n ¼ 52) 161–193 (173), X (n ¼ 28) 162–192 (175). Ethiopian birds are larger; wing, Y (n ¼ 8) 71–73, X (n ¼ 3) 66–69; Kasai birds intermediate in size between W African and E/S African birds (Kasai, wing, Y (n ¼ 25) 62–70). WEIGHT: Kenya, Y (n ¼ 8) 18–21 (198), X (n ¼ 6) 19–21 (196) (Williams and Keith 1962); Tanzania Y (n ¼ 5) 19–21 (198), 2 XX 21, 21; Transvaal, Y (n ¼ 4) 206–226 (215), X (laying) (n ¼ 6) 200–226 (214) (Payne 1977a). IMMATURE: juv. above tawny brown, centres of feathers blackish brown, edges tawny buff, face, throat and breast golden sandy buff, throat and belly pale buff, flanks buff with fine black streaks, greater primary coverts blackish with buff edges. Resembles juv. of hosts such as Winding Cisticola Cisticola galactotes. Juv. with rows of blackish scale-like tips to tawny crown feathers; upper mandible black, lower mandible pale, light brown; bill thick; eyes blackish brown; feet grey-brown. Juv. Y slightly yellower than juv. X from same nest. Plumage becomes darker with wear. Central tail feathers less pointed than in ads. Juv. Y moults directly into yellow-breasted plumage of ad. Y. NESTLING: at hatching differs from host young, skin dark purple on upperparts, dark flesh colour on underparts, tufts of fine white natal down on crown, back and wings. Mouth has broad border of deep orange-yellow, no palate or mouth spots, lining unmarked flesh or flesh-purple, tongue purplish pink with large black spots near rear, interior of both mandibles bright yellow. Bill laterally compressed in fledgling, but deep, like ad.; above, black and below yellow to straw-colour with black spots on gonys and near tip, becomes yellowish with dark brown culmen; eyes grey brown. A. i. butleri Sclater and Mackworth-Praed: W Africa to S Sudan (where some birds as large as E African birds) and N Zaı¨re (Uele). Small; bill horn-brown to brown with base of lower mandible whitish, eyes dark brown, feet horn brown. SIZE: wing, Y (n ¼ 29) 62–68 (656), X (n ¼ 7) 61–65 (637); tail, Y (n ¼ 25) 34–46 (364),

X (n ¼ 7) 34–41 (377); bill, Y (n ¼ 26) 97–124 (116), X (n ¼ 7) 103–122 (113); tarsus, Y (n ¼ 26) 142–171 (163), X (n ¼ 7) 152–167 (161). WEIGHT: Mali and Nigeria, Y (n ¼ 2) 173, 180.

Field Characters. Length 11–12 cm. Small and stocky bird with short tail and short, stubby bill. Culmen straight (slightly curved in bishops, weavers and canaries), length nearly same as depth at nostril (in bishops Euplectes half again as long). Breeding Y with yellow head and underparts resembles a short-tailed canary, but has robust bill, lower mandible dark with pale base, and black eye high on face. X streaked black on brown above, face yellowish white, cheeks buffy brown; resembles X Southern Red Bishop E. orix but with shorter and thicker bill, and tail extends well past wing tips at rest. Culmen straight; cutting edge of lower mandible angled at base about 60
from distal cutting edge (about 20
in bishops). Long claw on hallux visible at close range. Juvenile with crown streaked tawny and black, underparts tawny. Voice. Tape-recorded (99, 104, B, GIB, PAY). Song lasts 4 s and is in 3 parts, (1) series of chirping notes dropping in pitch from 8 to 2 kHz, lasting 10–20 s; (2) soft, descending whistle, ‘choo’, lasting 16 s, and (3) a final chirp. Some songs include only the chirping series. First part of song described as a thin, sibilant ‘tissiwick’ and ‘tissiway’, sometimes followed by a little jingle, ‘djzing-ji-ji’. Calls include a deliberate, 3-syllabled ‘dzi-bee-chew’ or ‘choopee choo’, and in flight call, a hard ‘jit-jit’ or ‘cheet-cheet’. Distress call a buzzy chatter. General Habits. Inhabits open, bushy grasslands, damp open meadows with high grasses, floodplains, dams, and edges of cultivation, rice fields, and weedy old cotton fields; around airfields during rains. Occurs singly or in small or large flocks in all seasons. In non-breeding season flocks of 8–50 birds, occasionally up to 200 (and over 1000 in Hwange Nat. Park, Zimbabwe). Feeds on ground, takes fallen grass seeds, moves awkwardly in upright posture and pecks seeds from surface. Also feeds on stems, clinging to erect stalk of grass or herb; perches on sunflower stalk, pulls seeds from fruiting heads, and crushes them in bill (Payne 1997a). Also takes sand and grit that pulverize hard seeds in stomach. Roosts in flocks in reedbeds. Flight direct and rapid, not undulating. Resident or moves locally with rains and grass seeding in most of range. Migratory in Zimbabwe, where it occurs in wet season (Irwin 1952), and in Natal and Transvaal, where most records Oct–May (Harrison et al. 1997). Food. Grass seeds (Setaria sphacelata, Sorghum verticilliflorum), other soft seeds, and hard seeds (sedges Scirpus, sunflowers Helianthus); also insects. X eats host’s eggs (shell found in crop, yolk seen on bill of laying X). Breeding Habits. Y sings from exposed perches on bushes and grass, or on ground in rainy season; YY, spaced on separate display territories, give territorial displays and chase each other. YY and laying XX also flock in breeding season.

Vidua

In courtship display, Y perches on twig or ground, faces X, arches wings high over back and slowly flicks, fans and flaps them; he stretches neck upward and sometimes arches it so that bill points to ground (Williams and Keith 1962). Y also flicks wings when display is not directed to X and when X is foraging nearby. X flicks wings over back. Copulation not described. Brood-parasitic, with host species attending and feeding young (Roberts 1917, 1939, van Someren 1922) (A). Lays in nests of warblers Prinia spp. and Cisticola spp., including those with ball nest with entrance hole at side, ‘soda bottle’ nest with vertically elongated opening at top, and ‘tailorbird’ nest attached or sewn to underside of large leaves. Known host species, mainly in grassland habitats, are Prinia subflava, P. flavicans, Cisticola chiniana, C. natalensis, C. cantans, C. tinniens, C. juncidis, C. aridula, C. ayresii, C. brunnescens, C. textrix, and C. galactotes. Host flies at X near nest, chases and gives alarm calls. X removes 1 or all host eggs before she lays her own (host eggs appearing in parasitized nests are laid later), but sometimes fails to remove eggs of largest hosts. Nestling apparently interferes with survival of host nestlings, though nest of C. galactotes had 1 host young and 2 Cuckoo-Finches all ready to fly (the latter misidentified as ‘Vidua macroura’: Cheesman and Sclater 1935–1936). Foster parents Cisticola juncidis feed insects, mainly caterpillars and grasshoppers, also snails, to fledglings. EGGS: sets (‘clutches’) of 1–4 (mean, 29), 1 a day, a few days between sets. A parasitized nest has 1–2 Anomalospiza eggs; a X lays c. 30 eggs in a season. Little gloss; colour and markings vary, often match hosts’ eggs (bluish white with no spots, or with speckles and spots; whitish with red spots; pinkish with faint purplish at thick end and purplish and reddish brown spots and blotches) (Payne

1944, Neuby-Varty 1950, Vincent 1949, Vernon 1964). SIZE: 17  13; weight 16 g; larger than all but the largest host species, but small in relation to X body size. LAYING DATES: breeds with rains, along with its host species. W Africa, no breeding records. Ethiopia (W Highlands, Aug, Nov; E Africa, north and east of L. Victoria, Feb–June; Kenya, Jan, May, June, Oct; Pemba Island, Sept–Jan; Zaı¨re (Kivu, suspected May–June; Uele, ˆ i, Apr; Botswana, Aug–Sept); Zambia, Jan–Apr; Malaw Feb and Apr; Zimbabwe, Nov–Mar, mainly Dec–Feb; Transvaal, Sept–Mar. INCUBATION: period undetermined, less than 14 days. DEVELOPMENT AND CARE OF YOUNG: nestling period 18 days, longer than young of host. Parental care is by hosts, which feed entire insects directly to nestlings. Host and parasite chicks rarely fledge together. Failure of host nestlings is due to removal of host eggs by X Cuckoo-Finch and by competition or interference by nestling parasites, not by eviction by nestlings insofar as 2 Cuckoo-Finch young often fledge from same nest. By day 2 after hatching, 2 chicks trampled a Cisticola ayresii chick, which disappeared 2 days later. By day 4, nestling is nearly black above, mouth flesh-purple with deep orange-yellow border formed a distinct gape flange; bill horn above and yellow below with a terminal black spot. By day 7, bill heavier and darker, tufts of natal down remain on crown, back and wings, featherquills broken out over upperparts; spine in distinct tracts on head and middle of back. By day 9, eyes still closed, feather tips emerge on back, and small orange-brown feather quills appear on throat. By day 11, back feathers emerge with sandy brown colour. By day 13, 2 chicks nearly fill nest, face feathered and legs pale grey. By day 18, young leave nest and are fed by host pair as far as 30 m from nest, and can fly low but strongly for 50 m (Vernon 1964). Begging posture is upright, without twists and turns of head and neck; begging calls are ‘tink’ or ‘jit . . . jit’, like calls of an ad. (and unlike begging calls of host young: Vernon 1987, Herremans et al. 1993, Brewster 2000). Fledgling remains with host for 10 days, when it begins to feed itself, clinging on grass stems to reach seed-heads while still accompanied by foster parents, then leaves the foster family and joins flocks of conspecifics (Chapin 1954). Key References Chapin, J.P. (1954), Friedmann, H. (1960), Lahti, D. M. and Payne, R. B. (2003), Roberts, A. (1917, 1939), Vernon, C.J. (1964, 1987), Williams, J.G. and Keith, G.S. (1962).

Genus Vidua Cuvier

Brood-parasitic finches with sexually dimorphic breeding plumage, YY glossy black, short-tailed (10 indigobirds), or dull black and elaborate-tailed (8 whydahs), or glossy black and elaborate-tailed (V. hypocherina). XX and non-breeding YY streaked brown above and whitish below; juvs less streaked, often resemble juvs of their host species. Wing rounded, P10 smaller than coverts and pointed, P9–P6 longest and subequal in length, axillaries and underwing-coverts usually whitish or white. Tail with 12 feathers, inner 2 pairs (T1, T2) moult twice a year in Y whydahs and indigobirds; other tail-feathers

419

Vidua

In courtship display, Y perches on twig or ground, faces X, arches wings high over back and slowly flicks, fans and flaps them; he stretches neck upward and sometimes arches it so that bill points to ground (Williams and Keith 1962). Y also flicks wings when display is not directed to X and when X is foraging nearby. X flicks wings over back. Copulation not described. Brood-parasitic, with host species attending and feeding young (Roberts 1917, 1939, van Someren 1922) (A). Lays in nests of warblers Prinia spp. and Cisticola spp., including those with ball nest with entrance hole at side, ‘soda bottle’ nest with vertically elongated opening at top, and ‘tailorbird’ nest attached or sewn to underside of large leaves. Known host species, mainly in grassland habitats, are Prinia subflava, P. flavicans, Cisticola chiniana, C. natalensis, C. cantans, C. tinniens, C. juncidis, C. aridula, C. ayresii, C. brunnescens, C. textrix, and C. galactotes. Host flies at X near nest, chases and gives alarm calls. X removes 1 or all host eggs before she lays her own (host eggs appearing in parasitized nests are laid later), but sometimes fails to remove eggs of largest hosts. Nestling apparently interferes with survival of host nestlings, though nest of C. galactotes had 1 host young and 2 Cuckoo-Finches all ready to fly (the latter misidentified as ‘Vidua macroura’: Cheesman and Sclater 1935–1936). Foster parents Cisticola juncidis feed insects, mainly caterpillars and grasshoppers, also snails, to fledglings. EGGS: sets (‘clutches’) of 1–4 (mean, 29), 1 a day, a few days between sets. A parasitized nest has 1–2 Anomalospiza eggs; a X lays c. 30 eggs in a season. Little gloss; colour and markings vary, often match hosts’ eggs (bluish white with no spots, or with speckles and spots; whitish with red spots; pinkish with faint purplish at thick end and purplish and reddish brown spots and blotches) (Payne

1944, Neuby-Varty 1950, Vincent 1949, Vernon 1964). SIZE: 17  13; weight 16 g; larger than all but the largest host species, but small in relation to X body size. LAYING DATES: breeds with rains, along with its host species. W Africa, no breeding records. Ethiopia (W Highlands, Aug, Nov; E Africa, north and east of L. Victoria, Feb–June; Kenya, Jan, May, June, Oct; Pemba Island, Sept–Jan; Zaı¨re (Kivu, suspected May–June; Uele, ˆ i, Apr; Botswana, Aug–Sept); Zambia, Jan–Apr; Malaw Feb and Apr; Zimbabwe, Nov–Mar, mainly Dec–Feb; Transvaal, Sept–Mar. INCUBATION: period undetermined, less than 14 days. DEVELOPMENT AND CARE OF YOUNG: nestling period 18 days, longer than young of host. Parental care is by hosts, which feed entire insects directly to nestlings. Host and parasite chicks rarely fledge together. Failure of host nestlings is due to removal of host eggs by X Cuckoo-Finch and by competition or interference by nestling parasites, not by eviction by nestlings insofar as 2 Cuckoo-Finch young often fledge from same nest. By day 2 after hatching, 2 chicks trampled a Cisticola ayresii chick, which disappeared 2 days later. By day 4, nestling is nearly black above, mouth flesh-purple with deep orange-yellow border formed a distinct gape flange; bill horn above and yellow below with a terminal black spot. By day 7, bill heavier and darker, tufts of natal down remain on crown, back and wings, featherquills broken out over upperparts; spine in distinct tracts on head and middle of back. By day 9, eyes still closed, feather tips emerge on back, and small orange-brown feather quills appear on throat. By day 11, back feathers emerge with sandy brown colour. By day 13, 2 chicks nearly fill nest, face feathered and legs pale grey. By day 18, young leave nest and are fed by host pair as far as 30 m from nest, and can fly low but strongly for 50 m (Vernon 1964). Begging posture is upright, without twists and turns of head and neck; begging calls are ‘tink’ or ‘jit . . . jit’, like calls of an ad. (and unlike begging calls of host young: Vernon 1987, Herremans et al. 1993, Brewster 2000). Fledgling remains with host for 10 days, when it begins to feed itself, clinging on grass stems to reach seed-heads while still accompanied by foster parents, then leaves the foster family and joins flocks of conspecifics (Chapin 1954). Key References Chapin, J.P. (1954), Friedmann, H. (1960), Lahti, D. M. and Payne, R. B. (2003), Roberts, A. (1917, 1939), Vernon, C.J. (1964, 1987), Williams, J.G. and Keith, G.S. (1962).

Genus Vidua Cuvier

Brood-parasitic finches with sexually dimorphic breeding plumage, YY glossy black, short-tailed (10 indigobirds), or dull black and elaborate-tailed (8 whydahs), or glossy black and elaborate-tailed (V. hypocherina). XX and non-breeding YY streaked brown above and whitish below; juvs less streaked, often resemble juvs of their host species. Wing rounded, P10 smaller than coverts and pointed, P9–P6 longest and subequal in length, axillaries and underwing-coverts usually whitish or white. Tail with 12 feathers, inner 2 pairs (T1, T2) moult twice a year in Y whydahs and indigobirds; other tail-feathers

419

420

VIDUIDAE

moult once a year. Skull incompletely pneumatized in ads until 4–9 years of age. Breed by 1 year of age (XX, some YY). Feed on fallen grass seeds; scratch with both feet (‘double scratch’) nearly simultaneously to uncover seeds on bare earth, hop backwards, then peck up the exposed seed. Most Vidua spp. are brood parasites of one particular estrildid (some occasionally parasitize other estrildids); V. macroura regularly uses more than one estrildid species. Nestling mouth patterns like those of their host species. Y sings; most species mimic songs of their host species, and also have locally characteristic non-mimetic songs. Vidua brood parasites have not speciated in parallel with their hosts, but have switched from one host to another, and some colonists have emerged as distinct species (Klein et al. 1993, Klein and Payne 1998). Paradise whydahs V. paradisaea and V. orientalis use same host species, Pytilia melba, but molecular genetics indicates they are distinct species and are not each other’s closest relative; they do not interbreed where their ranges meet in E Sudan and Eritrea. Indigobird species are known as distinct from consistent association of colour, form and mimicry song (and known assortative mating in southern Africa). Up to 4 species of indigobirds live together; YY are interspecifically territorial. X is selective in which kinds of Y she visits and mates. Most indigobirds use a single species of host estrildid, and their nestlings (where known) mimic mouth patterns of host nestlings; V. camerunensis mimics several species of estrildid hosts. V. camerunensis has no consistent morphological differences between song mimics of Lagonosticta rubricata, L. rara, Clytospiza monteiri and Euschistospiza dybowskii; their song populations represent recent host switches and colonizations. Recent colonists, including V. raricola which uses Sporaeginthus subflavus, and V. nigeriae which uses Ortygospiza atricollis, are not nestling mimics (Payne and Payne 1994). Adults mimic specific begging calls in their songs (Payne and Payne 2002). Molecular genetics of indigobirds indicates recent speciation, post-dating that of host species (Klein and Payne 1998, Sorenson and Payne 2002, Sorenson et al. 2003). Phylogenetic relationships among species have been estimated from mitochondrial DNA mapped restriction sites and sequence data (Klein and Payne 1998, Sorenson and Payne 2002). The 5 paradise whydahs (Payne 1997a: V. paradisaea, V. obtusa, V. orientalis, V. interjecta, V. togoensis) all use Pytilia species. The 10 indigobirds (Payne 1996, 1998a) use firefinches Lagonosticta and Clytospiza, Euschistospiza, Hypargos, Ortygospiza atricollis and Sporaeginthus subflavus (which are not all closely related to Lagonosticta or to each other). V. fischeri and V. regia use Granatina. V. macroura and V. hypocherina use mainly Estrilda species. Most hosts of song-mimicking Vidua spp. have elaborate songs themselves. These Vidua spp. may have colonized them to acquire their sexual songs, which Vidua copy and use as their own sexual songs, as well as to gain the hosts’ parental care. Short-tailed indigobirds are derived from long-tailed whydahs, as indicated by genetics (Klein and Payne 1998, Sorenson and Payne, 2002) and by the double annual moult of central tail-feathers. Sometimes an indigobird grows a long tail one year and a short one the next (Neunzig 1931, Payne 1973a). Indigobirds V. chalybeata, V. codringtoni, V. funerea, V. purpurascens in E and S Africa are more closely related to each other than any is to indigobirds in W Africa including V. c. chalybeata and V. c. neumanni; W African V. nigeriae, V. raricola and V. maryae are each others’ closest relatives. Indigobirds are most closely related to V. regia and V. fischeri, while V. hypocherina is more distantly related; V. macroura is basal to other species (Klein and Payne 1998). The close relationship of Viduidae to Estrildidae (Bentz 1979, Groth 1998, Sibley and Ahlquist 1990; Sorenson and Payne 1998, 2001, 2002) suggests that the first viduids were parasitic on estrildids, with similar nestling gape and palate colours: later, colours evolved better resemblances and mimetic young received more consistent parental care. Indigobirds are sometimes placed in Hypochera; paradise-whydahs in Steganura and Straw-tailed and Shaft-tailed Whydahs V. fischeri and V. regia in Tetraenura. XX of most species are similar, and hybrids are known between speciesgroups. We retain all species in Vidua. Endemic, 19 species. No superspecies recognized.

Plate 25

Vidua macroura (Pallas). Pin-tailed Whydah. Veuve dominicaine.

(Opp. p. 394)

Fringilla macroura Pallas, 1764. Adumbrat, in Vroeg, Catalogue de quadrupedes et d’insectes . . ., p. 3, no. 144; East Indies (¼Angola, ex Edwards and Brisson). Range and Status. Endemic resident and migrant, widespread in grassy and open shrubby habitats through most of subsaharan Africa, absent from driest parts of E and SW Africa. Introduced to Re´union Is. Throughout W Africa west of 20
E, north to S Mauritania (Rosso), 17
N in Mali, 14
N in Niger (and once at Agadez), Ouaddai region in Chad (once at Fada, at 17
N). Bioko, Sa˜o Tome´. Central African Republic, except E. Central Sudan. NW Eritrea. Ethiopia (rare above 2600 m). Somalia (a few in extreme NW, in S mainly along

Shabeelle R., Jubba R. and coast south of 2
N). Kenya (widespread in S, absent in arid N except along Tana R. and extreme NE and NW corners). Uganda (widespread, sparse in arid N). South of Equator, widespread east of 20
E, less so west of it. Zanzibar. Namibia (Orange R.). Botswana (only in N, E and S border areas, where sparse). South Africa (sparse in dry western areas). Suspected seasonal migrant into drier areas during rains, as in S Sudan. In dry areas of SW Africa such as Namibia, range has expanded in past 50–100 years with dams and irrigation and colonization

420

VIDUIDAE

moult once a year. Skull incompletely pneumatized in ads until 4–9 years of age. Breed by 1 year of age (XX, some YY). Feed on fallen grass seeds; scratch with both feet (‘double scratch’) nearly simultaneously to uncover seeds on bare earth, hop backwards, then peck up the exposed seed. Most Vidua spp. are brood parasites of one particular estrildid (some occasionally parasitize other estrildids); V. macroura regularly uses more than one estrildid species. Nestling mouth patterns like those of their host species. Y sings; most species mimic songs of their host species, and also have locally characteristic non-mimetic songs. Vidua brood parasites have not speciated in parallel with their hosts, but have switched from one host to another, and some colonists have emerged as distinct species (Klein et al. 1993, Klein and Payne 1998). Paradise whydahs V. paradisaea and V. orientalis use same host species, Pytilia melba, but molecular genetics indicates they are distinct species and are not each other’s closest relative; they do not interbreed where their ranges meet in E Sudan and Eritrea. Indigobird species are known as distinct from consistent association of colour, form and mimicry song (and known assortative mating in southern Africa). Up to 4 species of indigobirds live together; YY are interspecifically territorial. X is selective in which kinds of Y she visits and mates. Most indigobirds use a single species of host estrildid, and their nestlings (where known) mimic mouth patterns of host nestlings; V. camerunensis mimics several species of estrildid hosts. V. camerunensis has no consistent morphological differences between song mimics of Lagonosticta rubricata, L. rara, Clytospiza monteiri and Euschistospiza dybowskii; their song populations represent recent host switches and colonizations. Recent colonists, including V. raricola which uses Sporaeginthus subflavus, and V. nigeriae which uses Ortygospiza atricollis, are not nestling mimics (Payne and Payne 1994). Adults mimic specific begging calls in their songs (Payne and Payne 2002). Molecular genetics of indigobirds indicates recent speciation, post-dating that of host species (Klein and Payne 1998, Sorenson and Payne 2002, Sorenson et al. 2003). Phylogenetic relationships among species have been estimated from mitochondrial DNA mapped restriction sites and sequence data (Klein and Payne 1998, Sorenson and Payne 2002). The 5 paradise whydahs (Payne 1997a: V. paradisaea, V. obtusa, V. orientalis, V. interjecta, V. togoensis) all use Pytilia species. The 10 indigobirds (Payne 1996, 1998a) use firefinches Lagonosticta and Clytospiza, Euschistospiza, Hypargos, Ortygospiza atricollis and Sporaeginthus subflavus (which are not all closely related to Lagonosticta or to each other). V. fischeri and V. regia use Granatina. V. macroura and V. hypocherina use mainly Estrilda species. Most hosts of song-mimicking Vidua spp. have elaborate songs themselves. These Vidua spp. may have colonized them to acquire their sexual songs, which Vidua copy and use as their own sexual songs, as well as to gain the hosts’ parental care. Short-tailed indigobirds are derived from long-tailed whydahs, as indicated by genetics (Klein and Payne 1998, Sorenson and Payne, 2002) and by the double annual moult of central tail-feathers. Sometimes an indigobird grows a long tail one year and a short one the next (Neunzig 1931, Payne 1973a). Indigobirds V. chalybeata, V. codringtoni, V. funerea, V. purpurascens in E and S Africa are more closely related to each other than any is to indigobirds in W Africa including V. c. chalybeata and V. c. neumanni; W African V. nigeriae, V. raricola and V. maryae are each others’ closest relatives. Indigobirds are most closely related to V. regia and V. fischeri, while V. hypocherina is more distantly related; V. macroura is basal to other species (Klein and Payne 1998). The close relationship of Viduidae to Estrildidae (Bentz 1979, Groth 1998, Sibley and Ahlquist 1990; Sorenson and Payne 1998, 2001, 2002) suggests that the first viduids were parasitic on estrildids, with similar nestling gape and palate colours: later, colours evolved better resemblances and mimetic young received more consistent parental care. Indigobirds are sometimes placed in Hypochera; paradise-whydahs in Steganura and Straw-tailed and Shaft-tailed Whydahs V. fischeri and V. regia in Tetraenura. XX of most species are similar, and hybrids are known between speciesgroups. We retain all species in Vidua. Endemic, 19 species. No superspecies recognized.

Plate 25

Vidua macroura (Pallas). Pin-tailed Whydah. Veuve dominicaine.

(Opp. p. 394)

Fringilla macroura Pallas, 1764. Adumbrat, in Vroeg, Catalogue de quadrupedes et d’insectes . . ., p. 3, no. 144; East Indies (¼Angola, ex Edwards and Brisson). Range and Status. Endemic resident and migrant, widespread in grassy and open shrubby habitats through most of subsaharan Africa, absent from driest parts of E and SW Africa. Introduced to Re´union Is. Throughout W Africa west of 20
E, north to S Mauritania (Rosso), 17
N in Mali, 14
N in Niger (and once at Agadez), Ouaddai region in Chad (once at Fada, at 17
N). Bioko, Sa˜o Tome´. Central African Republic, except E. Central Sudan. NW Eritrea. Ethiopia (rare above 2600 m). Somalia (a few in extreme NW, in S mainly along

Shabeelle R., Jubba R. and coast south of 2
N). Kenya (widespread in S, absent in arid N except along Tana R. and extreme NE and NW corners). Uganda (widespread, sparse in arid N). South of Equator, widespread east of 20
E, less so west of it. Zanzibar. Namibia (Orange R.). Botswana (only in N, E and S border areas, where sparse). South Africa (sparse in dry western areas). Suspected seasonal migrant into drier areas during rains, as in S Sudan. In dry areas of SW Africa such as Namibia, range has expanded in past 50–100 years with dams and irrigation and colonization

Vidua macroura

Vidua macroura

of resulting new habitats by host species, Common Waxbill Estrilda astrild (Harrison et al. 1997). Absent in some areas where host waxbills Estrilda are common (N Namib Coast in Namibia, Namaqualand in W Cape). Regarded officially as a pest in Benin; abundant there, but sharp decline observed in 1997, possibly due to an epidemic or to increasing use of pesticides in agricultural production (Claffey 1999). Description. ADULT Y (breeding): forehead and crown black, face and collar white, back black, rump white; in tail, T1–T2 black, central feathers narrow, longer than rest of bird, T3 black, T4–T6 white on tips and inner webs; chin black, throat to undertailcoverts white; wings black with white patch formed by lesser and median upperwing-coverts, underwing-coverts white mottled with black. Bill red, eyes dark brown, feet blackish grey. ADULT Y (non-breeding): like X, head pattern bolder, stripe on crown whitish not buff, superciliary stripe black, bill red. ADULT X: crown to rump rufous buff with blackish streaks, well-defined dark brown stripe on side of crown and through eye, superciliary stripe rufous to buff; tail brown, T4–T6 with white inner margins; chin and throat buffy white, breast buff with dark streaks on side, belly to undertail-coverts whitish; wings brown with buffy wing-bars formed by upperwing-coverts, underwingcoverts grey; bill black in breeding season, black with red base at end of breeding season and red in non-breeding season. Form ‘V. m. arenosa’ Clancey 1977, described as distinct in colours of Y breeding plumage (chin white), Y non-breeding plumage and X plumage in southern Africa, is not supported in larger sample. YY with long tarsi have longer T1, T2 in breeding plumage, independent of geographic region (Alatalo et al. 1988). SIZE: (Kenya to South Africa) wing, Y (n ¼ 33) 67–76 (712), X (n ¼ 17) 66–68 (668); tail, Y (n ¼ 34) 44–54 (509), T1 and T2 (n ¼ 24) 180–248 (224), X (n ¼ 18) 47–55 (506); bill, Y (n ¼ 34) 88–102 (95), X (n ¼ 18) 90–98 (94); tarsus, Y (n ¼ 34) 145–168 (158), X (n ¼ 18) 15–16 (153). WEIGHT: Kenya, Y (n ¼ 33) 133–188 (160), X (n ¼ 46) 122–169 (155), Y (n ¼ 30) 120–155 (139), X (n ¼ 28) 115–160 (131) (Savalli 1995); Cameroon, X (n ¼ 3) 128–149 (136); Transvaal, Y (n ¼ 9) 138–178 (151); S Cape, Y

(n ¼ 9) 148–167 (159), X (n ¼ 18) 134–159 (149) (Barnard ˆ i to South Africa, laying X (n ¼ 31) 133–167 1991a); Malaw (Payne 1997a). IMMATURE: juv. upperparts uniform mouse-brown, cheek paler, tail uniform mouse-brown, throat whitish, breast to undertailcoverts light buff; bill black (after independence, bill red from base), gape with white swellings, eyes dark brown, feet black. NESTLING: hatchling skin mauve, head pink, bill black, tarsi mauve, feet pink, nostrils prominent, natal down on crown and back dusky grey, pale yellow on secondaries and white on rump, eyes closed. Later, skin dark pinkish mauve, paler below, natal down grey, bill black, feet grey. Base of upper mandible forms a white, shallow, raised, U-shaped flange; ends of U meet a broad papilla on lower bill; side of gape has 3 distinct gape swellings, the lower papillae fitting into each side of comma-shaped upper ridge, outlined in black (A – Nicolai 1964). Palate pale pinkish with ring of 5 black spots, tongue pinkish with 2 black spots, and lower mouth with sublingual black bar.

Field Characters. Length Y 30–32 cm, X 11–12 cm. Blackand-white breeding Y with red bill and long, floppy black tail unmistakable. X has head and face striped dark brown and buff, pale rufous central crown stripe, bill black when breeding, red in non-breeding season. Non-breeding Y like X but head stripes black and white, bill red at all times. Both X and non-breeding Y have inner half of outer 3 tail feathers white, apparent in the field, especially from below. Juvenile plain mouse-brown with pale cheek, whitish throat and short, stubby black or red-and-black bill. Nestling and juv. mouth similar to that of nestling host waxbills Estrilda spp., but young whydah larger, nostrils prominent (not flat), and nestling covered with natal down (waxbills hatch naked). Voice. Tape-recorded (86, 88, 99, 104, B, C, F, BAR, PAY, STJ). Song an uneven, jerky series of measured, single notes with sibilant quality, ‘tsip, tse-tsuc, tyap, tsrrr, wee, tip, jaa . . . ’, interspersed with trills (8–10 notes per s), and runs of rapid, tinny jingling, less often a sweet whistle, down-slurred, ‘tee-yew’ or up-slurred, ‘way-yee’, and a grating sparrow-like ‘jaa-jaa’ or ‘jajaja’. Y also directs quiet canary-like twittering at intruding Y, with head held back and bill raised, ‘whit-whit-whit’ and ‘chee-chee-chee,’ interspersed with short warbling passages (Shaw 1984). Both sexes give harsh chatter. Other calls include low ‘peeee’ and in flight, double ‘chip-chip’. General Habits. Inhabits cultivated land and gardens, disturbed clearings and bush tracks; around villages. Also in clearings near margins of forests and along larger rivers, especially the Congo. In E and central Africa generally occurs below 2300 m. Occurs in same grassy areas as Estrilda waxbill hosts. Feeds alone or in flocks with other Vidua and estrildid finches. Breeding Y aggressive to other

421

422

VIDUIDAE

estrildids on feeding areas (Friedmann 1960, Savalli 1990); he uses particular sites near feeding areas to attract a mate and prevent mating interference by other birds. Uncovers seeds by a rapid double-scratch: rapidly kicks aside top soil with one foot then the other, hops backwards, and pecks at exposed seeds at rate of 2 pecks in 5 s. Young whydahs may join family groups of other bird species after independence from waxbill hosts. Occurs in flocks of 100 or more birds in non-breeding season. One bird homed from 53 km away in 6 h (Steyn 2002). Roosts in dense canopy of leafy trees. Food. Grass seeds 1–2 mm in diam., taken on ground. Eats seeds of Echinochloa, Urochloa and Setaria spp. (Lochinvar Nat. Park, Zambia, Payne 1980), Digitaria exilis (upland Guinea and Sierra Leone), Eleusine colonara (Uele, Zaı¨re), and Panicum (Cape Prov.). Takes pounded maize meal and occasionally nutlets from stems of small sedge Cyperus flavescens, filamentous algae in shallow streams, and flying termites. Breeding X takes host eggs. Breeding Habits. Polygynous, territorial. Brood parasitic, lays in roofed nests of waxbills Estrilda and occasionally other estrildids; hosts rear young whydahs in mixed brood with their own young. X usually visits nests alone (Y remains on call-site and mates with other XX) or is accompanied by Y (Roberts 1913, van Someren 1956). X often removes egg from host clutch and eats it (Roberts 1917, 1939, Payne 1997a). Nesting host may chase whydah and damage its own eggs when defending nest (Roberts 1939). Parasitized nest has as many as 5 Vidua eggs (Skead 1957, Schmidt 1965). Y sings from a few favoured perches in trees or shrubs, all day, mainly in early morning; chases other YY and other species from display area; and flies conspicuously from tree to tree with tail flopping. Y displays to visiting X and to any other bird that perches in his tree, including sunbirds, up to size of domestic fowl. In courtship display, Y flies at X on ground near his singing site, bounces in flight over her and flaps long tail feathers; he flies in a circle, hovers c. 05 m from her, his body inclined forward at angle of 45
, wings alternately vibrated and closed as he jerks up and down, tail cascading towards her (B). X crouches, wings held slightly out and down and quivering, tail fanned and vibrated, head tipped upwards. At end of display, Y flies back to song perch. X, after being flapped at on ground for up to 1 min, follows him to perch and they mate there. Perched Y sometimes gives rapid bouts of wing-shaking, plumage fluffed, in horizontal stance; he tilts forward, vibrates wings (C), then approaches her on ground. After courting her, he leads X from his display tree to feeding ground nearby, where they feed together. Only c. 8% of displays result in copulation. Y is more successful in attracting X and copulating when he has a permanent food and water source near display site. Y often uses same call-site in successive years. When Y is removed from call-site, he is often replaced by Y from another site; the new Y increases his own sexual success when he shifts to a more active site (Shaw 1984). Develops into breeding condition before host species does (Immelmann 1969).

No strong pair bond. X visits several neighbouring YY, and several XX visit and copulate with 1 particular Y. Other YY sometimes visit a Y at his site at the same time as X and interfere with his courtship. As many as 16 XX in a season visit an unusually active breeding Y that displays and copulates all day, with av. 038 copulations per h. Young Y not in breeding plumage displays same behaviour to X, singing from call-site and hovering in jerky flight with short central tail-feathers erected and tail spread. Hosts include the waxbills Estrilda astrild, E. rhodopyga, E. melpoda, E. paludicola and E. troglodytes. In Senegal, and probably Benin, Nigeria and Cameroon, parasitizes both E. troglodytes and E. melpoda (Morel and Morel 1990); in coastal Ghana uses them both and Bronze Mannikin Spermestes cucullatus (Macdonald 1980). Hosts in Liberia probably E. melpoda and E. astrild (Gatter 1997); on Sa˜o Tome´ and Principe´ probably E. astrild. In Gabon, parasitizes S. cucullatus (whose young have a bar, not 5 spots, on palate, and lack the white gape papillae). In Kenya probably uses E. astrild and E. rhodopyga, and occasionally African Silverbill Euodice cantans. In southern Africa uses Estrilda spp. and swee waxbills Coccopygia melanotis and C. quartinia (whose young lack black palate spots). In NW Transvaal and Kwazulu-Natal may use Zebra Waxbill Sporaeginthus subflavus (Mines 1999). Claims of other hosts, based on eggs alone, are mostly misidentifications (the

Vidua orientalis most reliable identifications are from nestlings’ mouth markings or feathered plumage, and molecular genetics: Payne et al. 2000a). EGGS: sets of 3–4, 1 egg a day, a few days between sets. X lays av. 25 in a season (Payne 1977a). Little gloss; immaculate white. SIZE: 145–166  110–122. LAYING DATES: breeds during rains, in nesting season of Estrilda waxbill host species. Senegal, Aug–Sept; Mali, July–Sept; Liberia, (Y in breeding plumage Apr– Oct (–Nov), juv. from Sept–Nov); coastal Ghana, Mar–Apr to Sept (YY breeding plumage, juvs), in N, Apr–Oct (most YY displaying in July); E Nigeria, Mar–Nov (YY in breeding plumage, egg in oviduct Oct and Nov); Gabon, breeding plumage all year, lays mainly in long dry season, June–Sept; Sa˜o Tome´, (breeding plumage in Jan–June); SW Central African Republic, (breeding plumage June–Oct). Uganda, Feb, Apr, Oct and Nov; Kenya, Apr–July (YY in breeding plumage in most months; at Kakamega from Apr– July YY are in breeding plumage at onset of long rains, some with long tails until early Dec); Tanzania, Dec–Mar; ˆ i, Mar; N Angola, (breeding plumage Nov–May); Malaw Zimbabwe, Jan–Mar; South Africa, Nov–Apr, varying with rainfall and local breeding schedules of hosts.

INCUBATION:

423

period 11 days (van Someren 1956, Skead

1957). DEVELOPMENT AND CARE OF YOUNG: nestling period 20 days. Nestling eyes are half open and quills on primary and secondary upperwing-coverts emerge on day 5, quills on back, rump, tail, breast, flanks, and humerals emerge on day 7, crown and throat on day 8, some cheek feathers out and eyes well open on day 9, head well feathered on day 11 (when call is unlike that of young waxbills, and begging calls of a fledged whydah (‘wehk-wehk-wehk’) are unlike those of fledged Common Waxbills (‘chewnk’: Skead 1957, Friedmann 1960)). Most hosts (Estrilda spp.) have nestlings with similar mouth patterns; nestling V. macroura not known to mimic variations among these host species, but rather matches pattern of E. astrild. Young whydah is reared with young waxbills and often fledges with them; both species fed by parent waxbills; sometimes 2 young whydahs fledge in a brood (Friedmann 1960, Macdonald 1980). Young remains in host family group for more than a week, later joins whydah flock (Skead 1957).

Key References Barnard, P. (1989, 1991a, b), Barnard, P. and Markus, M.B. (1989), Chapin, J.P. (1954), Friedmann, H. (1960), Nicolai, J. (1964, 1968d), Shaw, P. (1984).

Vidua orientalis Heuglin. Sahel Paradise-Whydah. Veuve a ` collier d’or.

Plate 26

Vidua paradisaea orientalis Heuglin, 1871. Orn. Nordost Afrikas, vol. 1, p. 583; Bogosland, Keren ¼ Eritrea.

(Opp. p. 395)

Range and Status. Endemic resident. S Mauritania (Dosso). Senegal (Richard-Toll and St Louis south to Bao-Bolou). Gambia (Farafenni; Belel Forest Park). Mali (Boucle du Baoule´, Foreˆt de Tienfala, Dioura, Ke Macina, Niger Delta, Ban Markala, Fiko, Ansongo). Burkina Faso (De´digou, Ouagadougou, Kaya, Beni). Niger (south of c. 15
N near Filingue, Tawa, Park ‘W’, Zinder, Tahoua, Farak, Lagane to L. Chad). Nigeria (south to Sokoto, Gusau and Kano, Yankari Nat. Park, Maiduguri, Yo at Lake Chad). N Cameroon (Waza Nat. Park to Garoua). Chad (Ennedi: Ellala, Fada and Guelta Basso; Ndjamena, Maniling, Abe´che´). Central African Republic (Vakaga Pre´f.). S Sudan (Darfur, Bahr el Ghazal (Tonj, Kawajana), Kordofan (100 km west of Nahud, Kadugli, Talodi), White Nile (Renk, El Duien, Kawa, Jebel Ein, Fung), Blue Nile (Sennar, Abu Zor, Abu Haraz, Abu Usher, Karkoj, Roseires, Wad Medani, Dinder R.)). Eritrea (W third, east to 385
E). Ethiopia (once at Gimb west of L. Tana, also between 8
and 10
N and 35
and 365
E, within range of V. paradisaea). (V. orientalis and V. paradisaea reported together at Sennar, Sudan, but only the V. orientalis specimens have dates and are likely to have originated there.) Description. V. o. aucupum Neumann: Senegal to NW Nigeria. ADULT Y (breeding): upperparts black, tail black with central feathers (T1, T2) twisted vertically to form a flag, T2 very long and broad from base to tip, not tapered; nape dark brownish red, below throat to breast brownish red to dark maroon, belly yellowish. Bill black, eyes dark brown, feet black. ADULT Y (nonbreeding): upperparts brown streaked with blackish central shaft

Vidua orientalis

and light brown edges, whitish crown-stripe bordered by black stripes, white stripe above eye, light grey face with black mark around ear; bill grey with blackish culmen, eyes dark brown, feet grey. ADULT X: upperparts light brown streaked with black,

Vidua orientalis most reliable identifications are from nestlings’ mouth markings or feathered plumage, and molecular genetics: Payne et al. 2000a). EGGS: sets of 3–4, 1 egg a day, a few days between sets. X lays av. 25 in a season (Payne 1977a). Little gloss; immaculate white. SIZE: 145–166  110–122. LAYING DATES: breeds during rains, in nesting season of Estrilda waxbill host species. Senegal, Aug–Sept; Mali, July–Sept; Liberia, (Y in breeding plumage Apr– Oct (–Nov), juv. from Sept–Nov); coastal Ghana, Mar–Apr to Sept (YY breeding plumage, juvs), in N, Apr–Oct (most YY displaying in July); E Nigeria, Mar–Nov (YY in breeding plumage, egg in oviduct Oct and Nov); Gabon, breeding plumage all year, lays mainly in long dry season, June–Sept; Sa˜o Tome´, (breeding plumage in Jan–June); SW Central African Republic, (breeding plumage June–Oct). Uganda, Feb, Apr, Oct and Nov; Kenya, Apr–July (YY in breeding plumage in most months; at Kakamega from Apr– July YY are in breeding plumage at onset of long rains, some with long tails until early Dec); Tanzania, Dec–Mar; ˆ i, Mar; N Angola, (breeding plumage Nov–May); Malaw Zimbabwe, Jan–Mar; South Africa, Nov–Apr, varying with rainfall and local breeding schedules of hosts.

INCUBATION:

423

period 11 days (van Someren 1956, Skead

1957). DEVELOPMENT AND CARE OF YOUNG: nestling period 20 days. Nestling eyes are half open and quills on primary and secondary upperwing-coverts emerge on day 5, quills on back, rump, tail, breast, flanks, and humerals emerge on day 7, crown and throat on day 8, some cheek feathers out and eyes well open on day 9, head well feathered on day 11 (when call is unlike that of young waxbills, and begging calls of a fledged whydah (‘wehk-wehk-wehk’) are unlike those of fledged Common Waxbills (‘chewnk’: Skead 1957, Friedmann 1960)). Most hosts (Estrilda spp.) have nestlings with similar mouth patterns; nestling V. macroura not known to mimic variations among these host species, but rather matches pattern of E. astrild. Young whydah is reared with young waxbills and often fledges with them; both species fed by parent waxbills; sometimes 2 young whydahs fledge in a brood (Friedmann 1960, Macdonald 1980). Young remains in host family group for more than a week, later joins whydah flock (Skead 1957).

Key References Barnard, P. (1989, 1991a, b), Barnard, P. and Markus, M.B. (1989), Chapin, J.P. (1954), Friedmann, H. (1960), Nicolai, J. (1964, 1968d), Shaw, P. (1984).

Vidua orientalis Heuglin. Sahel Paradise-Whydah. Veuve a ` collier d’or.

Plate 26

Vidua paradisaea orientalis Heuglin, 1871. Orn. Nordost Afrikas, vol. 1, p. 583; Bogosland, Keren ¼ Eritrea.

(Opp. p. 395)

Range and Status. Endemic resident. S Mauritania (Dosso). Senegal (Richard-Toll and St Louis south to Bao-Bolou). Gambia (Farafenni; Belel Forest Park). Mali (Boucle du Baoule´, Foreˆt de Tienfala, Dioura, Ke Macina, Niger Delta, Ban Markala, Fiko, Ansongo). Burkina Faso (De´digou, Ouagadougou, Kaya, Beni). Niger (south of c. 15
N near Filingue, Tawa, Park ‘W’, Zinder, Tahoua, Farak, Lagane to L. Chad). Nigeria (south to Sokoto, Gusau and Kano, Yankari Nat. Park, Maiduguri, Yo at Lake Chad). N Cameroon (Waza Nat. Park to Garoua). Chad (Ennedi: Ellala, Fada and Guelta Basso; Ndjamena, Maniling, Abe´che´). Central African Republic (Vakaga Pre´f.). S Sudan (Darfur, Bahr el Ghazal (Tonj, Kawajana), Kordofan (100 km west of Nahud, Kadugli, Talodi), White Nile (Renk, El Duien, Kawa, Jebel Ein, Fung), Blue Nile (Sennar, Abu Zor, Abu Haraz, Abu Usher, Karkoj, Roseires, Wad Medani, Dinder R.)). Eritrea (W third, east to 385
E). Ethiopia (once at Gimb west of L. Tana, also between 8
and 10
N and 35
and 365
E, within range of V. paradisaea). (V. orientalis and V. paradisaea reported together at Sennar, Sudan, but only the V. orientalis specimens have dates and are likely to have originated there.) Description. V. o. aucupum Neumann: Senegal to NW Nigeria. ADULT Y (breeding): upperparts black, tail black with central feathers (T1, T2) twisted vertically to form a flag, T2 very long and broad from base to tip, not tapered; nape dark brownish red, below throat to breast brownish red to dark maroon, belly yellowish. Bill black, eyes dark brown, feet black. ADULT Y (nonbreeding): upperparts brown streaked with blackish central shaft

Vidua orientalis

and light brown edges, whitish crown-stripe bordered by black stripes, white stripe above eye, light grey face with black mark around ear; bill grey with blackish culmen, eyes dark brown, feet grey. ADULT X: upperparts light brown streaked with black,

424

VIDUIDAE

wings brown, crown with pale buff streak bordered by dark brown streaks, a pale streak above eye and a dark brown streak through eye, face whitish grey, below breast buff, belly and undertail-coverts white, tail brown, slightly graduated with T1 (central feathers) the longest, fresh T6 with 1-mm white margin; upperwing-coverts grey; bill grey, eyes dark brown or reddish brown, feet grey. SIZE: wing, Y (n ¼ 56) 72–78 (754), X (n ¼ 6) 71–75 (730); tail, Y T2 (n ¼ 39) 202–262 (233), width (depth) 26– 34, T3 (n ¼ 25) 56–67 (614), X (n ¼ 6) 51–54 (528); bill, Y (n ¼ 10) 94–109 (100), X (n ¼ 6) 91–104 (96); tarsus, Y (n ¼ 4) 16–18 (174), X (n ¼ 6) 15–17 (156). WEIGHT: Senegal, Y (n ¼ 3) 19–27 (240); N Nigeria, Y (n ¼ 4) 19–22 (205). IMMATURE: juv. above plain grey, rump grey, paler below, tail rounded to slightly graduated; bill black, eyes dark brown, feet grey. NESTLING: skin dark; down, gape colour and marks may be similar to host, Green-winged Pytilia Pytilia melba. V. o. orientalis Heuglin: Chad to Sudan and Eritrea. Y nape pale straw, X similar to aucupum. SIZE: Sudan and Chad, wing, Y (n ¼ 53) 74–79 (768), X (n ¼ 9) 71–74 (729). WEIGHT: Cameroon and Chad, Y (n ¼ 2) 170, 205 (187), X (n ¼ 2) 15–17 (160). Intergrades with V. o. aucupum near L. Chad.

Field Characters. Length Y 30–31 cm, X 13–14 cm. Breeding Y with long, broad tail; upperparts black with pale collar; below, rufous breast with buff belly; X with whydah head pattern. Differ by tail from other Y paradisewhydahs in its range, fully grown tail appears less than 3 times wing length (tail longer in Exclamatory ParadiseWhydah V. interjecta and Togo Paradise-Whydah V. togoensis), in the hand 3 times (vs 4 times) wing length. Head and upperparts black with pale collar, nape dark chestnut (V. o. aucupum, Senegal to Nigeria) or pale straw (nominate race, Chad to Eritrea), breast rufous, belly yellow-buff. X has head pattern of other West African paradise-whydahs, differs from Long-tailed Paradise-Whydah V. paradisaea by lack of dark cheek mark. Voice. Tape-recorded (104, BAR, PAY). Mimics songs and calls of its host Green-winged Pytilia; in W Africa songs of host and its mimic are ‘veet’ followed by 2 or 3 plaintive slurred whistles, shorter and more simple than the

complex song in E and S Africa. Mimics host calls; alarm ‘pik’ and contact ‘see-eh’. Also has non-mimetic wavery chatter, a rapid chatter, and a single ‘chuff’. General Habits. Inhabits grassy, sahel acacia savanna and woodland with scattered shrubs and open ground, sometimes with rocky pavement. Throughout range, occurs with host species, red-lored Green-winged Pytilia P. m. citerior (Nicolai 1977) and grey-lored P. m. jessei (V. paradisaea in Ethiopia and N and E Kenya occurs with red-lored P. m. soudanensis). Feeds on ground and termite mounds, alone or in small flocks. Resident, but a Y at 2130 m at Gimb, W Ethiopia, (the only whydah there in years: Cheesman and Sclater 1936), was probably a vagrant. Food. Grass seeds 1–2 mm in diam. Breeding Habits. Brood parasite of Green-winged Pytilia P. melba (as evidenced by geographic distribution and song mimicry). No field observations of brood parasitism. In captivity lays in P. melba nests and sometimes other estrildids. EGGS: little gloss; immaculate white. SIZE: unknown. LAYING DATES: Senegal, Nigeria and Sudan, (July–Dec, YY in breeding plumage, and juvs; copulation, Sudan, Sept); Gambia, (fledgling Jan); Mali, (Boucle de Baoule´, YY in breeding plumage Dec–Mar: de Bie and Morgan 1989). INCUBATION: period 11–13 days. DEVELOPMENT AND CARE OF YOUNG: no field information. In captivity, in absence of P. melba, has parasitized Redbilled Firefinch Lagonosticta senegala, Bengalese Finch Lonchura striata and canary Serinus canaria. Nestling period when reared by firefinch, 16 days; moult into ad. plumage begins by 6 weeks after fledging. Key References Benedetti, R. and Mignone, G.P. (1990), Capecchi, A. and Mignone, G.P. (1990), Garcia, L. (1996), Lloyd, T. (1955), Nicolai, J. (1964, 1968d), Payne, R.B. (1977a, 1985c, 1997a, b, 1998b).

Plate 26

Vidua interjecta (Grote). Exclamatory Paradise-Whydah. Veuve nige´rienne.

(Opp. p. 395)

Steganura paradisaea interjecta Grote, 1922. J. Orn., 70, p. 402; Weg Nola–Mbaika, ‘Kamerun’ ¼ Central African Republic. Range and Status. Endemic resident. Gambia (Kaur, Kiang West Nat. Park, Bansang). Senegal (Casamance). Guinea-Bissau (Mansoa, Dulombi-Bafata). Guinea (Labe´, Le´louma, Dalaba, Ria Bakanya southeast of Dabola, LayaFaranah, Kouroussa). Mali (Bamako, Foreˆt de Tienfala, Kulikoro). Burkina Faso (Arli Nat. Park). Ivory Coast (Korhogo, Bandama R.). Benin (Pendjari Nat. Park). Ghana (Mole Nat. Park). Togo (Naboulgou, Landa-Pozanda). Niger (Park ‘W’). Nigeria (Zaria, Upper Assop, Karu, Yankari Nat. Park, 176 km northwest of Yola, Baissa-Abong, Enugu, Abakaliki) (whydahs at Potiskum, Kainji and Igbetti (Elgood et al. 1994) are thought to be V. interjecta). Cameroon (Tibati, Kumbe, Ngaounde´re´-Ngaoundal). Chad (Banda ¼ Baala Uham ¼ Ouam ¼ Uamgebiet, Bahr Keita, Gore).

Central African Republic (Fort Sibut, Gaza, Nola-Mbaika, Zemio). Sudan (Bengengai, Boma, Logoforok, Torit, Roseires). Zaı¨re (Uele: Garamba, Gangala-na-Bodio, Faradje, Aba). Ethiopia (Borraga-Kokolata, Baro-Bonga fork, Gambela, Bahar Dar). Range occurs within that of host species Red-winged Pytilia Pytilia phoenicoptera, of Redbilled Pytilia P. lineata in Ethiopia, and from Nigeria eastwards it occurs also in range of Yellow-winged Pytilia P. hypogrammica. Description. ADULT Y (breeding): upperparts black, nape dark brownish red; throat to breast brownish red to dark maroon, belly yellowish; tail black, with T1 and T2 twisted vertically to form a flag, T2 very long, and broad to near tip. Bill black, eyes dark brown, feet dark brown-grey to grey. ADULT Y

424

VIDUIDAE

wings brown, crown with pale buff streak bordered by dark brown streaks, a pale streak above eye and a dark brown streak through eye, face whitish grey, below breast buff, belly and undertail-coverts white, tail brown, slightly graduated with T1 (central feathers) the longest, fresh T6 with 1-mm white margin; upperwing-coverts grey; bill grey, eyes dark brown or reddish brown, feet grey. SIZE: wing, Y (n ¼ 56) 72–78 (754), X (n ¼ 6) 71–75 (730); tail, Y T2 (n ¼ 39) 202–262 (233), width (depth) 26– 34, T3 (n ¼ 25) 56–67 (614), X (n ¼ 6) 51–54 (528); bill, Y (n ¼ 10) 94–109 (100), X (n ¼ 6) 91–104 (96); tarsus, Y (n ¼ 4) 16–18 (174), X (n ¼ 6) 15–17 (156). WEIGHT: Senegal, Y (n ¼ 3) 19–27 (240); N Nigeria, Y (n ¼ 4) 19–22 (205). IMMATURE: juv. above plain grey, rump grey, paler below, tail rounded to slightly graduated; bill black, eyes dark brown, feet grey. NESTLING: skin dark; down, gape colour and marks may be similar to host, Green-winged Pytilia Pytilia melba. V. o. orientalis Heuglin: Chad to Sudan and Eritrea. Y nape pale straw, X similar to aucupum. SIZE: Sudan and Chad, wing, Y (n ¼ 53) 74–79 (768), X (n ¼ 9) 71–74 (729). WEIGHT: Cameroon and Chad, Y (n ¼ 2) 170, 205 (187), X (n ¼ 2) 15–17 (160). Intergrades with V. o. aucupum near L. Chad.

Field Characters. Length Y 30–31 cm, X 13–14 cm. Breeding Y with long, broad tail; upperparts black with pale collar; below, rufous breast with buff belly; X with whydah head pattern. Differ by tail from other Y paradisewhydahs in its range, fully grown tail appears less than 3 times wing length (tail longer in Exclamatory ParadiseWhydah V. interjecta and Togo Paradise-Whydah V. togoensis), in the hand 3 times (vs 4 times) wing length. Head and upperparts black with pale collar, nape dark chestnut (V. o. aucupum, Senegal to Nigeria) or pale straw (nominate race, Chad to Eritrea), breast rufous, belly yellow-buff. X has head pattern of other West African paradise-whydahs, differs from Long-tailed Paradise-Whydah V. paradisaea by lack of dark cheek mark. Voice. Tape-recorded (104, BAR, PAY). Mimics songs and calls of its host Green-winged Pytilia; in W Africa songs of host and its mimic are ‘veet’ followed by 2 or 3 plaintive slurred whistles, shorter and more simple than the

complex song in E and S Africa. Mimics host calls; alarm ‘pik’ and contact ‘see-eh’. Also has non-mimetic wavery chatter, a rapid chatter, and a single ‘chuff’. General Habits. Inhabits grassy, sahel acacia savanna and woodland with scattered shrubs and open ground, sometimes with rocky pavement. Throughout range, occurs with host species, red-lored Green-winged Pytilia P. m. citerior (Nicolai 1977) and grey-lored P. m. jessei (V. paradisaea in Ethiopia and N and E Kenya occurs with red-lored P. m. soudanensis). Feeds on ground and termite mounds, alone or in small flocks. Resident, but a Y at 2130 m at Gimb, W Ethiopia, (the only whydah there in years: Cheesman and Sclater 1936), was probably a vagrant. Food. Grass seeds 1–2 mm in diam. Breeding Habits. Brood parasite of Green-winged Pytilia P. melba (as evidenced by geographic distribution and song mimicry). No field observations of brood parasitism. In captivity lays in P. melba nests and sometimes other estrildids. EGGS: little gloss; immaculate white. SIZE: unknown. LAYING DATES: Senegal, Nigeria and Sudan, (July–Dec, YY in breeding plumage, and juvs; copulation, Sudan, Sept); Gambia, (fledgling Jan); Mali, (Boucle de Baoule´, YY in breeding plumage Dec–Mar: de Bie and Morgan 1989). INCUBATION: period 11–13 days. DEVELOPMENT AND CARE OF YOUNG: no field information. In captivity, in absence of P. melba, has parasitized Redbilled Firefinch Lagonosticta senegala, Bengalese Finch Lonchura striata and canary Serinus canaria. Nestling period when reared by firefinch, 16 days; moult into ad. plumage begins by 6 weeks after fledging. Key References Benedetti, R. and Mignone, G.P. (1990), Capecchi, A. and Mignone, G.P. (1990), Garcia, L. (1996), Lloyd, T. (1955), Nicolai, J. (1964, 1968d), Payne, R.B. (1977a, 1985c, 1997a, b, 1998b).

Plate 26

Vidua interjecta (Grote). Exclamatory Paradise-Whydah. Veuve nige´rienne.

(Opp. p. 395)

Steganura paradisaea interjecta Grote, 1922. J. Orn., 70, p. 402; Weg Nola–Mbaika, ‘Kamerun’ ¼ Central African Republic. Range and Status. Endemic resident. Gambia (Kaur, Kiang West Nat. Park, Bansang). Senegal (Casamance). Guinea-Bissau (Mansoa, Dulombi-Bafata). Guinea (Labe´, Le´louma, Dalaba, Ria Bakanya southeast of Dabola, LayaFaranah, Kouroussa). Mali (Bamako, Foreˆt de Tienfala, Kulikoro). Burkina Faso (Arli Nat. Park). Ivory Coast (Korhogo, Bandama R.). Benin (Pendjari Nat. Park). Ghana (Mole Nat. Park). Togo (Naboulgou, Landa-Pozanda). Niger (Park ‘W’). Nigeria (Zaria, Upper Assop, Karu, Yankari Nat. Park, 176 km northwest of Yola, Baissa-Abong, Enugu, Abakaliki) (whydahs at Potiskum, Kainji and Igbetti (Elgood et al. 1994) are thought to be V. interjecta). Cameroon (Tibati, Kumbe, Ngaounde´re´-Ngaoundal). Chad (Banda ¼ Baala Uham ¼ Ouam ¼ Uamgebiet, Bahr Keita, Gore).

Central African Republic (Fort Sibut, Gaza, Nola-Mbaika, Zemio). Sudan (Bengengai, Boma, Logoforok, Torit, Roseires). Zaı¨re (Uele: Garamba, Gangala-na-Bodio, Faradje, Aba). Ethiopia (Borraga-Kokolata, Baro-Bonga fork, Gambela, Bahar Dar). Range occurs within that of host species Red-winged Pytilia Pytilia phoenicoptera, of Redbilled Pytilia P. lineata in Ethiopia, and from Nigeria eastwards it occurs also in range of Yellow-winged Pytilia P. hypogrammica. Description. ADULT Y (breeding): upperparts black, nape dark brownish red; throat to breast brownish red to dark maroon, belly yellowish; tail black, with T1 and T2 twisted vertically to form a flag, T2 very long, and broad to near tip. Bill black, eyes dark brown, feet dark brown-grey to grey. ADULT Y

Vidua interjecta

Vidua interjecta

exclamation mark. Nape dark brownish red, as in W African race of Sahel, darker than in Togo and E race of Sahel; maroon on breast extends farther back than in Togo, giving 2-tone appearance to underparts, which are more uniformly pale in Togo (Payne 1985c). X has upperparts streaked greyish brown (yellowish or buffy in some Sahel), whydah face pattern, with dark line behind eye as in Sahel and Broad-tailed Paradise-Whydah V. obtusa, not forming a ‘C’ mark behind ear as in Longtailed Paradise-Whydah V. paradisaea. Most distinctive feature is reddish colour of bill and feet (grey in Sahel). Voice. Tape-recorded (104, BAR, PAY). Mimics songs and calls of host, Red-winged Pytilia (Gambia and Guinea to N Nigeria, Payne 1991, 1997a). Mimicry songs include a sharp ‘pik’ or ‘tik’ followed by slurred whistle that rises then falls, ‘tik-feew’; short whistle followed by short buzz, a pair of loud descending whistles (double ‘chuck’) or a train of these whistles, a 2-part buzz-trill ‘peeeezyoooo’, low, grating ‘churr’, and ‘chink’ alarm call. Mimicry of Yellow-winged Pytilia would be similar except that song is 2-part trill, first slow then fast, with no change in pitch, rather than buzz-trill of Red-winged Pytilia. Adult Y whydah also gives non-mimetic calls, a soft chatter in flight, a loud rapid chatter ‘dzidzidzit’ when perched, and loud complex chatter before flying.

(non-breeding): upperparts brown streaked with blackish (feathers with blackish central shaft and light brown edges), crown with medial whitish stripe bordered by black stripes, a white stripe above eye, light orange-grey face with black mark around ear; bill orange-yellow with blackish culmen, feet fleshy grey. ADULT X: upperparts light brown streaked with black, tail brown, slightly graduated with inner feathers the longest, fresh outer ones with 1 mm white margin; crown with pale buff streak, bordered by dark brown crown streaks, face plain buff except for pale streak above eye and dark brown streak through eye; breast buff, belly and undertail-coverts white; wings brown, upperwing-coverts grey; bill light orange, eyes dark brown or reddish brown, feet pastel red. SIZE: wing, Y (n ¼ 41) 75–79 (762), X (n ¼ 4) 74–76 (748); tail, Y T2 (n ¼ 23) 270–340 (292), width 28–36, T3 (n ¼ 27) 56–65 (587), X (n ¼ 4) 51–58 (547); bill, Y (n ¼ 13) 95–116 (106), X (n ¼ 4) 91–104 (100); tarsus, Y (n ¼ 13) 160–184 (167), X (n ¼ 3) 150–155 (152). WEIGHT: Nigeria, Y (n ¼ 5) 20– 21 (206). IMMATURE: juv. plain grey-brown, like juv. Pytilia hosts but without barring or red rump (Nicolai 1977). NESTLING: skin dark, natal down on upperparts thick and grey. Gape swellings purplish white, palate whitish pink with purplish lateral patches and no central black spot. Nestling and young juv. have mouth like that of hosts, Red-winged and Yellow-winged Pytilias. Reddish palate, black lining to tip of bill, and whitish tip of tongue (like both hosts), small central black spot on palate (missing in hosts), lateral to median fissure in palate has pair of large, elongate blue-violet patches (as in P. phoenicoptera, not small and round as in P. hypogrammica). At fledging, bill short, stubby and deep at level of nostrils, and head large and broad (compared with Pytilia, Nicolai 1964, 1977).

Field Characters. Length Y 38–40 cm, X 13–14 cm. Tail of breeding Y broad, about the same as body depth and constant throughout (not tapering), similar in shape to that of Sahel Paradise-Whydah V. orientalis but longer, more than 3 times wing length (less in Sahel); broader than tail of Togo Paradise-Whydah. In flight bird looks like a flying

General Habits. Inhabits guinean woodlands, open grassy woodlands with bare ground, tall grass, and scattered trees and bushes, and rocky hillsides. Occurs within range and in same habitats as Red-winged and Yellow-winged Pytilias. Occurs in flocks, in non-breeding season. Feeds on the ground, alone or in small groups. Food. Grass seeds 1–2 mm in diam., taken from ground. In upland Guinea and Sierra Leone, feeds on Digitaria exilis seeds during and harvesting in Sept–Oct. Breeding Habits. Brood-parasitic on Pytilia phoenicoptera and (from Nigeria eastward) P. hypogrammica (2 species with nearly identical nestling mouth patterns and colours). Parasitized nests had 3 whydah and 4 pytilia eggs, 1 whydah and 3 pytilia eggs, and 2 whydah and 4 pytilia eggs (Nicolai 1977). In courtship display, Y flies around territory with bursts of wingbeats, tail trailing, in fore-and-aft rocking motion he circles 50 m above ground, moves from tree to tree, then perches in tree top and sings mimicry songs (Barlow et al. 1997). He chases intruding Y for 200–300 m out of territory, flying below it. Y courts perched X in frontal display; he turns c. 90
to each side, throws back head, and gives mimicry song, hovering in front of her and over her. EGGS: little gloss; immaculate white. SIZE: (n ¼ 1) 172  133 (in nest of Pytilia hypogrammica, Enugu, Nigeria; host eggs smaller, 148–152  115–119: Serle 1957). LAYING DATES: Gambia and Guinea (YY in breeding plumage, July to Nov–Dec); Nigeria, Nov–Jan, (courting YY, Zaria, Aug–Sept). INCUBATION: period 1175 days (shorter than 12–13 days of pytilia eggs when reared together by foster Bengalese Finches Lonchura striata: Nicolai 1977).

425

426

VIDUIDAE

DEVELOPMENT AND CARE OF YOUNG: nestling period 14 days (shorter than 16–18 days in P. hypogrammica when reared together by foster Bengalese Finch: Nicolai 1977), though P. hypogrammica reared by their own parents fledge in 21 days. Nestling swings head from side to side as it begs. Calls of fledged young are like those of P. hypogrammica. Feeds independently by day 26 or 27 (11–12 days after it leaves the nest) (pytilia young by day

36). Chatters like ad. by day 30. Y moults from juv. plumage to X-like striped plumage in a few weeks, and into full breeding plumage in 135 months (Nicolai 1977). Key References Chapin, J.P. (1929a, b), Delacour, J. and Edmond-Blanc, F. (1933–1934), Nicolai, J. (1964, 1968b, d, 1977), Payne, R.B. (1985c, 1991, 1994b, 1997a, b), Serle, W. (1957).

Plate 26

Vidua togoensis (Grote). Togo Paradise-Whydah. Veuve du Togo.

(Opp. p. 395)

Steganura paradisaea togoensis Grote, 1923. Orn. Monatsb., 31, p. 43; Kete, Togoland [ ¼ Ghana]. Range and Status. Endemic resident, W Africa. Guinea (Rio Bakanya, Fouta Djalon). Sierra Leone (Kabala, Karina). Mali, Bougouni, and reported from 6 localities including Se´nomango and Be´li depression, in Dogon region (Lamarche 1993). Ivory Coast (Bandama R.). Ghana (Gambaga, Mole Nat. Park, Keta, Ke´te´ Kratchi, Yendi, Tumu). Togo (Paio, Mangu, Sansanne-Mango). Cameroon, recently reported from 7 localities in Mbam R. drainage, between Foumban and E end of L. M’Bakaou (M. Languy, pers. comm.). Except for Dogon region, Mali, range is within that of its host species, Yellow-winged Pytilia Pytilia hypogrammica. Description. ADULT Y (breeding): upperparts black, nape yellow; below, throat and upper breast light rust, lower breast and belly yellowish, tail black, T1 and T2 twisted vertically to form a flag, T2 very long and narrow, feather width nearly equal throughout its length. Bill black, eyes dark brown, feet dark brown-grey to grey. ADULT Y (non-breeding): similar to non-breeding V. interjecta. ADULT X: unknown. SIZE (Y): wing, (n ¼ 8) 74–78 (768); tail, T2 length (n ¼ 4) 290–370 (320), width 22–24, T6 length (n ¼ 7) 57–60 (506); bill, (n ¼ 10) 92–103 (96); tarsus, (n ¼ 2) 166–170 (168). IMMATURE: juv. upperparts plain grey, paler below; tail rounded to slightly graduated; bill with reddish base and dark tip (presumably black in younger bird) (Payne 1991, 1997a). NESTLING: unknown.

Vidua togoensis

Mali: see Introduction.

Field Characters. Length Y 40–43 cm. Woodlands from Guinea to Togo. Breeding Y has long tail narrower than depth of body (tails broader in Exclamatory and Sahel Paradise-Whydahs V. interjecta and V. orientalis), nape yellow (red-brown in Exclamatory and W race of Sahel), yellow of underparts more extensive. Voice. Tape-recorded (PAY). Song thought to mimic Yellow-winged Pytilia Pytilia hypogrammica, q.v. In Mali and Ghana also gives chatters and songs like Exclamatory Paradise-Whydah. General Habits. Open woodland and cultivation. Shares habitat and breeding ecology with its apparent host species P. hypogrammica, in moist open woodland. Feeds on ground, alone or in small groups. Food. Grass seeds 1–2 mm in diam.

Breeding Habits. Unknown. By distribution in W upper guinean region, presumably brood-parasitic on P. hypogrammica (Friedmann 1960, Nicolai 1964, Payne 1985c). (A record of brood parasitism of ‘togoensis’ on P. hypogrammica in SE Nigeria (Nicolai 1977) applies to V. interjecta). LAYING DATES: Mali, (YY in breeding plumage, Sept); Ghana, (YY in breeding plumage and courting XX, Oct); Sierra Leone (Y in breeding plumage). Key References 1991, 1997a).

Nicolai, J. (1964, 1977), Payne, R.B. (1985c,

426

VIDUIDAE

DEVELOPMENT AND CARE OF YOUNG: nestling period 14 days (shorter than 16–18 days in P. hypogrammica when reared together by foster Bengalese Finch: Nicolai 1977), though P. hypogrammica reared by their own parents fledge in 21 days. Nestling swings head from side to side as it begs. Calls of fledged young are like those of P. hypogrammica. Feeds independently by day 26 or 27 (11–12 days after it leaves the nest) (pytilia young by day

36). Chatters like ad. by day 30. Y moults from juv. plumage to X-like striped plumage in a few weeks, and into full breeding plumage in 135 months (Nicolai 1977). Key References Chapin, J.P. (1929a, b), Delacour, J. and Edmond-Blanc, F. (1933–1934), Nicolai, J. (1964, 1968b, d, 1977), Payne, R.B. (1985c, 1991, 1994b, 1997a, b), Serle, W. (1957).

Plate 26

Vidua togoensis (Grote). Togo Paradise-Whydah. Veuve du Togo.

(Opp. p. 395)

Steganura paradisaea togoensis Grote, 1923. Orn. Monatsb., 31, p. 43; Kete, Togoland [ ¼ Ghana]. Range and Status. Endemic resident, W Africa. Guinea (Rio Bakanya, Fouta Djalon). Sierra Leone (Kabala, Karina). Mali, Bougouni, and reported from 6 localities including Se´nomango and Be´li depression, in Dogon region (Lamarche 1993). Ivory Coast (Bandama R.). Ghana (Gambaga, Mole Nat. Park, Keta, Ke´te´ Kratchi, Yendi, Tumu). Togo (Paio, Mangu, Sansanne-Mango). Cameroon, recently reported from 7 localities in Mbam R. drainage, between Foumban and E end of L. M’Bakaou (M. Languy, pers. comm.). Except for Dogon region, Mali, range is within that of its host species, Yellow-winged Pytilia Pytilia hypogrammica. Description. ADULT Y (breeding): upperparts black, nape yellow; below, throat and upper breast light rust, lower breast and belly yellowish, tail black, T1 and T2 twisted vertically to form a flag, T2 very long and narrow, feather width nearly equal throughout its length. Bill black, eyes dark brown, feet dark brown-grey to grey. ADULT Y (non-breeding): similar to non-breeding V. interjecta. ADULT X: unknown. SIZE (Y): wing, (n ¼ 8) 74–78 (768); tail, T2 length (n ¼ 4) 290–370 (320), width 22–24, T6 length (n ¼ 7) 57–60 (506); bill, (n ¼ 10) 92–103 (96); tarsus, (n ¼ 2) 166–170 (168). IMMATURE: juv. upperparts plain grey, paler below; tail rounded to slightly graduated; bill with reddish base and dark tip (presumably black in younger bird) (Payne 1991, 1997a). NESTLING: unknown.

Vidua togoensis

Mali: see Introduction.

Field Characters. Length Y 40–43 cm. Woodlands from Guinea to Togo. Breeding Y has long tail narrower than depth of body (tails broader in Exclamatory and Sahel Paradise-Whydahs V. interjecta and V. orientalis), nape yellow (red-brown in Exclamatory and W race of Sahel), yellow of underparts more extensive. Voice. Tape-recorded (PAY). Song thought to mimic Yellow-winged Pytilia Pytilia hypogrammica, q.v. In Mali and Ghana also gives chatters and songs like Exclamatory Paradise-Whydah. General Habits. Open woodland and cultivation. Shares habitat and breeding ecology with its apparent host species P. hypogrammica, in moist open woodland. Feeds on ground, alone or in small groups. Food. Grass seeds 1–2 mm in diam.

Breeding Habits. Unknown. By distribution in W upper guinean region, presumably brood-parasitic on P. hypogrammica (Friedmann 1960, Nicolai 1964, Payne 1985c). (A record of brood parasitism of ‘togoensis’ on P. hypogrammica in SE Nigeria (Nicolai 1977) applies to V. interjecta). LAYING DATES: Mali, (YY in breeding plumage, Sept); Ghana, (YY in breeding plumage and courting XX, Oct); Sierra Leone (Y in breeding plumage). Key References 1991, 1997a).

Nicolai, J. (1964, 1977), Payne, R.B. (1985c,

Vidua obtusa

427

Vidua obtusa (Chapin). Broad-tailed Paradise-Whydah. Veuve de Chapin.

Plate 26

Steganura aucupum obtusa Chapin, 1922. Amer. Mus. Nov., 43, p. 6; Luchenza, Nyasaland.

(Opp. p. 395)

Range and Status. Endemic resident. Uganda (observed Ruwenzori Nat. Park and Mbarara – no recent records). Kenya (E slope Mt Kenya: Meru, Chuka – only 3 records, most recent 1947). S and E Zaı¨re (Kivu and Katanga, Marungu Mts and Kasai (Kabinda, Bakwanga and Luluabourg to Chute Guillaume on Kwango R.)). Angola (central and N Huı´la north to Bie´, Luanda, Bengo, Cuanza Norte and Malanje, and east to Lunda Norte and Moxico). Tanzania (Mbeya north to Rukwa and Malangali and Iringa highlands regions southward, near L. Tanganyika at Ngara, Kibondo and Ufipa Plateau). Zambia (throughout, except for Luangwa Valley and sandy soils west of upper ˆ i (Rumphi, Nkhata Zambezi R. basin in W Zambia). Malaw Bay, Nkota-kota, Monkey Bay, Zomba, Blantyre, Chididi hills). Botswana (Chobe). Namibia (Caprivi). Zimbabwe (upper Zambezi R. from Kazungula to Victoria Falls, middle Zambezi R. along E end of L. Kariba and in wooded N, W and SE). Mozambique (Inhaminga, Beira; Tete, Mocuba, Malemo). Transvaal (Soutpansberg, Tzaneen). Range falls within that of host, Orange-winged Pytilia Pytilia afra. Description. ADULT Y (breeding): upperparts black; tail black, long and broad, T2 broad from base to near tip, at 40 mm width greater than body of bird as seen from side. Nape dark coppery rufous, breast dark maroon, belly and undertail-coverts buff. Bill black, eyes dark brown, feet dark brown to black. ADULT Y (nonbreeding): upperparts brown streaked with blackish (feathers with blackish shaft and light brown edges), crown with medial whitish stripe bordered by black stripes, white stripe above eye, face light grey; bill grey above and whitish below, feet fleshy grey to light grey. ADULT X: upperparts grey-brown, streaked blackish, crown with whitish streak bordered by blackish one, face with whitish superciliary streak, face lacks vertical marks; breast light grey, sometimes with indistinct streaks, belly white, undertailcoverts whitish, underwing-coverts grey; bill light grey to pinkish grey (paler on base of lower mandible), paler in nonbreeding season, eyes dark brown, feet light grey or grey. SIZE: wing, Y (n ¼ 142) 81–87 (837), X (n ¼ 38) 77–86 (800); tail, Y T2 (n ¼ 92) 175–228 (199), width (depth) 33–43, T3 (n ¼ 40) 60–66 (626), X (n ¼ 15) 51–61 (563); bill, Y (n ¼ 16) 100–127 (112), X (n ¼ 15) 93–100 (95); tarsus, Y (n ¼ 136) 15–19 (172), X (n ¼ 15) 15–18 (165). WEIGHT: Y (n ¼ 8) 195–260 (222), X (n ¼ 8) 178–212 (laying) (193); juv., 2 YY 176, 178, 1 X 176. IMMATURE: juv. upperparts plain grey-brown, rump grey, face nearly unmarked (with slight pale grey eye-stripe), wings and tail brownish grey, underparts pale grey, belly and undertail-coverts white, tail rounded to slightly graduated. Mouth like that of host, Orange-winged Pytilia: whitish gape ridges and light pink palate with lateral purple spots and without medial black spot. Bill black, eyes dark brown, feet grey. NESTLING: skin blackish, natal down on crown and back whitish, gape flange whitish lined with 2 black spots on inner margin above and below, back of palate whitish with large elongated violet spot at either side; no black spot on hard pinkish palate (very similar to nestling Pytilia afra: Nicolai 1964) (fig. p. 252).

Field Characters. Length Y 26–28 cm, X 13–14 cm. Breeding Y has shorter and broader tail than Eastern Paradise-Whydah V. paradisaea, central feathers deeper than body and equally wide from base to tip, nape darker, coppery rufous rather than yellow, and breast deeper

Vidua obtusa

chestnut. Early in breeding season, growing feathers are broad to tips (and grow to full length in 45–50 days: Verheyen 1953); late in season, long tail-feathers lost (T2 first), when bird distinguished from Eastern ParadiseWhydah by nape and breast. X lacks dark C mark on cheek of Eastern, and bill is pale not dark in breeding season. Non-breeding Y has grey (not black) bill and less black on cheek. Juvenile has mouth similar to young Orange-winged Pytilia in lacking black central palate spot; whydah differs in having rump grey-brown not red. Voice. Tape-recorded (99, 104, B, PAY). Songs and calls mimic those of Orange-winged Pytilia, including: song with rattling note and fluty descending whistles followed by a crackling note, and simple descending fluty whistle with buzzy overtone, ‘zheee-zhoooow’, and harsh contact call. Non-mimicry calls include soft chatter and ‘whoooeee’ in flight, loud rapid chatter when perched and loud complex chatter before flying. General Habits. Lives in dry miombo woodland and old cultivated fields, also grassland with drainage lines. In some areas occurs together with Long-tailed Paradiseˆ i, both whydahs and their Whydah V. paradisaea. In Malaw Pytilia hosts are common near Monkey Bay and Rumphi; in Nsanje district, V. obtusa is at higher elevations than V. paradisaea. In Tanzania, both occur in Mikumi Nat. Park, near Iringa, at Rukwa, Nyaviumbu and Kifuru, and Masasi; in Selous Game Res. they occupy different habitats (V. paradisaea in acacia bushland). In Zambia, the 2 are

428

VIDUIDAE

largely allopatric, V. obtusa (and Pytilia afra) in northern miombo woodlands, both in Lochinvar Nat. Park (where V. obtusa and P. afra are less common). May be migratory; some birds are extremely fat (Traylor 1965). Feeds alone, or Y with 1 or more XX, in breeding season. Near end of breeding season, occurs in flocks of 20–40 or more, often on burned ground (Payne 1971, Randall et al. 1994). Food. Grass seeds 1–2 mm in diam., taken from ground. Breeding Habits. Brood parasitic on Orange-winged Pytilia Pytilia afra (Nicolai 1969, Payne 1971). Juveniles occur in mixed-species broods. Has bred in captivity under P. afra; induced to lay in nestboxes after it hears songs of P. afra (Nicolai 1964, Baptista 1992). In courtship display, Y flies slowly and conspicuously over singing area with inner tail feathers T1 enclosed by longer T2, and does not erect T1 (as does V. paradisaea). Y perches in top of tree, mimics songs of P. afra. X whydah flies to Y; then (1) Y approaches X, his central tail feather exposed and held above drooping long tail, stretches head and body upright with legs stiffly extended, bill directed toward her; corrugated central tail feathers produce slight rustling sound; (2) he perches with body and tail presented sideways to X, bows head and neck up and down, flexing legs; bill goes below level of perch, bird in a choking posture (A); Y opens and closes bill as if courtship feeding or feeding a nestling; (3) he swings head from side to side, still holding body sideways to X, turns to face her, turns away and then to her again (Nicolai 1964, 1968d, 1969, Tweedy 1965). Y then flies to X, hovers, courts from above. EGGS: sets of 3, 1 a day, a few days between sets (Payne 1967, 1977a). Little gloss; immaculate white. SIZE: 185  14, 185  135, 175  13, 175  13 (large eggs in nests of P. afra: Belcher 1930); 182  131, 179  130 (large eggs in nest of P. afra: Vincent 1949); 176  130 (hard egg in oviduct: Payne 1967). LAYING DATES: Jan–July, from end of rains to well into dry season. S Zaı¨re (Katanga) (Mar–July, YY in breeding

plumage: Ruwet 1965; large egg in nest of P. afra: Vincent 1949); Angola (Aug–Sept, YY in breeding and eclipse plumage: Dean et al. 1988); SW Tanzania (YY in breeding plumage at end of rains and early dry season: Meise 1937); Zambia (NW, fledglings fed by ad. P. afra, mid-Sept; postjuvenile moult in late Sept, indicating laying July–Aug (Payne 1971); at Kafue in breeding plumage Feb–June (Leonard 1998); at Lochinvar Nat. Park, juvs appear in Apr–June with laying 1–2 months earlier, ad. YY moult ˆ i (eggs in P. afra nests, Apr–June from July); Malaw (Belcher 1930)); Zimbabwe (hard egg in oviduct, Mar (Payne 1977a)), YY in breeding plumage, Victoria Falls, Aug (Tree 1994); NE Transvaal (oviduct egg, Feb (Payne 1967)). DEVELOPMENT AND CARE OF YOUNG: unknown, except that young whydahs and host Pytilia afra in a mixed brood fledge together. Y in first year sometimes has incompletelydeveloped breeding plumage; develops ad. breeding plumage in 2nd breeding season. Key References Chapin, J.P. (1922, 1929a, b), Nicolai, J. (1964, 1968d, 1969), Payne, R.B. (1967, 1971, 1977a, 1997a).

Plate 26

Vidua paradisaea (Linnaeus). Long-tailed Paradise-Whydah. Veuve de paradis.

(Opp. p. 395)

Emberiza paradisaea Linnaeus, 1758. Syst. Nat., ed. 10, p. 178; Africa (¼ Angola, Linnaeus). Range and Status. Endemic resident. SE Sudan (Mongalla, Juba, Lado, Boma, Torit). Eritrea (Mai Uassen, Ailet, Salamona, Komayli, Danakil). Ethiopia (Rift Valley and south and east of northern highlands; absent Ogaden desert and E Bale). Somalia (west of 47
E). Uganda (Murchison Falls, Kidepo Valley Nat. Park, Moroto). Kenya (coastal bush to lower Tana R., Voi, Samburu, L. Baringo to Turkana, Kerio and rift valleys, Tsavo and Amboseli Nat. Parks). Tanzania (widespread in bush country; absent northeast of L. Tanganyika; few in S in miombo woodlands; present in Rovuma R. lowlands to coast at

ˆ i. Angola (Huı´la Plateau and woodlands Mikindani). Malaw of western littoral). SE Zaı¨re (L. Tanganyika: Bredo, Musosa). Zambia (between L. Mweru and L. Tanganyika; mainly in E and S including Luangwa, Kafue and Middle Zambezi valleys up to Sinjembela and Sioma). Namibia (Windhoek north to Cunene, Caprivi). Zimbabwe (throughout, sparse in E highlands). Botswana. Mozambique. Swaziland. South Africa (Natal, E and N Transvaal to bushveld north of Magaliesburg Mts). Density, in Lochinvar Nat. Park, Zambia, 2–4 displaying YY per km2 in 1972–1976, less numerous in 1997.

428

VIDUIDAE

largely allopatric, V. obtusa (and Pytilia afra) in northern miombo woodlands, both in Lochinvar Nat. Park (where V. obtusa and P. afra are less common). May be migratory; some birds are extremely fat (Traylor 1965). Feeds alone, or Y with 1 or more XX, in breeding season. Near end of breeding season, occurs in flocks of 20–40 or more, often on burned ground (Payne 1971, Randall et al. 1994). Food. Grass seeds 1–2 mm in diam., taken from ground. Breeding Habits. Brood parasitic on Orange-winged Pytilia Pytilia afra (Nicolai 1969, Payne 1971). Juveniles occur in mixed-species broods. Has bred in captivity under P. afra; induced to lay in nestboxes after it hears songs of P. afra (Nicolai 1964, Baptista 1992). In courtship display, Y flies slowly and conspicuously over singing area with inner tail feathers T1 enclosed by longer T2, and does not erect T1 (as does V. paradisaea). Y perches in top of tree, mimics songs of P. afra. X whydah flies to Y; then (1) Y approaches X, his central tail feather exposed and held above drooping long tail, stretches head and body upright with legs stiffly extended, bill directed toward her; corrugated central tail feathers produce slight rustling sound; (2) he perches with body and tail presented sideways to X, bows head and neck up and down, flexing legs; bill goes below level of perch, bird in a choking posture (A); Y opens and closes bill as if courtship feeding or feeding a nestling; (3) he swings head from side to side, still holding body sideways to X, turns to face her, turns away and then to her again (Nicolai 1964, 1968d, 1969, Tweedy 1965). Y then flies to X, hovers, courts from above. EGGS: sets of 3, 1 a day, a few days between sets (Payne 1967, 1977a). Little gloss; immaculate white. SIZE: 185  14, 185  135, 175  13, 175  13 (large eggs in nests of P. afra: Belcher 1930); 182  131, 179  130 (large eggs in nest of P. afra: Vincent 1949); 176  130 (hard egg in oviduct: Payne 1967). LAYING DATES: Jan–July, from end of rains to well into dry season. S Zaı¨re (Katanga) (Mar–July, YY in breeding

plumage: Ruwet 1965; large egg in nest of P. afra: Vincent 1949); Angola (Aug–Sept, YY in breeding and eclipse plumage: Dean et al. 1988); SW Tanzania (YY in breeding plumage at end of rains and early dry season: Meise 1937); Zambia (NW, fledglings fed by ad. P. afra, mid-Sept; postjuvenile moult in late Sept, indicating laying July–Aug (Payne 1971); at Kafue in breeding plumage Feb–June (Leonard 1998); at Lochinvar Nat. Park, juvs appear in Apr–June with laying 1–2 months earlier, ad. YY moult ˆ i (eggs in P. afra nests, Apr–June from July); Malaw (Belcher 1930)); Zimbabwe (hard egg in oviduct, Mar (Payne 1977a)), YY in breeding plumage, Victoria Falls, Aug (Tree 1994); NE Transvaal (oviduct egg, Feb (Payne 1967)). DEVELOPMENT AND CARE OF YOUNG: unknown, except that young whydahs and host Pytilia afra in a mixed brood fledge together. Y in first year sometimes has incompletelydeveloped breeding plumage; develops ad. breeding plumage in 2nd breeding season. Key References Chapin, J.P. (1922, 1929a, b), Nicolai, J. (1964, 1968d, 1969), Payne, R.B. (1967, 1971, 1977a, 1997a).

Plate 26

Vidua paradisaea (Linnaeus). Long-tailed Paradise-Whydah. Veuve de paradis.

(Opp. p. 395)

Emberiza paradisaea Linnaeus, 1758. Syst. Nat., ed. 10, p. 178; Africa (¼ Angola, Linnaeus). Range and Status. Endemic resident. SE Sudan (Mongalla, Juba, Lado, Boma, Torit). Eritrea (Mai Uassen, Ailet, Salamona, Komayli, Danakil). Ethiopia (Rift Valley and south and east of northern highlands; absent Ogaden desert and E Bale). Somalia (west of 47
E). Uganda (Murchison Falls, Kidepo Valley Nat. Park, Moroto). Kenya (coastal bush to lower Tana R., Voi, Samburu, L. Baringo to Turkana, Kerio and rift valleys, Tsavo and Amboseli Nat. Parks). Tanzania (widespread in bush country; absent northeast of L. Tanganyika; few in S in miombo woodlands; present in Rovuma R. lowlands to coast at

ˆ i. Angola (Huı´la Plateau and woodlands Mikindani). Malaw of western littoral). SE Zaı¨re (L. Tanganyika: Bredo, Musosa). Zambia (between L. Mweru and L. Tanganyika; mainly in E and S including Luangwa, Kafue and Middle Zambezi valleys up to Sinjembela and Sioma). Namibia (Windhoek north to Cunene, Caprivi). Zimbabwe (throughout, sparse in E highlands). Botswana. Mozambique. Swaziland. South Africa (Natal, E and N Transvaal to bushveld north of Magaliesburg Mts). Density, in Lochinvar Nat. Park, Zambia, 2–4 displaying YY per km2 in 1972–1976, less numerous in 1997.

Vidua paradisaea

Vidua paradisaea

NESTLING: skin black, hatchling with grey natal down on crown and back; gape flange whitish with upper and lower swellings lined with 2 black spots on inner margin above and below, palate black at tip, white with a single black spot in centre, and pink with a large elongated violet-blue spot on either side at rear (Nicolai 1964, 1974, Skead 1975, Payne 1998b); exactly like mouth of P. melba (A: P. melba on left, V. paradisaea on right). Pattern of natal down nearly identical with that of P. melba with rows across crown (10 feathers on each side), 3 feathers on wing, spinal tract with diamond of down in front and 6 downy tufts on rear femoral tract (does not have down along each side of lower abdomen as in P. melba (D.N. Mansfield, pers. comm., Markus 1970)). Feathered nestling nearly identical to P. melba except in rump colour (brown in whydah, red in pytilia); whydah larger (Nicolai 1964, 1965a, 1969, 1991).

Range falls within that of its host, Green-winged Pytilia Pytilia melba. Unknown in part of range of P. melba in lower Congo basin. Description. ADULT Y (breeding): upperparts black; tail black, 3 times length of body, T1 and T2 long, broad at base, tapered to tip, twisted longitudinally through 90
, and prominent in lateral view; T1 shorter, enclosed by T2. Edges of T2 form a zipper that binds T1 and T2 together below (Rutschke and Stresemann 1961). Nape yellow (golden when fresh, straw when faded); upper breast maroon chestnut, lower breast, belly and undertail-coverts buff. Bill black, eyes dark brown, feet blackish, dark grey or brown. ADULT Y (non-breeding): black crown-stripes and black face marks broader and more distinct than in X, head stripe whiter, less buffy, more distinctly streaked black on breast; bill grey. ADULT X: upperparts grey-brown streaked blackish, crown with buffy whitish streak bordered by blackish one, face with pale superciliary streak and vertical ‘C’ mark over ear, open towards bill, breast light grey to buffy with indistinct streaks, belly white, undertail-coverts whitish, underwing-coverts light grey; bill dark grey, lighter on base of lower mandible. SIZE: wing, Y (n ¼ 246) 74–82 (786), X (n ¼ 35) 73–79 (757); tail, Y (n ¼ 14) 53–63 (588), T2 (n ¼ 57) 243–346 (288), width (depth) 23–32, T3 (n ¼ 25) 54– 71 (636), X (n ¼ 27) 50–65 (555); bill, Y (n ¼ 14) 107–121 (113), X (n ¼ 26) 91–103 (97); tarsus, Y (n ¼ 233) 15–19 (170), X (n ¼ 26) 16–19 (168). WEIGHT: Zambia to Transvaal, Y (n ¼ 6) 197–225 (212), 1st-year (n ¼ 4) 175–194 (186); X (n ¼ 25) 172–248 (laying) (199); Namibia, Y (n ¼ 102) 185–293 (218), X (n ¼ 98) 158–247 (190); fledglings (n ¼ 16) 147–232. IMMATURE: juv. upperparts unmarked grey-brown, rump grey, face nearly unmarked (with slight paler grey eye-stripe), wings and tail brownish grey, underparts paler grey, belly and undertail-coverts white, tail rounded to slightly graduated; bill black to blackish with brown on top centre and at base of lower mandible, eyes dark brown, feet brownish grey to dark grey. Fledgling has mouth like that of Pytilia melba, with whitish gape ridges, and light pink palate with median black spot and purple-blue lateral spots.

Field Characters. Length Y 36–39 cm, X 13–14 cm. Breeding Y told from Broad-tailed Paradise-Whydah V. obtusa by longer, tapered tail, straw-yellow nape and paler chestnut breast. Arabic name Abu Mus (‘father of knife’) describes the blade-like shape of tail. X differs from other paradise-whydahs in having dark C-shaped mark on cheeks; bill dark when breeding (pale in Broad-tailed), paler when in moult (but darker than moulting Broadtailed). Juvenile unmarked grey, with black bill and white gape ridge; differs from Broad-tailed by presence of black palate spot. Nestling plumage and mouth colours similar to those of P. melba. Newly hatched whydah distinguished from pytilia by larger size, darker skin, grey-white not sandy white natal down, and broader more conical bill (pointed in host). Feathered nestling and juv. larger (wing 80 and tail 60, vs 60 and 52 in host), plumage grey-brown (vs olivegrey), rump brown (vs red), tarsus with 6 scales (vs 8), and bill shorter, deeper and lacking convex culmen of P. melba (B: P. melba on left, V. paradisaea on right).

429

430

VIDUIDAE

Voice. Tape-recorded (86, 99, 104, B, PAY). Y mimics calls and songs of its host, Green-winged Pytilia (whose songs vary regionally). Songs of particular Y vary in elements and sequence (as do songs of individual pytilia). Mimicry songs last up to 16 s, begin with sound like drop of water falling onto water, followed by long and short whistles, then gurgling and trilling, ending with 3 fluty notes; often repeated several times (all as in pytilia, Immelmann et al. 1965). Locally, birds match their mimicked songs; which differ from songs some km away. Y also gives non-mimetic call, a high, thin, sweet, hissing, ‘weeeee’, ‘weee-tssss’ or ‘seeee-tsi-tseeeew’, triple hissing call ‘sss-sss-ssssss’ or buzzier ‘zzzeeee’, sometimes in mixture of chatters and short notes. Both sexes give nonmimetic chatters similar to those of other whydah species, soft in flight, loud and rapid when perched, and loud and complex before flying.

General Habits. Inhabits woodland with scattered trees and bushes, acacia woodland and other semi-arid scrubland; in E Africa generally dry country below 1400 m; Acacia tortilis savanna in central Transvaal (Barnard 1989); Brachystegia, Baikiaea and low mopane Colophospermum mopane woodland, and lightly cultivated habitats derived from these woodlands. Feeds in grassy areas and along roadsides. Normal flight rapid, with deeper wingbeats than in display flight; in tail, T1 enclosed within long T2. Display flight and song occur mainly in late morning and late afternoon; territorial behaviour ceases in evening when breeding birds gather to feed and roost (C). Occurs in flocks of >100 with other Vidua species (mainly V. chalybeata) at Lochinvar Nat. Park in July–Aug, in large mixed roosts in dense leafy trees (pers. obs.). Flocks at end of breeding season also in Zimbabwe. Throughout range, occurs with grey-lored races of Green-winged Pytilia; but in S and E Sudan, Ethiopia and N and E Kenya occurs with red-lored P. m. somaliensis.

Food. Grass seeds 1–2 mm in diam., taken from ground. Laying X eats host eggs. Feeding on seeds is most intense just before flying to roost at dusk; 20 min before sunset a laying X had >400 small grass seeds in crop.

Breeding Habits. Brood parasitic on Pytilia melba. X visits nests of pytilia and occasionally other species, e.g. Violeteared Waxbill Granatina granatina (Skead 1975). Proportion of nests parasitized varies and is sometimes high: in Transvaal, 28% of 75 P. melba nests were parasitized; in ˆ i, 47% of 15 nests, in Tanzania, 95% of 36. Often Malaw more than 1 whydah egg or chick in nest (in Transvaal, 11 nests had 1, 7 had 2, and 3 had 3 whydah eggs; in Tanzania, 9 had 1, 18 had 2, 5 had 3, 1 had 4 and 1 had 5). Nests with more than 1 whydah egg sometimes involve more than 1 X; in one nest 3 whydah eggs were laid in 1 day. Both whydah and host young fledge from parasitized nests. Occasionally lays in nest of other species and young are given parental care even when mouth colours differ: in Tanzania a nestling V. paradisaea and a nestling V. fischeri were in brood with 2 Purple Grenadiers G. ianthinogaster (Nicolai 1969). Courtship behaviour: Y gives aerial displays twice a day for c. 20 min, around 10h00 and 16h00 (Kenya, Tanzania, Zambia and Transvaal). During those periods Y hardly feeds at all; YY display within sight of each other each over separate display territories, of c.3 ha. Display arena has trees and shrubs that Y uses as song perches and to launch his display flights. In aerial display, Y flies up from tree at angle of 30–60
, then flies slowly with rapid wingbeats at height of 20–30 m, circling an area of 50– 100 m radius in horizontal flight (D); cruises slowly across territory, giving whistled ‘whee-hew’s, with central pair of tail-feathers (T1) held upright, vertical to axis of bird, and the long T2 trailing behind. After 20 s or so he folds wings to sides then flaps again, descending jerkily, then plunges head-first, with T1 concealed by T2, and perches in another tree or bush. There he gives a long, sizzling, warbling song, of 10–12 s, that mimics song of P. melba (Nicolai 1969, Payne 1997a). If no X approaches, he flies up at sharp angle, rustling tail, ascends to 20–30 m, and repeats slow flight display and ‘whee-hew’ call. Tail produces quiet rustling sound, surfaces of T1 and T2

Vidua hypocherina

being corrugated (Koenig 1962). 1st year Y in sparrowy plumage lifts central T1 in display like ad. Y. Mating occurs when X visits singing Y at his site. X approaches and perches, Y bows head and neck low over perch towards her, tosses head up and over back, bill tilted and mouth alternately opened and closed, breast thrust forward (E); he holds this posture, then arches head high and raises long tail feathers vertically, so that body and tail form a ‘U’; he raises his tail, exposes central T1, while long T2 feathers arch behind, and turns to present first one side then the other to X, then bows. He swings head from side to side, flies to her, hovers over her, and copulates (Nicolai 1964, 1969). During display period, if intruding Y appears overhead, resident Y accompanies him away, flying low and keeping between intruder and song trees.

431

X perches in top of bush, often for 15 min or longer, looking for pytilia nesting activity, then flies to investigate, not accompanied by Y. X lays any time from morning to early afternoon (Payne 1977a). EGGS: sets of 3–4, laid 1 a day, a few days between sets. X lays av. 22 in a season. Little gloss; immaculate white. SIZE: (n ¼ 5, hard eggs in oviduct) 173–184  120–141 (177  129) (Payne 1977a); (n ¼ 2) 19  13, 183  124 (Belcher 1930). LAYING DATES: Ethiopia, May–June; Kenya, June–July; Tanzania, (Iringa, breeds Feb–Mar, N Pare Mts, Apr–May, fledgling in July); Zambia, (Lochinvar Nat. Park, YY displaying and courting Mar–May; oviduct egg May; ˆ i, Feb–June (D.N. Mansfield, pers. comm.); S Malaw Mozambique, (YY in breeding plumage until May–June; Zimbabwe, (oviduct eggs Mar–Apr (Payne 1977a), YY in breeding plumage until Aug); Botswana, (fledged young, May); W Angola, (YY in breeding plumage from July); N Transvaal, (oviduct eggs, Feb–Mar); southern Africa, Jan– June. INCUBATION: period (n ¼ 3) 11 days (2 days shorter than host’s). DEVELOPMENT AND CARE OF YOUNG: from hatching, nestling larger than P. melba nestling, and bill form differs; skin, natal down and gape colours nearly identical (Nicolai 1965a). Whydahs that hatch 2 days later than host nestlings do not survive (D.N. Mansfield, pers. comm.). In first 8–10 days, foster parents feed insects to young (Nicolai 1969); young are also fed seeds throughout nestling period. Begging call at first like that of nestling P. melba; later it becomes more distinct, a husky, harsh ‘chew chew chew’ (P. melba: ‘chuc, chuc, pett’). Nestling period: (in captivity, reared by Bengalese Finch Lonchura striata) 16 days (Nicolai 1969). Whydahs and pytilias leave nest together. Young beg just like host young, by crouching and twisting neck around to face foster parent, who regurgitates food. Independent 27–30 days after hatching; begin moulting 1–2 months after fledging. 1st year Y sometimes has incomplete breeding plumage; ad. breeding plumage develops in 2nd breeding season. Key References Barnard, P. (1995), Friedmann, H. (1960), Nicolai, J. (1964, 1965a, b, 1968d, 1969, 1974, 1977, 1991), Payne, R.B. (1971, 1973b, 1977a, 1998b).

Vidua hypocherina J. and E. Verreaux. Steel-blue Whydah. Veuve me ´tallique.

Plate 25

Vidua hypocherina J. and E. Verreaux, 1856. Rev. Mag. Zool., p. 260, pl. 165; ‘West Africa’.

(Opp. p. 394)

Range and Status. Endemic resident, E Africa. Sudan (east of Kapoeta). NE Uganda (Kidepo Valley Nat. Park). Ethiopia (SE and S, Gallaland, Boran and Harar). Somalia, in NW Hargesia and Ethiopian border and in S, mainly along Jubba valley. Kenya (Wajir, Marsabit, Tana R. to Lamu, Samburu Nat. Park, Turkwel R., L. Baringo, L. Bogoria, Nyanza, rift areas from Mara to Olorgesailie and Magadi, Tsavo West Nat. Park). Tanzania (dry central plateau from Serengeti, Arusha, L. Manyara, Tarangire

Nat. Park, Mkomazi in Usambara and Masai Steppe south to Tabora, Morogoro, Mikumi Nat. Park and Usangu Flats). Breeding range is within ranges of main host species, Black-faced Waxbill Estrilda charmosyna and Blackcheeked Waxbill E. erythronotos. Description. ADULT Y (breeding): glossy blue-black; central tailfeathers (T1,T2) black, narrow (5–6 mm near tip, where broader than near base) and longer than the rest of bird; T3–T6 black with buff-white margin on inner webs; wing dark brown, with broad

Vidua hypocherina

being corrugated (Koenig 1962). 1st year Y in sparrowy plumage lifts central T1 in display like ad. Y. Mating occurs when X visits singing Y at his site. X approaches and perches, Y bows head and neck low over perch towards her, tosses head up and over back, bill tilted and mouth alternately opened and closed, breast thrust forward (E); he holds this posture, then arches head high and raises long tail feathers vertically, so that body and tail form a ‘U’; he raises his tail, exposes central T1, while long T2 feathers arch behind, and turns to present first one side then the other to X, then bows. He swings head from side to side, flies to her, hovers over her, and copulates (Nicolai 1964, 1969). During display period, if intruding Y appears overhead, resident Y accompanies him away, flying low and keeping between intruder and song trees.

431

X perches in top of bush, often for 15 min or longer, looking for pytilia nesting activity, then flies to investigate, not accompanied by Y. X lays any time from morning to early afternoon (Payne 1977a). EGGS: sets of 3–4, laid 1 a day, a few days between sets. X lays av. 22 in a season. Little gloss; immaculate white. SIZE: (n ¼ 5, hard eggs in oviduct) 173–184  120–141 (177  129) (Payne 1977a); (n ¼ 2) 19  13, 183  124 (Belcher 1930). LAYING DATES: Ethiopia, May–June; Kenya, June–July; Tanzania, (Iringa, breeds Feb–Mar, N Pare Mts, Apr–May, fledgling in July); Zambia, (Lochinvar Nat. Park, YY displaying and courting Mar–May; oviduct egg May; ˆ i, Feb–June (D.N. Mansfield, pers. comm.); S Malaw Mozambique, (YY in breeding plumage until May–June; Zimbabwe, (oviduct eggs Mar–Apr (Payne 1977a), YY in breeding plumage until Aug); Botswana, (fledged young, May); W Angola, (YY in breeding plumage from July); N Transvaal, (oviduct eggs, Feb–Mar); southern Africa, Jan– June. INCUBATION: period (n ¼ 3) 11 days (2 days shorter than host’s). DEVELOPMENT AND CARE OF YOUNG: from hatching, nestling larger than P. melba nestling, and bill form differs; skin, natal down and gape colours nearly identical (Nicolai 1965a). Whydahs that hatch 2 days later than host nestlings do not survive (D.N. Mansfield, pers. comm.). In first 8–10 days, foster parents feed insects to young (Nicolai 1969); young are also fed seeds throughout nestling period. Begging call at first like that of nestling P. melba; later it becomes more distinct, a husky, harsh ‘chew chew chew’ (P. melba: ‘chuc, chuc, pett’). Nestling period: (in captivity, reared by Bengalese Finch Lonchura striata) 16 days (Nicolai 1969). Whydahs and pytilias leave nest together. Young beg just like host young, by crouching and twisting neck around to face foster parent, who regurgitates food. Independent 27–30 days after hatching; begin moulting 1–2 months after fledging. 1st year Y sometimes has incomplete breeding plumage; ad. breeding plumage develops in 2nd breeding season. Key References Barnard, P. (1995), Friedmann, H. (1960), Nicolai, J. (1964, 1965a, b, 1968d, 1969, 1974, 1977, 1991), Payne, R.B. (1971, 1973b, 1977a, 1998b).

Vidua hypocherina J. and E. Verreaux. Steel-blue Whydah. Veuve me ´tallique.

Plate 25

Vidua hypocherina J. and E. Verreaux, 1856. Rev. Mag. Zool., p. 260, pl. 165; ‘West Africa’.

(Opp. p. 394)

Range and Status. Endemic resident, E Africa. Sudan (east of Kapoeta). NE Uganda (Kidepo Valley Nat. Park). Ethiopia (SE and S, Gallaland, Boran and Harar). Somalia, in NW Hargesia and Ethiopian border and in S, mainly along Jubba valley. Kenya (Wajir, Marsabit, Tana R. to Lamu, Samburu Nat. Park, Turkwel R., L. Baringo, L. Bogoria, Nyanza, rift areas from Mara to Olorgesailie and Magadi, Tsavo West Nat. Park). Tanzania (dry central plateau from Serengeti, Arusha, L. Manyara, Tarangire

Nat. Park, Mkomazi in Usambara and Masai Steppe south to Tabora, Morogoro, Mikumi Nat. Park and Usangu Flats). Breeding range is within ranges of main host species, Black-faced Waxbill Estrilda charmosyna and Blackcheeked Waxbill E. erythronotos. Description. ADULT Y (breeding): glossy blue-black; central tailfeathers (T1,T2) black, narrow (5–6 mm near tip, where broader than near base) and longer than the rest of bird; T3–T6 black with buff-white margin on inner webs; wing dark brown, with broad

432

VIDUIDAE

Vidua hypocherina

white band formed by inner webs of primaries and secondaries, underwing-coverts white; upper flank with white spot, normally concealed by back feathers. Bill short and stubby, white (but in museum specimens red, grey or black), eyes dark brown, feet grey. ADULT Y (non-breeding): similar to X, head markings more distinctly blackish and white; bill light grey or horn colour, feet grey. ADULT X: upperparts grey-brown with black streaks, crown dark brown with whitish central streak and whitish superciliary streak, tail dark with white trim formed by white edge on inside and tips of feathers and narrower white edge on outer margin (300 m from other YY, and feeds in territory. Occurs singly or in flocks, often with other species. Scratches bare ground rapidly with both feet, hops backwards, then pecks up the exposed seed. Visits waterholes to drink. Food. Grass seeds 1–2 mm in diam., taken from ground. Breeding Habits. Polygynous, territorial. Brood parasite of Black-faced and Black-cheeked Waxbills. Parasitized waxbill nest has 1 or 2 whydah eggs (Nicolai 1989). X once visited nest of Bronze Mannikin Spermestes cucullatus (van Someren 1956). Usually does not mimic songs and calls of waxbills, but known to mimic voices of Red-cheeked Cordon-bleu (W Kenya, though not known to parasitize it: Payne 1997a). Courting Y sings from a high perch. When X visits, he flies to ground, where he sings; when she joins him, he makes a quick jump over her as though pricked by a pin, and flutters his wings and bobs tail in a tight circling flight, for up to 30 s. He repeats display or forages with her, then leads her in flight back to his perch. EGGS: sets of 3–4, laid 1 a day, a few days between sets (Payne 1977a). Little gloss; immaculate white. SIZE: c. 171  121, larger than eggs of waxbill host (Nicolai 1989).

Vidua fischeri LAYING DATES: S Sudan, (Y in breeding plumage, May); S Somalia, (fledglings, Sept); Kenya, (L. Baringo, June– Sept; YY in breeding plumage, May–Sept; Kisumu, June– July, S Kenya, May–June). DEVELOPMENT AND CARE OF YOUNG: nestling period, c. 20 days. Foster parent waxbills feed half-ripe seeds and insects to nestlings. By day 12 nestlings are in pinfeathers, by day 16 they are well feathered, by day 28 the

433

fledged young are independent of their foster parents (Nicolai 1989, 1990). Y develops breeding plumage or partial breeding plumage when a year old (Y with skull 40% pneumatized is only 10% black). Key References Cunningham-van Someren, G.R. (1974, 1978), Friedmann, H. (1960), Neunzig, R. (1929), Nicolai, J. (1964, 1989, 1990), Payne, R.B. (1977a, 1997a, b).

Vidua fischeri (Reichenow). Straw-tailed Whydah. Veuve de Fischer.

Plate 25

Linura fischeri Reichenow, 1882. Orn. Centralbl., 7, p. 91; Usegua [Tanzania].

(Opp. p. 394)

Range and Status. Endemic resident, E Africa. Sudan (south of 7
N: E side of Dongotona Mts, E side of Didinga Mts, Kapoeta and Kenyan border). Ethiopia (S Rift Valley at L. Stephanie, Wobok, Tertale, Owisa, also Dire Dawa, NE Harar and Jubba R.). Somalia (in N Hargesia and Ethiopian border, and in S mainly in lower Shabeelle R. and Jubba R. areas). NE Uganda (Kidepo Valley Nat. Park). Kenya (Marsabit, Wajir, Garissa, Kongelai escarpment, Kerio Valley, L. Baringo, Samburu Nat. Park, Loita plains, Olorgesailie, Magadi, Namanga, Amboseli Nat. Park, Tsavo Nat. Parks, Voi, Winam Gulf on L. Victoria). Tanzania (dry central plateau from Serengeti and Masai steppe south to Tabora, Morogoro, Mikumi Nat. Park and Iringa region). Breeding range is within the range of its host species, Purple Grenadier Granatina ianthinogaster. Density, c. 3 singing YY per km2 at L. Baringo (Payne et al. 2000b). Description. ADULT Y (breeding): bright yellow forehead and crown, rest of upperparts black; tail short, brown (tips and edges buff when fresh), except for T1 and T2, which are narrow, strawcoloured and longer than rest of bird (usually >200 long and 2–3 wide with fine barbs along shafts); face and chin to upper breast black, rest of underparts yellowish buff; wings brown, underwingcoverts dark grey. Bill orange to coral red, eyes dark brown, feet orange. ADULT Y (non-breeding): similar to X, but head more boldly marked. ADULT X: upperparts streaky brown, head nearly unstreaked dark rufous to pale chestnut, wearing and fading to rust or buff, nape streaked, rump unstreaked, tail brown (tips and edges buff when fresh); face pale buff, plain, without distinct superciliary stripe or eye-stripe, nearly same colour as breast, throat whitish, breast sandy buff, belly whitish, wings brown, underwing-coverts brownish grey. SIZE: wing, Y (n ¼ 34) 64–70 (673), X (n ¼ 16) 63–67 (650); tail, Y (n ¼ 34) 40–48 (424), T1, T2 (n ¼ 21) 182–260 (213), X (n ¼ 18) 36–44 (399); bill, Y (n ¼ 33) 78–101 (85), X (n ¼ 18) 76–99 (85); tarsus, Y (n ¼ 34) 14–17 (156), X (n ¼ 18) 14–16 (144). WEIGHT: Kenya, Y (n ¼ 13) 100–178 (132), X (n ¼ 11) 90–179 (113). IMMATURE: juv. rusty brown, unmarked or with indistinct streaks on back, rump rusty brown, tail brown; bill black, eyes dark brown, feet brown; bill and feet change to orange when bird begins to moult into streaked plumage. NESTLING: skin purplish black, natal down grey; palate centre whitish with 3 black spots, tip of palate dark grey, behind the spots orange, grading to whitish and pale blue on sides, rest of palate black, oral surface of gape black. Dorsal gape swelling large and blue, ventral one small and pale blue, band around gape dark blue to purplish black. Tongue black with white edges, lower mandible black. Mouth pattern similar to that of host nestling, Purple Grenadier, but in 1st days of life whydah has round black

Vidua fischeri

palate spot (lateral ellipse in host); by day 12, feathered whydah has rufous-brown rump and uppertail-coverts (blue in host) (Neunzig 1929, Nicolai 1969).

Field Characters. Length Y 30–32 cm, X 10–11 cm. Breeding Y unmistakable, black with yellow-buff crown and belly, orange bill and feet, and long, thin, strawcoloured tail. X and non-breeding Y differ from Redbilled Quelea Quelea quelea in orange-red, not pinkish red, bill and feet, unstreaked head with rufous wash, brown (not yellow) wing-edgings. Juvenile plain rusty brown, paler below, similar to Purple Grenadier but rump and tail brown (not rump blue and tail black). Voice. Tape-recorded (B, McVIC, PAY, PEA, STJ). Mimics songs and calls of Purple Grenadier: (1) loud song, series of short notes ending with 4–5 high, rising whistles, ‘cheerer cheet tsee-tsee sur-chit-cheet-chit-tsereea’; the long whistle ‘tsereea’ glides up or down in pitch, varying regionally; (2) trill song, introduced by whispers,

Vidua fischeri LAYING DATES: S Sudan, (Y in breeding plumage, May); S Somalia, (fledglings, Sept); Kenya, (L. Baringo, June– Sept; YY in breeding plumage, May–Sept; Kisumu, June– July, S Kenya, May–June). DEVELOPMENT AND CARE OF YOUNG: nestling period, c. 20 days. Foster parent waxbills feed half-ripe seeds and insects to nestlings. By day 12 nestlings are in pinfeathers, by day 16 they are well feathered, by day 28 the

433

fledged young are independent of their foster parents (Nicolai 1989, 1990). Y develops breeding plumage or partial breeding plumage when a year old (Y with skull 40% pneumatized is only 10% black). Key References Cunningham-van Someren, G.R. (1974, 1978), Friedmann, H. (1960), Neunzig, R. (1929), Nicolai, J. (1964, 1989, 1990), Payne, R.B. (1977a, 1997a, b).

Vidua fischeri (Reichenow). Straw-tailed Whydah. Veuve de Fischer.

Plate 25

Linura fischeri Reichenow, 1882. Orn. Centralbl., 7, p. 91; Usegua [Tanzania].

(Opp. p. 394)

Range and Status. Endemic resident, E Africa. Sudan (south of 7
N: E side of Dongotona Mts, E side of Didinga Mts, Kapoeta and Kenyan border). Ethiopia (S Rift Valley at L. Stephanie, Wobok, Tertale, Owisa, also Dire Dawa, NE Harar and Jubba R.). Somalia (in N Hargesia and Ethiopian border, and in S mainly in lower Shabeelle R. and Jubba R. areas). NE Uganda (Kidepo Valley Nat. Park). Kenya (Marsabit, Wajir, Garissa, Kongelai escarpment, Kerio Valley, L. Baringo, Samburu Nat. Park, Loita plains, Olorgesailie, Magadi, Namanga, Amboseli Nat. Park, Tsavo Nat. Parks, Voi, Winam Gulf on L. Victoria). Tanzania (dry central plateau from Serengeti and Masai steppe south to Tabora, Morogoro, Mikumi Nat. Park and Iringa region). Breeding range is within the range of its host species, Purple Grenadier Granatina ianthinogaster. Density, c. 3 singing YY per km2 at L. Baringo (Payne et al. 2000b). Description. ADULT Y (breeding): bright yellow forehead and crown, rest of upperparts black; tail short, brown (tips and edges buff when fresh), except for T1 and T2, which are narrow, strawcoloured and longer than rest of bird (usually >200 long and 2–3 wide with fine barbs along shafts); face and chin to upper breast black, rest of underparts yellowish buff; wings brown, underwingcoverts dark grey. Bill orange to coral red, eyes dark brown, feet orange. ADULT Y (non-breeding): similar to X, but head more boldly marked. ADULT X: upperparts streaky brown, head nearly unstreaked dark rufous to pale chestnut, wearing and fading to rust or buff, nape streaked, rump unstreaked, tail brown (tips and edges buff when fresh); face pale buff, plain, without distinct superciliary stripe or eye-stripe, nearly same colour as breast, throat whitish, breast sandy buff, belly whitish, wings brown, underwing-coverts brownish grey. SIZE: wing, Y (n ¼ 34) 64–70 (673), X (n ¼ 16) 63–67 (650); tail, Y (n ¼ 34) 40–48 (424), T1, T2 (n ¼ 21) 182–260 (213), X (n ¼ 18) 36–44 (399); bill, Y (n ¼ 33) 78–101 (85), X (n ¼ 18) 76–99 (85); tarsus, Y (n ¼ 34) 14–17 (156), X (n ¼ 18) 14–16 (144). WEIGHT: Kenya, Y (n ¼ 13) 100–178 (132), X (n ¼ 11) 90–179 (113). IMMATURE: juv. rusty brown, unmarked or with indistinct streaks on back, rump rusty brown, tail brown; bill black, eyes dark brown, feet brown; bill and feet change to orange when bird begins to moult into streaked plumage. NESTLING: skin purplish black, natal down grey; palate centre whitish with 3 black spots, tip of palate dark grey, behind the spots orange, grading to whitish and pale blue on sides, rest of palate black, oral surface of gape black. Dorsal gape swelling large and blue, ventral one small and pale blue, band around gape dark blue to purplish black. Tongue black with white edges, lower mandible black. Mouth pattern similar to that of host nestling, Purple Grenadier, but in 1st days of life whydah has round black

Vidua fischeri

palate spot (lateral ellipse in host); by day 12, feathered whydah has rufous-brown rump and uppertail-coverts (blue in host) (Neunzig 1929, Nicolai 1969).

Field Characters. Length Y 30–32 cm, X 10–11 cm. Breeding Y unmistakable, black with yellow-buff crown and belly, orange bill and feet, and long, thin, strawcoloured tail. X and non-breeding Y differ from Redbilled Quelea Quelea quelea in orange-red, not pinkish red, bill and feet, unstreaked head with rufous wash, brown (not yellow) wing-edgings. Juvenile plain rusty brown, paler below, similar to Purple Grenadier but rump and tail brown (not rump blue and tail black). Voice. Tape-recorded (B, McVIC, PAY, PEA, STJ). Mimics songs and calls of Purple Grenadier: (1) loud song, series of short notes ending with 4–5 high, rising whistles, ‘cheerer cheet tsee-tsee sur-chit-cheet-chit-tsereea’; the long whistle ‘tsereea’ glides up or down in pitch, varying regionally; (2) trill song, introduced by whispers,

434

VIDUIDAE

has short trills in middle, and ends with a short trill or single whistle on one pitch; (3) contact call, thin ‘wis-wis’; (4) contact trills (c. 10 notes per s); (5) excitement trills ‘chay-chay-chay’; (6) sharp ‘tsek’ of alarm, with notes often run together in a chatter; and nest and begging calls. Y gives medley of these songs and short chatters as he displays to X when she visits his call site, also when he is alone. Non-mimetic song is different, a variable repertoire of chatters, whistles and harsh notes. This song alternates with bouts of mimicry, and is given when one Y chases another, or when one Y gives wing-whir or ‘hover-hold’ display to X. Non-mimetic calls include chatters, rapid (30 notes per s) or slow (15–16 notes per s), and buzzy ‘chuz’ given singly or in series. Each Y has 3–4 variations of loud song (1); songs of nearby YY are almost identical, but populations 2–4 km apart have different songs, and songs differ regionally. In a given area songs change slightly over 12 years (Payne et al. 2000b). The chatter is innate; songs are not but are copied from other whydahs rather than from their hosts. General Habits. Inhabits short-grass plains habitat with patches of open ground in arid and semi-arid dry thorn scrub and bush country, below 1600 m. Y sings on display area, using several large bushes or small trees; out of sight, or several YY within sight and sound of each other. He sings from sunrise to sunset, through heat of mid-day, on a bush for up to 60 min at a time. Territorial; defends song perch as a mating site; feeds in territory. In non-breeding season, occurs in flocks of its own species and with other species. In all seasons, YY and XX occur with Purple Grenadiers. Food. Grass seeds 1–2 mm in diam., larval and ad. insects, all taken from bare ground (Archer and Godman 1961). Breeding Habits. Polygynous, territorial. Brood parasite of Purple Grenadier (Nicolai 1969, 1973a, Payne et al. 2000b);

perhaps also of Blue-capped Cordon-bleu Uraeginthus cyanocephalus (Nikolaus 1979, 1987). Y sings in top of a bush, usually out of sight of other YY, 100 m or more away, in display area of c. 2500 m2 in open habitat, and flies from bush to bush with tail jerking. X visits Y, he displays and takes her to ground where they feed, then go to a call-site bush to copulate. Courtship display: Y sings on perch, beats his wings, holding perch with extended legs as he hovers for up to a min, looking around. When X approaches, Y chatters then directs ‘hover-hold’ display to her, bows his head, showing his yellow cap and breast. If she stays on perch, Y flies to her and dances with hovering wings over her, bobbing up and down, quivering and flopping long tail. X may crouch and quiver wings, Y mounts; more often, Y hovers then flies to ground and sings, she joins him, and they forage while Y continues to sing. X then flies to sample the behaviour of another Y. X copulates any time up to sunset, and even that late in the day Y sings to attract additional mates. EGGS: sets of 3, laid 1 a day, a few days between sets. Little gloss; immaculate white. SIZE: (n ¼ 2) both 155  125. LAYING DATES: S Sudan, (YY in breeding condition, Dec, May); Ethiopia, (YY in breeding plumage, Apr–June, Dec; juvs. Aug–Nov); Somalia, (YY in breeding plumage, May– Dec); S Kenya, Feb–June, W Kenya, (copulation Aug– Sept); Tanzania, Mar–June. DEVELOPMENT AND CARE OF YOUNG: nestling period, c. 18 days. When reared in captivity by Bengalese Finch Lonchura striata, young grow up with host young and fledge at 16 days. Begging call of nestlings like that of host young. Chatter calls start at 32–35 days of age. Y is in breeding plumage and sings when 1 year old (Nicolai 1969, 1973a, b). Key References Cunningham-van Someren, G.R. (1973), Neunzig, R. (1929), Nicolai, J. (1964, 1968c, d, 1969, 1973a, b), Payne, R.B. (1977a, 1997a), Payne, R.B. et al. (2000b, 2003).

Plate 25

Vidua regia (Linnaeus). Shaft-tailed Whydah. Veuve royale.

(Opp. p. 394)

Emberiza regia Linnaeus, 1766. Syst. Nat., 12th ed., 1, p. 313; (S Angola). Range and Status. Endemic resident. S Angola (SW Namibe, S and central Huı´la, Cuando Cubango at Cuangar and Guma). Zambia (Western Prov. from Senanga to Chunga Pools, Katombora, Livingstone). Zimbabwe. Botswana. Namibia (widespread except coastal 120 km strip wide, south of 24
S). South Africa (N, E and W Transvaal, N and W Free State, N and NW Cape Prov.). S Mozambique (Inhambane district, and border of Kruger Nat. Park). Breeding range is within range of its host species, Violeteared Waxbill Granatina granatina. Numbers undergo annual fluctuations; common some years and nearly missing in others (e.g. Kalahari Nat. Park in South Africa and Namibia, and Springbok flats in E Transvaal). Population density, only 1 call-site in area of 12 km2, although in some areas several YY each use nearby trees (Barnard 1989).

Description. ADULT Y (breeding): forehead and crown black, xcollar rusty, mantle to rump black; tail short, pale brown, 4 central feathers black, long and slender, with shafts 20 km2 in extent, as in Lochinvar Nat. Park, Zambia; YY a few km further away have different songs. Y sings 10–50 songs in 10 min, may sing continuously for an hour; usually begins in early morning and may continue through the day and until 30 min before sunset. General Habits. Widespread in drier brushy woodlands, thorn bush, and riverine habitats where their host species, Jameson’s Firefinch, occurs (Payne 1973a; Payne et al. 1993). Less tolerant of heavy grazing by cattle than are Village Indigobird V. chalybeata and its host Red-billed Firefinch L. senegala; tends to be in denser bushland and taller grass than Village Indigobird (Payne 1987). Territorial. Y defends his song perch from own species and other species of Vidua as a mating site. Forages on ground, Y sometimes on territory. Moves on ground by hopping; feeding bout lasts up to 20 min; scratches with both feet nearly simultaneously to uncover seeds on bare earth, hops backwards, then pecks up exposed seeds. Husks grass seed in bill, rolls it forward and back with tongue against ridge of palate; gathers hundreds of seeds in crop in a feeding bout. Takes grass seeds piled at entrance burrows of small Pheidole ants, and small termites during emergences. Roosts in dense trees in mixed-species flocks in breeding season and especially in dry season. Food. Grass seeds 1–2 mm in diam.; at Lochinvar Nat. Park, Zambia, seeds of annual grasses Echinochloa colonum, Setaria spp., Urochloris and Dactycynodon. Ants and termites. Breeding Habits. Brood parasitic. Polygynous, territorial. Lays in nests of Jameson’s Firefinch L. rhodopareia, which rear young with their own. Y sings in morning or all day, courts visiting X with aerial hover in front of her, mounts, sings mimicry songs while pair is in cloacal contact, then Y flies to ground near his call-site, continuing to sing, X follows and they feed there together. X then flies to another Y and is courted by each of several YY in turn. Y attracts as many as 6 XX in a day and copulates with 3 of them. EGGS: sets of 3–4, laid 1 a day, a few days between sets. Little gloss; immaculate white. More rounded at both ends and with larger pores than firefinch eggs (ColebrookRobjent 1977). SIZE: (n ¼ 1) 154  120, (3 oviduct eggs) 15  125 (Payne 1977a). LAYING DATES: based mainly on XX with oviduct eggs: N Kenya, June; Zambia, (Choma), Mar, (Lochinvar Nat. Park), Feb–May (Payne 1977a, 1985b, 1987); Botswana, Apr; Zimbabwe, Jan–Apr; Transvaal, Jan–Mar. In

449

450

VIDUIDAE

Zimbabwe, (Chirundu, Victoria Falls) some birds in breeding plumage as late as Aug and Oct (Tree 1994). DEVELOPMENT AND CARE OF YOUNG: nestling period unknown. Fledged young occur in family groups of Jameson’s Firefinches, indicating that parasite and host young are reared together.

Plate 27 (Opp. p. 410)

BREEDING SUCCESS/SURVIVAL: 2 colour-ringed YY (Lochinvar Nat. Park, Zambia) sang on same call-site in breeding season for 4 successive years.

Key References Payne, R.B. (1968a, 1973a, 1977a, 1982, 1985b, 1987, 1994a, 1996), Payne, R.B. et al. (1993).

Vidua codringtoni (Neave). Zambezi Indigobird; Peters’s Twinspot Indigobird. Combassou de Codrington. Hypochera codringtoni Neave, 1907. Mem. Manchester Lit. Phil. Soc., 51; Molilo’s, Loangwa Valley, northeast Rhodesia [¼ Zambia]. (V. funerea lusituensis Payne 1973 is a synonym). Range and Status. Endemic resident. S Tanzania (Mikindani, Iringa, L. Rukwa). Zambia (Lusaka, Chilanga, ˆ i (Zomba, Lochinvar Nat. Park, Luangwa Valley). Malaw Namadzi, Limbe, Mt Mulanje at Likulezi Mission, Lengwe Nat. Park, Chididi hills). Zimbabwe (Gwaai R., Kwekwe, Kadoma, Penhalonga, Mutare, Rusitu R. at 380 m, Chipinge). Range is within that of presumptive host species, Peters’s Twinspot Hypargos niveoguttatus. Description. ADULT Y (breeding): black with bright blue or green gloss, tail black, concealed flank spot white; wing with inner 3 secondaries (tertials) and their coverts glossy black, other secondaries and primaries and upperwing-coverts black, underwingcoverts light grey (sometimes some coverts black). Bill white, eyes dark brown, feet bright orange or red. ADULT Y (non-breeding): like X but more strongly marked. ADULT X: upperparts brown with blackish shaft streaks, crown with central pale brown stripe streaked black, bordered by dark brown stripe, rump unstreaked brown, tail dark brown, nearly black, without white border; face pale with whitish supraorbital stripe and dark stripe from eye to lighter brown unstreaked nape, ear-spot blackish and indistinct; chin and throat whitish, breast unstreaked grey, sides of breast and flanks buffy grey, undertail-coverts white, wings dark brown with buffy wing-bars formed by upperwing-coverts, underwing-coverts grey. Bill whitish to grey or grey-brown, upper mandible darker than lower; eyes dark; feet orange, duller in non-breeding season. ˆ i, Zimbabwe) wing, Y (n ¼ 31) 66–70 (675), X (n ¼ 4) SIZE: (Malaw 65–66 (655); tail, Y (n ¼ 30) 37–41 (391), X (n ¼ 4) 39–41 (398); bill, Y (n ¼ 29) 81–100 (89), X (n ¼ 4) 80–89 (86); tarsus, Y ˆ i, (n ¼ 31) 142–160 (146), X (n ¼ 4) 14 (140). WEIGHT: Malaw Zimbabwe, Y (n ¼ 29) 114–141 (128), X (n ¼ 4) 123–133 (129). IMMATURE: juv. upperparts less streaked than in ad. X, crown nearly uniformly grey-brown (streaked in ad.), face with indistinct buffy supraorbital stripe, pattern less distinct than in ad. X, breast buffy grey, belly and undertail-coverts whitish. Mouth yellow with 3 black palate spots and without white gape papillae and blue base. NESTLING: unknown.

Field Characters. Length 10–11 cm. Breeding Y black with blue or green gloss, white bill, orange or red legs and feet; told from sympatric species by much brighter gloss, black (not brown) wings. Also identified by imitations of Peters’s Twinspot. X has whitish bill, orange feet and more distinct separation of grey breast and white belly than other indigobirds. Voice. Tape-recorded (86, 102, 103, PAY, STJ). Song churring and scratchy as in other indigobirds, mixed with

Vidua codringtoni

imitations of songs and calls of Peters’s Twinspot. Each Y has repertoire of c. 20 song types, a third of which are mimetic. Slow and fast chatters similar to those used in songs are given in aggressive contexts, interspersed with complex song. YY within a few km of each other match all their song types; YY a few km away have different sets of songs. Most distinctive and frequently heard mimicked twinspot vocalization is alarm trill ‘trrrrrrrreee’, higher in pitch (7–8 kHz) than firefinch alarm trills (4 kHz). Also mimicked are: (1) simple song, high ‘treeee’ followed by lower whistles and rapid chatter; (2) long, complex, soft song (used in twinspot sexual display) with whistles, trills and chatters; (3) short excitement calls, ‘sip’ and ‘tsisi’; and (4) begging calls, which resemble those of young firefinches (Payne et al. 1992, 1993, Payne and Payne 2001). Occasional birds imitate hosts other than twinspots – one in Zimbabwe mimicked African Firefinch Lagonosticta rubricata and matched its songs with local Dusky Indigobirds V. funerea; another imitated Jameson’s Firefinch L. rhodopareia. A Y in South Africa (KwaZulu-Natal, Phinda

450

VIDUIDAE

Zimbabwe, (Chirundu, Victoria Falls) some birds in breeding plumage as late as Aug and Oct (Tree 1994). DEVELOPMENT AND CARE OF YOUNG: nestling period unknown. Fledged young occur in family groups of Jameson’s Firefinches, indicating that parasite and host young are reared together.

Plate 27 (Opp. p. 410)

BREEDING SUCCESS/SURVIVAL: 2 colour-ringed YY (Lochinvar Nat. Park, Zambia) sang on same call-site in breeding season for 4 successive years.

Key References Payne, R.B. (1968a, 1973a, 1977a, 1982, 1985b, 1987, 1994a, 1996), Payne, R.B. et al. (1993).

Vidua codringtoni (Neave). Zambezi Indigobird; Peters’s Twinspot Indigobird. Combassou de Codrington. Hypochera codringtoni Neave, 1907. Mem. Manchester Lit. Phil. Soc., 51; Molilo’s, Loangwa Valley, northeast Rhodesia [¼ Zambia]. (V. funerea lusituensis Payne 1973 is a synonym). Range and Status. Endemic resident. S Tanzania (Mikindani, Iringa, L. Rukwa). Zambia (Lusaka, Chilanga, ˆ i (Zomba, Lochinvar Nat. Park, Luangwa Valley). Malaw Namadzi, Limbe, Mt Mulanje at Likulezi Mission, Lengwe Nat. Park, Chididi hills). Zimbabwe (Gwaai R., Kwekwe, Kadoma, Penhalonga, Mutare, Rusitu R. at 380 m, Chipinge). Range is within that of presumptive host species, Peters’s Twinspot Hypargos niveoguttatus. Description. ADULT Y (breeding): black with bright blue or green gloss, tail black, concealed flank spot white; wing with inner 3 secondaries (tertials) and their coverts glossy black, other secondaries and primaries and upperwing-coverts black, underwingcoverts light grey (sometimes some coverts black). Bill white, eyes dark brown, feet bright orange or red. ADULT Y (non-breeding): like X but more strongly marked. ADULT X: upperparts brown with blackish shaft streaks, crown with central pale brown stripe streaked black, bordered by dark brown stripe, rump unstreaked brown, tail dark brown, nearly black, without white border; face pale with whitish supraorbital stripe and dark stripe from eye to lighter brown unstreaked nape, ear-spot blackish and indistinct; chin and throat whitish, breast unstreaked grey, sides of breast and flanks buffy grey, undertail-coverts white, wings dark brown with buffy wing-bars formed by upperwing-coverts, underwing-coverts grey. Bill whitish to grey or grey-brown, upper mandible darker than lower; eyes dark; feet orange, duller in non-breeding season. ˆ i, Zimbabwe) wing, Y (n ¼ 31) 66–70 (675), X (n ¼ 4) SIZE: (Malaw 65–66 (655); tail, Y (n ¼ 30) 37–41 (391), X (n ¼ 4) 39–41 (398); bill, Y (n ¼ 29) 81–100 (89), X (n ¼ 4) 80–89 (86); tarsus, Y ˆ i, (n ¼ 31) 142–160 (146), X (n ¼ 4) 14 (140). WEIGHT: Malaw Zimbabwe, Y (n ¼ 29) 114–141 (128), X (n ¼ 4) 123–133 (129). IMMATURE: juv. upperparts less streaked than in ad. X, crown nearly uniformly grey-brown (streaked in ad.), face with indistinct buffy supraorbital stripe, pattern less distinct than in ad. X, breast buffy grey, belly and undertail-coverts whitish. Mouth yellow with 3 black palate spots and without white gape papillae and blue base. NESTLING: unknown.

Field Characters. Length 10–11 cm. Breeding Y black with blue or green gloss, white bill, orange or red legs and feet; told from sympatric species by much brighter gloss, black (not brown) wings. Also identified by imitations of Peters’s Twinspot. X has whitish bill, orange feet and more distinct separation of grey breast and white belly than other indigobirds. Voice. Tape-recorded (86, 102, 103, PAY, STJ). Song churring and scratchy as in other indigobirds, mixed with

Vidua codringtoni

imitations of songs and calls of Peters’s Twinspot. Each Y has repertoire of c. 20 song types, a third of which are mimetic. Slow and fast chatters similar to those used in songs are given in aggressive contexts, interspersed with complex song. YY within a few km of each other match all their song types; YY a few km away have different sets of songs. Most distinctive and frequently heard mimicked twinspot vocalization is alarm trill ‘trrrrrrrreee’, higher in pitch (7–8 kHz) than firefinch alarm trills (4 kHz). Also mimicked are: (1) simple song, high ‘treeee’ followed by lower whistles and rapid chatter; (2) long, complex, soft song (used in twinspot sexual display) with whistles, trills and chatters; (3) short excitement calls, ‘sip’ and ‘tsisi’; and (4) begging calls, which resemble those of young firefinches (Payne et al. 1992, 1993, Payne and Payne 2001). Occasional birds imitate hosts other than twinspots – one in Zimbabwe mimicked African Firefinch Lagonosticta rubricata and matched its songs with local Dusky Indigobirds V. funerea; another imitated Jameson’s Firefinch L. rhodopareia. A Y in South Africa (KwaZulu-Natal, Phinda

Fringilla

Food. Grass seeds 1–2 mm in diam.

range; song mimicry) (Payne et al. 1992, 1993). In captivity, parasitizes this twinspot (Colahan 1995). EGGS: sets of 3, laid 1 a day, a few days between sets. Little gloss; immaculate white. SIZE: unknown. LAYING DATES: E Zimbabwe, (oviduct eggs, Mar–Apr); ˆ i, (YY in breeding plumage sing, court Zambia and Malaw and mate, Mar–Apr); South Africa (Y in breeding plumage, in song, Mar). Nothing further known.

Breeding Habits. Brood parasite. Polygynous, territorial. Evidently parasitizes Peters’s Twinspot (distributional

Key References Payne, R.B. (1973a, 1977a, 1994a, 1996), Payne, R.B. et al. (1992, 1993).

Nature Reserve) mimicked songs of Peters’s Twinspot (D. Allan, pers. comm.). General Habits. Inhabits riverine and flooded thickets, and dense acacia woodland thickets. Uses same habitat as Peters’s Twinspot. Territorial Y defends his song perch as a mating site. Feeds on ground, Y often in territory.

Family FRINGILLIDAE: finches

The term ‘finch’ has been used for stout-billed seed-eating birds of many unrelated families, including some in Africa (e.g. Estrildidae, ‘weaver-finches’). On the basis of DNA hybridization studies, Sibley and Monroe (1990) include some 900 species in Fringillidae in their subfamilies Peucedraminae (the New World Olive Warbler Peucedramus taeniatus, a sisterspecies of all of the others), Fringillinae and Emberizinae. Earlier authorities (e.g. Newton 1972, 1985) and some later ones (e.g. Clement et al. 1993) restrict the family to the ‘true’ finches, i.e. the 3 chaffinches Fringilla (Fringillinae sens. str.) and the 130 or so cardueline species known, somewhat arbitrarily, as finches, serins, canaries, siskins, seedeaters, grosbeaks, linnets and crossbills (Carduelinae). We do likewise, if for no better reason than convenience. Size range c. 10–100 g; bills generally stout and conical; large jaw muscles; skulls strong (and differing from skulls of other stout-billed granivorous birds in some details of morphology); groove at edge of palate; seed wedged in bill using the grooves, crushed by raising mandible up onto it, and dehusked using tongue. Bill form varies with diet: large, deep and powerful in grosbeaks (adapted for crushing large, hard tree-fruits); with maxilla and mandible short and rounded (for eating buds); with tips crossed asymmetrically (for extracting seeds from conifer cones); or rather long and slender (for extracting seeds from fruiting heads of plants such as Compositae). 9 (not 10) large primaries, 12 large rectrices; nest a neat, open cup; built by X only; incubation by X only. About 135 spp., 61 in Africa (41 endemic), nearly all others in Nearctic or Palearctic.

Subfamily FRINGILLINAE: chaffinches

A single genus of sexually dimorphic finches; YY bluish and orange-pink, blue, or black and orange, with prominent shoulder patches; XX rather less colourful; bill  half length of head, sharp-pointed, culmen almost straight; tail only slightly indented, white-sided; do not possess a crop. Forage on the ground; feed their young entirely on insects, carried in bill to the nest, 1 at a time; do not feed by regurgitation; territories large; sexual behaviour and (in breeding season) foodgathering entirely within territory; territory defended by singing only at perch, i.e. no song-flights; song loud, short, highly stereotyped; no courtship feeding.

Genus Fringilla Linnaeus

Characters those of the subfamily. 3 species: one in W Palearctic, breeding in N Africa; another confined to W Canary Is; the 3rd N Palearctic, migrating into S Europe, N Africa and S and E Asia. They are sometimes held to compose a superspecies (e.g. Cramp and Perrins 1994), from which we exclude F. montifringilla.

451

Fringilla

Food. Grass seeds 1–2 mm in diam.

range; song mimicry) (Payne et al. 1992, 1993). In captivity, parasitizes this twinspot (Colahan 1995). EGGS: sets of 3, laid 1 a day, a few days between sets. Little gloss; immaculate white. SIZE: unknown. LAYING DATES: E Zimbabwe, (oviduct eggs, Mar–Apr); ˆ i, (YY in breeding plumage sing, court Zambia and Malaw and mate, Mar–Apr); South Africa (Y in breeding plumage, in song, Mar). Nothing further known.

Breeding Habits. Brood parasite. Polygynous, territorial. Evidently parasitizes Peters’s Twinspot (distributional

Key References Payne, R.B. (1973a, 1977a, 1994a, 1996), Payne, R.B. et al. (1992, 1993).

Nature Reserve) mimicked songs of Peters’s Twinspot (D. Allan, pers. comm.). General Habits. Inhabits riverine and flooded thickets, and dense acacia woodland thickets. Uses same habitat as Peters’s Twinspot. Territorial Y defends his song perch as a mating site. Feeds on ground, Y often in territory.

Family FRINGILLIDAE: finches

The term ‘finch’ has been used for stout-billed seed-eating birds of many unrelated families, including some in Africa (e.g. Estrildidae, ‘weaver-finches’). On the basis of DNA hybridization studies, Sibley and Monroe (1990) include some 900 species in Fringillidae in their subfamilies Peucedraminae (the New World Olive Warbler Peucedramus taeniatus, a sisterspecies of all of the others), Fringillinae and Emberizinae. Earlier authorities (e.g. Newton 1972, 1985) and some later ones (e.g. Clement et al. 1993) restrict the family to the ‘true’ finches, i.e. the 3 chaffinches Fringilla (Fringillinae sens. str.) and the 130 or so cardueline species known, somewhat arbitrarily, as finches, serins, canaries, siskins, seedeaters, grosbeaks, linnets and crossbills (Carduelinae). We do likewise, if for no better reason than convenience. Size range c. 10–100 g; bills generally stout and conical; large jaw muscles; skulls strong (and differing from skulls of other stout-billed granivorous birds in some details of morphology); groove at edge of palate; seed wedged in bill using the grooves, crushed by raising mandible up onto it, and dehusked using tongue. Bill form varies with diet: large, deep and powerful in grosbeaks (adapted for crushing large, hard tree-fruits); with maxilla and mandible short and rounded (for eating buds); with tips crossed asymmetrically (for extracting seeds from conifer cones); or rather long and slender (for extracting seeds from fruiting heads of plants such as Compositae). 9 (not 10) large primaries, 12 large rectrices; nest a neat, open cup; built by X only; incubation by X only. About 135 spp., 61 in Africa (41 endemic), nearly all others in Nearctic or Palearctic.

Subfamily FRINGILLINAE: chaffinches

A single genus of sexually dimorphic finches; YY bluish and orange-pink, blue, or black and orange, with prominent shoulder patches; XX rather less colourful; bill  half length of head, sharp-pointed, culmen almost straight; tail only slightly indented, white-sided; do not possess a crop. Forage on the ground; feed their young entirely on insects, carried in bill to the nest, 1 at a time; do not feed by regurgitation; territories large; sexual behaviour and (in breeding season) foodgathering entirely within territory; territory defended by singing only at perch, i.e. no song-flights; song loud, short, highly stereotyped; no courtship feeding.

Genus Fringilla Linnaeus

Characters those of the subfamily. 3 species: one in W Palearctic, breeding in N Africa; another confined to W Canary Is; the 3rd N Palearctic, migrating into S Europe, N Africa and S and E Asia. They are sometimes held to compose a superspecies (e.g. Cramp and Perrins 1994), from which we exclude F. montifringilla.

451

Fringilla

Food. Grass seeds 1–2 mm in diam.

range; song mimicry) (Payne et al. 1992, 1993). In captivity, parasitizes this twinspot (Colahan 1995). EGGS: sets of 3, laid 1 a day, a few days between sets. Little gloss; immaculate white. SIZE: unknown. LAYING DATES: E Zimbabwe, (oviduct eggs, Mar–Apr); ˆ i, (YY in breeding plumage sing, court Zambia and Malaw and mate, Mar–Apr); South Africa (Y in breeding plumage, in song, Mar). Nothing further known.

Breeding Habits. Brood parasite. Polygynous, territorial. Evidently parasitizes Peters’s Twinspot (distributional

Key References Payne, R.B. (1973a, 1977a, 1994a, 1996), Payne, R.B. et al. (1992, 1993).

Nature Reserve) mimicked songs of Peters’s Twinspot (D. Allan, pers. comm.). General Habits. Inhabits riverine and flooded thickets, and dense acacia woodland thickets. Uses same habitat as Peters’s Twinspot. Territorial Y defends his song perch as a mating site. Feeds on ground, Y often in territory.

Family FRINGILLIDAE: finches

The term ‘finch’ has been used for stout-billed seed-eating birds of many unrelated families, including some in Africa (e.g. Estrildidae, ‘weaver-finches’). On the basis of DNA hybridization studies, Sibley and Monroe (1990) include some 900 species in Fringillidae in their subfamilies Peucedraminae (the New World Olive Warbler Peucedramus taeniatus, a sisterspecies of all of the others), Fringillinae and Emberizinae. Earlier authorities (e.g. Newton 1972, 1985) and some later ones (e.g. Clement et al. 1993) restrict the family to the ‘true’ finches, i.e. the 3 chaffinches Fringilla (Fringillinae sens. str.) and the 130 or so cardueline species known, somewhat arbitrarily, as finches, serins, canaries, siskins, seedeaters, grosbeaks, linnets and crossbills (Carduelinae). We do likewise, if for no better reason than convenience. Size range c. 10–100 g; bills generally stout and conical; large jaw muscles; skulls strong (and differing from skulls of other stout-billed granivorous birds in some details of morphology); groove at edge of palate; seed wedged in bill using the grooves, crushed by raising mandible up onto it, and dehusked using tongue. Bill form varies with diet: large, deep and powerful in grosbeaks (adapted for crushing large, hard tree-fruits); with maxilla and mandible short and rounded (for eating buds); with tips crossed asymmetrically (for extracting seeds from conifer cones); or rather long and slender (for extracting seeds from fruiting heads of plants such as Compositae). 9 (not 10) large primaries, 12 large rectrices; nest a neat, open cup; built by X only; incubation by X only. About 135 spp., 61 in Africa (41 endemic), nearly all others in Nearctic or Palearctic.

Subfamily FRINGILLINAE: chaffinches

A single genus of sexually dimorphic finches; YY bluish and orange-pink, blue, or black and orange, with prominent shoulder patches; XX rather less colourful; bill  half length of head, sharp-pointed, culmen almost straight; tail only slightly indented, white-sided; do not possess a crop. Forage on the ground; feed their young entirely on insects, carried in bill to the nest, 1 at a time; do not feed by regurgitation; territories large; sexual behaviour and (in breeding season) foodgathering entirely within territory; territory defended by singing only at perch, i.e. no song-flights; song loud, short, highly stereotyped; no courtship feeding.

Genus Fringilla Linnaeus

Characters those of the subfamily. 3 species: one in W Palearctic, breeding in N Africa; another confined to W Canary Is; the 3rd N Palearctic, migrating into S Europe, N Africa and S and E Asia. They are sometimes held to compose a superspecies (e.g. Cramp and Perrins 1994), from which we exclude F. montifringilla.

451

Fringilla

Food. Grass seeds 1–2 mm in diam.

range; song mimicry) (Payne et al. 1992, 1993). In captivity, parasitizes this twinspot (Colahan 1995). EGGS: sets of 3, laid 1 a day, a few days between sets. Little gloss; immaculate white. SIZE: unknown. LAYING DATES: E Zimbabwe, (oviduct eggs, Mar–Apr); ˆ i, (YY in breeding plumage sing, court Zambia and Malaw and mate, Mar–Apr); South Africa (Y in breeding plumage, in song, Mar). Nothing further known.

Breeding Habits. Brood parasite. Polygynous, territorial. Evidently parasitizes Peters’s Twinspot (distributional

Key References Payne, R.B. (1973a, 1977a, 1994a, 1996), Payne, R.B. et al. (1992, 1993).

Nature Reserve) mimicked songs of Peters’s Twinspot (D. Allan, pers. comm.). General Habits. Inhabits riverine and flooded thickets, and dense acacia woodland thickets. Uses same habitat as Peters’s Twinspot. Territorial Y defends his song perch as a mating site. Feeds on ground, Y often in territory.

Family FRINGILLIDAE: finches

The term ‘finch’ has been used for stout-billed seed-eating birds of many unrelated families, including some in Africa (e.g. Estrildidae, ‘weaver-finches’). On the basis of DNA hybridization studies, Sibley and Monroe (1990) include some 900 species in Fringillidae in their subfamilies Peucedraminae (the New World Olive Warbler Peucedramus taeniatus, a sisterspecies of all of the others), Fringillinae and Emberizinae. Earlier authorities (e.g. Newton 1972, 1985) and some later ones (e.g. Clement et al. 1993) restrict the family to the ‘true’ finches, i.e. the 3 chaffinches Fringilla (Fringillinae sens. str.) and the 130 or so cardueline species known, somewhat arbitrarily, as finches, serins, canaries, siskins, seedeaters, grosbeaks, linnets and crossbills (Carduelinae). We do likewise, if for no better reason than convenience. Size range c. 10–100 g; bills generally stout and conical; large jaw muscles; skulls strong (and differing from skulls of other stout-billed granivorous birds in some details of morphology); groove at edge of palate; seed wedged in bill using the grooves, crushed by raising mandible up onto it, and dehusked using tongue. Bill form varies with diet: large, deep and powerful in grosbeaks (adapted for crushing large, hard tree-fruits); with maxilla and mandible short and rounded (for eating buds); with tips crossed asymmetrically (for extracting seeds from conifer cones); or rather long and slender (for extracting seeds from fruiting heads of plants such as Compositae). 9 (not 10) large primaries, 12 large rectrices; nest a neat, open cup; built by X only; incubation by X only. About 135 spp., 61 in Africa (41 endemic), nearly all others in Nearctic or Palearctic.

Subfamily FRINGILLINAE: chaffinches

A single genus of sexually dimorphic finches; YY bluish and orange-pink, blue, or black and orange, with prominent shoulder patches; XX rather less colourful; bill  half length of head, sharp-pointed, culmen almost straight; tail only slightly indented, white-sided; do not possess a crop. Forage on the ground; feed their young entirely on insects, carried in bill to the nest, 1 at a time; do not feed by regurgitation; territories large; sexual behaviour and (in breeding season) foodgathering entirely within territory; territory defended by singing only at perch, i.e. no song-flights; song loud, short, highly stereotyped; no courtship feeding.

Genus Fringilla Linnaeus

Characters those of the subfamily. 3 species: one in W Palearctic, breeding in N Africa; another confined to W Canary Is; the 3rd N Palearctic, migrating into S Europe, N Africa and S and E Asia. They are sometimes held to compose a superspecies (e.g. Cramp and Perrins 1994), from which we exclude F. montifringilla.

451

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FRINGILLIDAE

Plate 34

Fringilla coelebs Linnaeus. Chaffinch. Pinson des arbres.

(Opp. p. 523)

Fringilla coelebs Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 179; Sweden. Forms a superspecies with F. teydea (Canary Is). Range and Status. W Palearctic; breeding summer visitor, W and N Norway and Belorussya east to L. Baykal; resident and partial migrant, rest of Europe and N Africa east to Caspian Sea and SW Turkmenistan, south to Canary Is and S Sinai. 7 island endemic subspecies, 3 in Canary Is and 1 each in Azores, Madeira, Sardinia and Crete. F. c. africana is common, widespread resident and F. c. coelebs an abundant winter visitor in Maghreb. A sparse resident (spodiogenys) and irregular winter visitor, coastal Libya and N Egypt. Vagrant Mauritania (1, Nouadhibou, Oct 1988), Libyan Desert (1, Jaghbub, Feb 1928) and S Egypt (1, Aswan, Feb 1976). Morocco, africana widespread in Souss and south to Haut Atlas and Tiznit; from coast up to 2200 m in Rif (Jebel Lakraa, Jebel Tidighine), 2100 m in Moyen Atlas and 1800 m in Haut Atlas; in flocks of up to 60–300 in Nov and Dec, e.g. in Aleppo pines in Ras Negri, Aguelmane Afenourir and Aı¨n Nokra; commoner in mountain-side woods than on plains; coelebs mixes with africana and occurs also in large flocks on its own, e.g. 200 near Ifrane in Jan, more near Rabat in Feb–Mar. Algeria, africana widespread from coast to Atlas Mts and Aure`s, up to 2200– 2300 m; in winter south to N oases (e.g. Biskra) and exceptionally to Oued Ameskiki and Tinrhert Hamada; coelebs regular from Oct/Nov to Feb/Mar, from coast to High Plateaux, but abundance hard to estimate (Isenmann and Moali 2000). Tunisia, africana in NW and spodiogenys elsewhere, common south to Chebka and Mahdia, local and rare south to Kettana and Gafsa; perhaps absent from central plateau; coelebs an uncommon winter visitor in N. Libya, spodiogenys breeds sparsely on Jebel Nafusa west of Garian (Tripoli); africana (or spodiogenys?) breeds in Jebel Akhdar (Cyrenaica); coelebs winters in coastal zone, in W sometimes commonly, in flocks of up to 100, in E sparsely. Egypt, coelebs winters regularly in small flocks along N coast, in Nile Delta and Suez Canal area (tens of thousands once arrived in N Sinai in Oct, in 1942); rare in Western Desert (Wadi Natrun, Baharia) and Eastern Desert (Gebel el Galala el Qibliya). F. c. gengleri introduced from Britain to Groot Schuur Estate, Cape Town, South Africa, in 1897; became established in Cape Peninsula, where some 50 pairs survive on E slopes between Newlands and Silvermine and on W slopes between Constantia Nek and Hout Bay; vagrant to Cape of Good Hope Nat. Res. (1 at Kenton-on-Sea doubtless an escapee); numbers have remained stable, perhaps with some decline since 1960s. Density in maquis in Morocco of 375 pairs per km2. Density in Pinus radiata of 9 birds per km2 in Tenerife (Canary Is) but 10 times higher in NW Spain. Description. F. c. africana Levaillant: Morocco to NW Tunisia; perhaps this subspecies in Cyrenaica, NE Libya. ADULT Y: forehead black, feathers fringed buff-brown when fresh; crown to hindneck, sides of neck, cheeks and ear-coverts blue-grey, fringed

Fringilla coelebs

buff-brown when fresh, especially on sides of crown and neck; whitish spot on centre of nape; mantle and inner scapulars olivegreen; outer scapulars blue-grey, tinged green; back and rump brighter olive-green or mixed green and blue-grey; uppertailcoverts blue-grey, tinged green on inner webs. Central tail feathers blackish on outer web, dark grey on inner web, narrowly fringed whitish; T2–T4 black, inner web of T2 sometimes with small white subterminal spot, of T3 with white spot or distal wedge, of T4 with large white wedge extending up to 40 mm along shaft; T5–T6 white except black distal third of outer web, black along shaft and black base. Lores black; narrow whitish eye-ring, broken at front and rear. Chin to breast, flanks and side of belly pale cinnamon-pink, merging into creamy white centre of belly, thighs and undertail-coverts. Remiges blackish, primaries narrowly edged yellowish white, secondaries and outer tertial edged pale yellowish, central and inner tertial broadly fringed cinnamon on outer web; bases of middle and inner primaries white, forming narrow bar along tips of primary coverts; bases of secondaries and outer web of outer tertial more extensively white (hidden on closed wing). Greater coverts blackish, broadly tipped yellowish white, forming conspicuous wing-bar; primary coverts and alula black, the former often narrowly edged green; median coverts and larger lesser coverts white, forming second, broader, wing-bar; smaller lesser coverts blue-grey. Coverts along leading edge of underwing black and white; rest of underwing-coverts and axillaries white; inner borders of undersides of flight feathers white. Bill in breeding season light blue-grey with black tip and cutting edges; otherwise dull grey, greyish horn or pale horn with flesh or mauve tinge, palest at base of lower mandible. Eyes dark brown; legs pale grey, brown-grey or blue-grey. ADULT X: forehead to upper back and scapulars greyish olive-brown with centre of crown and nape greyer (forming more prominent pale patch in worn plumage); lower back, rump and shorter uppertailcoverts bright olive-green; longer uppertail-coverts grey-brown.

Fringilla coelebs Tail patterned as in ad. Y, but T1 grey-brown; T2–T6 brownish black, white wedges less strongly contrasting, that on T4 usually smaller. Lores and sides of head grey-brown, with diffuse paler superciliary stripe and purer grey sides to neck. Chin to center of throat and breast greyish white, sides of breast and flanks pale grey-brown, centre of belly to undertail-coverts creamy white. Wing similar to ad. Y, but remiges and primary coverts blackish brown; primary coverts edged olive; greater coverts edged olive and white tips tinged olive-yellow; smaller lesser coverts dark greyish. Bare parts as non-breeding ad. Y. SIZE (10 YY, 10 XX): wing, Y 87–95 (908), X 81–89 (842); tail, Y 67–74 (704), X 62– 68 (649); bill, Y 145–16 (151), X 14–16 (147); tarsus, Y 185–20 (192), X 18–20 (192). Larger in Atlas Saharien area of Algeria, smaller in Tanger and Er Rif area of N Morocco. WEIGHT: Haut Atlas, Sept, Y (n ¼ 14) av. 234, X (n ¼ 8) av. 220. IMMATURE: juv. like ad. X but head and body feathering short and soft; tail feathers narrower, more pointed. First-winter like ad., but juv. tail retained. NESTLING (nominate coelebs): down long and plentiful, on upperparts, upperwing, thigh and vent, pale smoke-grey; bare skin flesh-pink; mouth red, orange on palate, grey along inside of mandibles. Gape flanges whitish or yellow-white (Cramp and Perrins 1994). F. c. spodiogenys Bonaparte: Tunisia (except NW), NW Libya. Like africana but Y paler blue-grey on head, paler green on mantle and paler pinkish below; white nape spot more pronounced; more white in tail (T3 with large white wedge and T4 mainly white), and in lesser coverts; underwing purer white. X paler, more greyish above, more greyish white below. Wing, Y (n ¼ 10) 86–92 (900). WEIGHT: Tunisia, May, 1Y 258, 1 X 265. F. c. coelebs Linnaeus: continental Europe to Siberia; winters NW Africa. Y differs from africana in having mantle and scapulars dark umber-brown, underparts much darker vinouscinnamon; cheeks and ear-coverts deep vinous-pink (not bluegrey); tail with greyer centre feathers, T4 with little or no white, white on T5 and T6 less extensive. X darker and browner. Wing (Y, n ¼ 10) 86–92 (890). WEIGHT: SE France, winter, Y (n ¼ 87) 19–27 (231), X (n ¼ 45) 18–25 (210). (F. c. balearica Von Jordans: Portugal, Spain south to Gibraltar, Balearic Is. Like coelebs but slightly paler and larger. Not yet found in Africa but almost certainly visits, mixing with migrant coelebs.) F. c. gengleri Kleinschmidt: Britain and Ireland; introduced in Cape Peninsula, South Africa. Like nominate coelebs, but underparts slightly darker, more uniform rufous-cinnamon; wing shorter, Britain and Ireland, Y (n ¼ 39) 82–90 (858). WEIGHT: South Africa, unsexed, 19–31. TAXONOMIC NOTE: although YY of African africana and spodiogenys look very alike and can readily be distinguished in the field from YY of European coelebs, mitochondrial DNA evidence shows that spodiogenys is far more distantly related with africana than africana is with coelebs (Marshall and Baker 1999). In fact spodiogenys is a sister taxon to all other chaffinches in the region, including the distinctive Azores and Canary Is races (moreletti, maderensis, palmae, ombriosa, canariensis); it is derivative of an ancestral population linking Chaffinch with Blue Chaffinch F. teydea of W Canary Is (Collinson 2001). We follow Baker et al. (1990) and Isenmann and Moali (2000) in treating all continental chaffinches as a single species.

Field Characters. Length 145 cm. Y of nominate race (winter visitor to N Africa) has blue-grey head, brown back, brownish pink cheeks and underparts, white shoulder patch and bar on greater coverts, much white in tail. Y of race gengleri (introduced Cape Town) similar, but resident N African birds have grey cheeks, green back and pale pink underparts. XX of all races are drab brownish birds with 2 prominent white wing-bars on dark wing, in flight showing greenish rump and white in tail.

Voice. Tape-recorded (62–73, 76–89, 99, 105, 110–113, 112, B, C, F). Cheerful bustling song of 10–18 notes, descending and accelerating, with a terminal flourish, ‘che-wee-o’; typically in 2 parts, initial notes slower, sometimes slightly liquid, second part faster, more of a rattle, but extremely variable both between individuals and regionally, so much so that local dialects hard to describe (Cramp and Perrins 1994, and see sonagrams therein). N African birds recorded by J.C. Roche (112) have a very short song, ‘jrree, jrree, jrree, chacha-chaaa’, the final ‘chaaa’ taking the place of usual ‘che-wee-o’, but others in Morocco (105) sound like European birds. Common call ‘pink, pink’ similar to Great Tit Parus major; other calls include ‘tyup’ in flight, ‘seee’ in alarm, and plaintive ‘wheat’. General Habits. In Morocco inhabits all areas with broadleaved trees, also conifers and palms: woods, thickets, gardens, orchards, farms, hillsides and gorges; occurs in maquis, and in oases (e.g. Skoura); commonest in cork oaks, cedars, Aleppo pines Pinus halepensis, cypresses Thuja, eucalypts, Argania spinosa (Sapotaceae, a Moroccan endemic), walnuts in valleys in Haut Atlas, and olives. In Algeria occurs in Tetraclinis articulata and xerophytic woodlands and upland cedars; in Tunisia inhabits olive groves in particular; in Libya breeds in olive groves in Tripoli and in juniper woodland in Cyrenaica. In winter, more in farmland and stubble fields. Around Newlands, South Africa, oak and pine forests and plantations, parks and large gardens with alien trees. In pairs, small flocks in winter, often mainly of one sex (coelebs ¼ celibate). Can mix with European Greenfinches Carduelis chloris, Common Linnets C. cannabina and European Goldfinches C. carduelis in winter to form large flocks in fields and at roosts. Often rather shy, but can become very tame when fed around human habitation and in parks. In Cape Peninsula generally in pairs; does not form flocks; said to be rather secretive. Forages mainly on the ground, especially where tilled or with only sparse herb cover, often under large trees. Gait a short, shuffling, jerky hop-walk, bird half crouching. Feeds with rapid pecking action. Flight bounding, undulating; quite fast in the open. Roosts in small or large, monospecific or mixed flocks, in leafy tree or conifer plantation; in Israel in citrus groves and avocado orchards. Migrates by night and day, in small flocks often moving visibly along coasts, river valleys and through mountain passes. Daytime migrations of European coelebs often noted in Morocco, late Oct to mid Nov, e.g. at Cap des Trois Fourches, Ouezzane, Moulay-Bousselham, Saı¨dia, and Marja Mellah; wintering birds quit Moyen Atlas in mid Mar. 200 Chaffinches, race not determined, in Taffert cedars in mid Sept. Autumn passage at Gibraltar and doubtless into Africa starts in Oct and peaks in Nov with some in Dec; 11,000 birds in autumn 1977; populations involved are coelebs from central and E Europe (Finnish and Baltic birds winter in W Iberia). Passage occurs at Gibraltar also in Jan (birds presumably northbound), diminishing in Feb, Mar and Apr (Finlayson 1992). Bird ringed near Kalinigrad recovered in Morocco, 3250 km to southwest; recoveries in Algeria from Germany (1), Switzerland (3), France (1), and Italy (6). Small flocks of africana are caught

453

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FRINGILLIDAE

up in migration of coelebs in spring and appear in Gibraltar. A spodiogenys vagrant to E England in Apr 1994 and 1–3 africana to NW England in Feb–Apr 1998 (arrival coinciding with Saharan red dust there). F. c. coelebs winters in NE Tunisia in Oct–Apr with a few until early May, in Tripoli in Nov–Mar, in Cyrenaica in Oct–Mar and in Egypt from late Oct to late Mar/early Apr; bird ringed in Cairo recovered in Siberia 18 months later, in 1939. Food. In Europe, in breeding season mainly invertebrates (young fed almost entirely on invertebrates, particularly aphids and caterpillars) and in rest of year mainly seeds and other plant material (grass seeds and cereal grain, beechmast, crucifer and chenopod seeds; buds, fruits). Diet more varied than that of any other W Palearctic finch; for inventories, see Cramp and Perrins (1994). In Cape Peninsula eats seeds, grain, small fruits, buds; insects, spiders, snails, worms. Breeding Habits. Solitary nester; monogamous; territorial. Territory size varies greatly, higher in N than S Europe; av. 270 m2 in Moroccan maquis; territory demarcated by Y’s border song-posts, used for nesting and feeding, defended by both Y and X vigorously chasing intruders away. Y sings from top of shrub, telephone wire or exposed position at side of tree, in upright posture with head thrown back. Early in breeding season Y can sing several times a minute for hours on end, patrolling territory by changing song-perch regularly, flying with wings fully extended and beating rapidly through a shallow arc. Displays, not studied in Africa, include Y’s Crouching-lopsided and Upright-lopsided courtship and X’s wing-shivering Soliciting-display (Cramp and Perrins 1994.) 2 clutches per year in NW Africa.

NEST: a neat, compact, firm-walled, deep cup, made of mosses, lichens, rootlets and fibres bound with cobweb, clad with moss and lichen and (SW Cape) sometimes paper, lined with hair, wool, feathers, rootlets and fine grasses; in SW Cape, ext. diam. 90–100, ext. depth 70–80, int. diam. c. 50, int. depth 40–50; sited 3–10 m above ground, in fork in bush or small tree; in SW Cape, nests in cork, oak and pine trees (Tarboton 2001). EGGS: Morocco and Algeria, 3–5, rarely 6, av. (11 clutches) 395; SW Cape, 3–6. Sub-elliptical, smooth, slightly glossy. Colours and markings variable; NW Africa, pale coffee-coloured, spotted and streaked with dark purplebrown, often clouded; occasionally pale blue-green, marked similarly; SW Cape, probably just the same, described as bluish white, brownish or greenish grey, lined and spotted with brown, grey and violet. SIZE: (africana, n ¼ 101) 185– 245  139–163 (207  152); (gengleri, SW Cape, n ¼ 27) 178–210  143–152 (194  146). WEIGHT: av. 26. LAYING DATES: Morocco and Algeria, late Mar to early June, SW Cape, Sept–Nov. INCUBATION: by X period, SW Cape 12–13 days; Europe av. 126 days. DEVELOPMENT AND CARE OF YOUNG: fed and cared for by both parents, mainly X. Nestling period, SW Cape, 12–13 days; Europe 139 days. Young can feed themselves from 2 weeks after leaving nest but are fed by parents for 3 weeks. BREEDING SUCCESS/SURVIVAL: no African data. Adult annual mortality, Spain, 48%. Oldest wild bird 14 years, but few live beyond c. 5 years. Nestling mortality high in years when main food, caterpillars, in short supply.

Key References Clement, P. et al. (1993), Collinson, M. (2001), Cramp, S. and Perrins, C.M. (1994), Isenmann, P. and Moali, A. (2000).

Plate 34

Fringilla montifringilla Linnaeus. Brambling. Pinson du Nord.

(Opp. p. 523)

Fringilla montifringilla Linnaeus, 1758. Syst. Nat., et. 10, 1, p. 179; Sweden. Breeds from Norway to Siberia, mainly north of Arctic Circle except in N-central Asia; wholly migratory, wintering south to Spain, Turkey, Israel, N Iran, Himalayas, China and Japan. Winter vagrant to Iceland, Madeira, Canary Is., N Africa, Crete, Cyprus, Jordan, E and SE Arabia, Philippines and Aleutian Is. Senegal, one visited a ship off S Senegal in Nov 1976; Mauritania, singles Nouadhibou Jan and Apr 1987; Morocco, 32 records up to 1990 and av. of 4 per winter during 1990s, mostly singletons, rarely 2–3 together, south to Oukaı¨meden, Fkihben-Salah, Col du Zad and Agouim; Algeria, regular in small number Oct–Mar, mainly Nov–Apr, in Tell, up to 10 in Apr 1992 near Tizi Ouzou, once near Be´ni Abbe`s; Tunisia, 1, Lac Tunis, Apr 1970; Libya, 1 Benghazi 1977; Egypt, 11 old records in Nile Delta (Meinertzhagen 1930) require validation; 2, Port Said, Nov 1959; 2, Safaga, Mar 1982; others in NW and S Sinai. Adult Y distinctive, with black head, rust-orange breast and scapulars, white rump, black wings and tail with feathers conspicuously tipped and edged with white or pale orange, and dark-spotted flanks; bill yellow in winter. X similar but head pale, with soft, dark band from forehead and above eyebrow to side of neck; ear-coverts dusky; flanks often spotted; best told from X Chaffinch F. coelebs by white rump and orange scapulars. Length 145–175 cm; wing (S France, winter) Y (n ¼ 34) 88– 97 (934), X (n ¼ 45) 85–93 (882); tail (NW Europe, summer), Y (n ¼ 22) 63–71 (663), X (n ¼ 17) 57–66 (620); weight, Y averages 22–28, X averages c. 1 g less. In winter more commonly in flock of Chaffinches than on its own, but wary, and much less around human habitation; inhabits wooded farmland, hillsides and shrubby wadis; uses trees freely but forages on ground, eating seeds. Less vocal than Chaffinch; flight call a hard, clipped, slightly hoarse ‘tyek’. Another common call, nasal, rising ‘dzhayek’. Often sings in winter quarters: long, buzzy, nasal ‘dzhweeee’, alternating with short dry rattle, reminiscent of European Greenfich Carduelis chloris.

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FRINGILLIDAE

up in migration of coelebs in spring and appear in Gibraltar. A spodiogenys vagrant to E England in Apr 1994 and 1–3 africana to NW England in Feb–Apr 1998 (arrival coinciding with Saharan red dust there). F. c. coelebs winters in NE Tunisia in Oct–Apr with a few until early May, in Tripoli in Nov–Mar, in Cyrenaica in Oct–Mar and in Egypt from late Oct to late Mar/early Apr; bird ringed in Cairo recovered in Siberia 18 months later, in 1939. Food. In Europe, in breeding season mainly invertebrates (young fed almost entirely on invertebrates, particularly aphids and caterpillars) and in rest of year mainly seeds and other plant material (grass seeds and cereal grain, beechmast, crucifer and chenopod seeds; buds, fruits). Diet more varied than that of any other W Palearctic finch; for inventories, see Cramp and Perrins (1994). In Cape Peninsula eats seeds, grain, small fruits, buds; insects, spiders, snails, worms. Breeding Habits. Solitary nester; monogamous; territorial. Territory size varies greatly, higher in N than S Europe; av. 270 m2 in Moroccan maquis; territory demarcated by Y’s border song-posts, used for nesting and feeding, defended by both Y and X vigorously chasing intruders away. Y sings from top of shrub, telephone wire or exposed position at side of tree, in upright posture with head thrown back. Early in breeding season Y can sing several times a minute for hours on end, patrolling territory by changing song-perch regularly, flying with wings fully extended and beating rapidly through a shallow arc. Displays, not studied in Africa, include Y’s Crouching-lopsided and Upright-lopsided courtship and X’s wing-shivering Soliciting-display (Cramp and Perrins 1994.) 2 clutches per year in NW Africa.

NEST: a neat, compact, firm-walled, deep cup, made of mosses, lichens, rootlets and fibres bound with cobweb, clad with moss and lichen and (SW Cape) sometimes paper, lined with hair, wool, feathers, rootlets and fine grasses; in SW Cape, ext. diam. 90–100, ext. depth 70–80, int. diam. c. 50, int. depth 40–50; sited 3–10 m above ground, in fork in bush or small tree; in SW Cape, nests in cork, oak and pine trees (Tarboton 2001). EGGS: Morocco and Algeria, 3–5, rarely 6, av. (11 clutches) 395; SW Cape, 3–6. Sub-elliptical, smooth, slightly glossy. Colours and markings variable; NW Africa, pale coffee-coloured, spotted and streaked with dark purplebrown, often clouded; occasionally pale blue-green, marked similarly; SW Cape, probably just the same, described as bluish white, brownish or greenish grey, lined and spotted with brown, grey and violet. SIZE: (africana, n ¼ 101) 185– 245  139–163 (207  152); (gengleri, SW Cape, n ¼ 27) 178–210  143–152 (194  146). WEIGHT: av. 26. LAYING DATES: Morocco and Algeria, late Mar to early June, SW Cape, Sept–Nov. INCUBATION: by X period, SW Cape 12–13 days; Europe av. 126 days. DEVELOPMENT AND CARE OF YOUNG: fed and cared for by both parents, mainly X. Nestling period, SW Cape, 12–13 days; Europe 139 days. Young can feed themselves from 2 weeks after leaving nest but are fed by parents for 3 weeks. BREEDING SUCCESS/SURVIVAL: no African data. Adult annual mortality, Spain, 48%. Oldest wild bird 14 years, but few live beyond c. 5 years. Nestling mortality high in years when main food, caterpillars, in short supply.

Key References Clement, P. et al. (1993), Collinson, M. (2001), Cramp, S. and Perrins, C.M. (1994), Isenmann, P. and Moali, A. (2000).

Plate 34

Fringilla montifringilla Linnaeus. Brambling. Pinson du Nord.

(Opp. p. 523)

Fringilla montifringilla Linnaeus, 1758. Syst. Nat., et. 10, 1, p. 179; Sweden. Breeds from Norway to Siberia, mainly north of Arctic Circle except in N-central Asia; wholly migratory, wintering south to Spain, Turkey, Israel, N Iran, Himalayas, China and Japan. Winter vagrant to Iceland, Madeira, Canary Is., N Africa, Crete, Cyprus, Jordan, E and SE Arabia, Philippines and Aleutian Is. Senegal, one visited a ship off S Senegal in Nov 1976; Mauritania, singles Nouadhibou Jan and Apr 1987; Morocco, 32 records up to 1990 and av. of 4 per winter during 1990s, mostly singletons, rarely 2–3 together, south to Oukaı¨meden, Fkihben-Salah, Col du Zad and Agouim; Algeria, regular in small number Oct–Mar, mainly Nov–Apr, in Tell, up to 10 in Apr 1992 near Tizi Ouzou, once near Be´ni Abbe`s; Tunisia, 1, Lac Tunis, Apr 1970; Libya, 1 Benghazi 1977; Egypt, 11 old records in Nile Delta (Meinertzhagen 1930) require validation; 2, Port Said, Nov 1959; 2, Safaga, Mar 1982; others in NW and S Sinai. Adult Y distinctive, with black head, rust-orange breast and scapulars, white rump, black wings and tail with feathers conspicuously tipped and edged with white or pale orange, and dark-spotted flanks; bill yellow in winter. X similar but head pale, with soft, dark band from forehead and above eyebrow to side of neck; ear-coverts dusky; flanks often spotted; best told from X Chaffinch F. coelebs by white rump and orange scapulars. Length 145–175 cm; wing (S France, winter) Y (n ¼ 34) 88– 97 (934), X (n ¼ 45) 85–93 (882); tail (NW Europe, summer), Y (n ¼ 22) 63–71 (663), X (n ¼ 17) 57–66 (620); weight, Y averages 22–28, X averages c. 1 g less. In winter more commonly in flock of Chaffinches than on its own, but wary, and much less around human habitation; inhabits wooded farmland, hillsides and shrubby wadis; uses trees freely but forages on ground, eating seeds. Less vocal than Chaffinch; flight call a hard, clipped, slightly hoarse ‘tyek’. Another common call, nasal, rising ‘dzhayek’. Often sings in winter quarters: long, buzzy, nasal ‘dzhweeee’, alternating with short dry rattle, reminiscent of European Greenfich Carduelis chloris.

Serinus

Fringilla montifringilla

Subfamily CARDUELINAE: serins, canaries, Old World seedeaters, goldfinches, grosbeaks and others

Sexually dimorphic or nearly monomorphic; YY often strikingly patterned and coloured, in reds, yellows and greens; XX and juvs of most spp. brownish, streaky; bill shape varied (see Family Fringillidae diagnosis); tail usually deeply notched. Possess a crop. Forage on ground and at fruiting heads of herbs, shrubs and trees; feed their young mainly on seeds, carried in crop, given by regurgitation. Many species loosely colonial; territories small; sexual behaviour not entirely within territory; food gathered outside territory; territory defended by singing at perch and in song-flights; song often quite protracted and varied, not as stereotyped as in Fringillinae; Y courtship-feeds X. 60 spp. in Africa (42 endemic), nearly all other species being Nearctic or Palearctic. 20 genera are small; Serinus, Carduelis and Carpodacus contain nearly 100 species between them.

Genus Serinus Koch

An essentially African group of small or medium-sized yellow-olive or brown finches with short, stumpy bills: the serins, canaries and Old World seedeaters. Culmen decurved or (in a few species) straight; bill horn-coloured, grey or pink; bill large and stout in some spp., relatively fine and long in a few others. Lowland and montane; open country, forest edge, scrub, grassland; one species associated with papyrus, another with ilala palm. Arboreal, foraging partly in trees, shrubs and herbs although mainly on ground. Nest a neat, open cup, snugly lined, usually in bush or tree. Characters of bill morphology, plumage colour and pattern have a mosaic distribution, such that very different classifications were reached by earlier systematists. Plumage and breeding behaviour characters studied by Nicolai (1960, investigating 9 spp.) and van den Elzen (1985, 18 spp.), and morphological characters by van den Elzen et al. (1987). Van den Elzen (1985) pointed out

455

Serinus

Fringilla montifringilla

Subfamily CARDUELINAE: serins, canaries, Old World seedeaters, goldfinches, grosbeaks and others

Sexually dimorphic or nearly monomorphic; YY often strikingly patterned and coloured, in reds, yellows and greens; XX and juvs of most spp. brownish, streaky; bill shape varied (see Family Fringillidae diagnosis); tail usually deeply notched. Possess a crop. Forage on ground and at fruiting heads of herbs, shrubs and trees; feed their young mainly on seeds, carried in crop, given by regurgitation. Many species loosely colonial; territories small; sexual behaviour not entirely within territory; food gathered outside territory; territory defended by singing at perch and in song-flights; song often quite protracted and varied, not as stereotyped as in Fringillinae; Y courtship-feeds X. 60 spp. in Africa (42 endemic), nearly all other species being Nearctic or Palearctic. 20 genera are small; Serinus, Carduelis and Carpodacus contain nearly 100 species between them.

Genus Serinus Koch

An essentially African group of small or medium-sized yellow-olive or brown finches with short, stumpy bills: the serins, canaries and Old World seedeaters. Culmen decurved or (in a few species) straight; bill horn-coloured, grey or pink; bill large and stout in some spp., relatively fine and long in a few others. Lowland and montane; open country, forest edge, scrub, grassland; one species associated with papyrus, another with ilala palm. Arboreal, foraging partly in trees, shrubs and herbs although mainly on ground. Nest a neat, open cup, snugly lined, usually in bush or tree. Characters of bill morphology, plumage colour and pattern have a mosaic distribution, such that very different classifications were reached by earlier systematists. Plumage and breeding behaviour characters studied by Nicolai (1960, investigating 9 spp.) and van den Elzen (1985, 18 spp.), and morphological characters by van den Elzen et al. (1987). Van den Elzen (1985) pointed out

455

Serinus

Fringilla montifringilla

Subfamily CARDUELINAE: serins, canaries, Old World seedeaters, goldfinches, grosbeaks and others

Sexually dimorphic or nearly monomorphic; YY often strikingly patterned and coloured, in reds, yellows and greens; XX and juvs of most spp. brownish, streaky; bill shape varied (see Family Fringillidae diagnosis); tail usually deeply notched. Possess a crop. Forage on ground and at fruiting heads of herbs, shrubs and trees; feed their young mainly on seeds, carried in crop, given by regurgitation. Many species loosely colonial; territories small; sexual behaviour not entirely within territory; food gathered outside territory; territory defended by singing at perch and in song-flights; song often quite protracted and varied, not as stereotyped as in Fringillinae; Y courtship-feeds X. 60 spp. in Africa (42 endemic), nearly all other species being Nearctic or Palearctic. 20 genera are small; Serinus, Carduelis and Carpodacus contain nearly 100 species between them.

Genus Serinus Koch

An essentially African group of small or medium-sized yellow-olive or brown finches with short, stumpy bills: the serins, canaries and Old World seedeaters. Culmen decurved or (in a few species) straight; bill horn-coloured, grey or pink; bill large and stout in some spp., relatively fine and long in a few others. Lowland and montane; open country, forest edge, scrub, grassland; one species associated with papyrus, another with ilala palm. Arboreal, foraging partly in trees, shrubs and herbs although mainly on ground. Nest a neat, open cup, snugly lined, usually in bush or tree. Characters of bill morphology, plumage colour and pattern have a mosaic distribution, such that very different classifications were reached by earlier systematists. Plumage and breeding behaviour characters studied by Nicolai (1960, investigating 9 spp.) and van den Elzen (1985, 18 spp.), and morphological characters by van den Elzen et al. (1987). Van den Elzen (1985) pointed out

455

456

FRINGILLIDAE

that generic limits are not defined by any rules; that, for practical reasons, the lumping of several distinct species-groups into a single genus is currently in vogue; and that the large genus Serinus sens. lat., as it has been widely recognized in recent years, is probably polyphyletic. She recognized 7 groups or genera, much like the arrangement of Hall and Moreau (1970). Sibley and Monroe (1990), Short et al. (1990), Clement et al. (1993) and Dowsett and Dowsett-Lemaire (1993a) have all retained the large genus Serinus sens. lat. Short et al. and Dowsett and Dowsett-Lemaire accepted van den Elzen’s arrangement but reduced her genera to subgenera, for reasons of practicality. We do the same, thinking that overt differences between species and groups are less than those within e.g. Nectarinia sens. lat. (sunbirds), whose species we distributed into multiple genera (see Vol. 6, p. 135). We diagnose the subgenera separately in the following species accounts; we feel that the appearance of alario and the behaviours of totta/symonsi are sufficiently distinct to warrant generic separation as Alario and Pseudochloroptila respectively, and we transfer African ankoberensis from Serinus into the probably derivative Carduelis. Serinus sens. lat. and Carduelis would appear to be closely allied and are difficult to diagnose separately, merging through forms such as [Serinus] pusillus, citrinella and nigriceps and [Carduelis] spinus, ankoberensis and estherae. Little is yet known about species-isolating mechanisms in these finches and, in the absence of molecular studies, the ranking of taxa presents particular difficulties. Forms like (canicollis) flavivertex, (citrinelloides) frontalis, (c.) hypostictus, (atrogularis) xanthopygius, (a.) reichenowi, (donaldsoni) buchanani, (gularis) elgonensis, (g.) reichardi, (g.) striatipectus, (striolatus) whytii, (burtoni) melanochrous, (b.) tanganjicae and Alario (Serinus) (alario) leucolaema have previously been treated variously as subspecies, megasubspecies, allospecies and full species, with a recent tendency to upgrading. We continue that tendency but are aware that we may not have been entirely consistent in our treatment, keeping some difficult taxa down and promoting others. By no means is this the last word on serin classification. 45 spp. (Sibley and Monroe 1990) or 37 (Clement et al. 1993), all breeding in or visiting Africa except canaria (Canary Is), thibetanus (Tibet), menachensis (Arabia) and estherae (Sumatra, Java, Philippines). We recognize 37 in Africa (or 40 globally, including canaria, corsicana and menachensis, excluding thibetanus and estherae which may belong in Carduelis). Subgenera Dendrospiza, Ochrospiza and Crithagra are endemic and Poliospiza nearly so; they embrace 30 spp. in Africa and 1 in Arabia. Of the 7 African spp. in subgenus Serinus, 3 are endemic, 1 (serinus) is N African, Mediterranean and European, and 3 are European or Middle East vagrants to Africa.

Serinus citrinelloides superspecies

Serinus atrogularis superspecies

1

1

1

3 2 2

2,4 1 S. citrinelloides 2 S. frontalis 3 S. hypostictus 4 S. capistratus

3

4

3

1

2 3

1 S. xanthopygius 2 S. reichenowi 3 S. atrogularis

3

3

2

Serinus

Serinus sulphuratus superspecies

Serinus flavigula superspecies

1

3

2

1 1 1,3 2,3

2 1 S. donaldsoni 2 S. buchanani 3 S. sulphuratus

1 S. flavigula 2 S. xantholaemus

Serinus striolatus superspecies

3

Serinus gularis superspecies

1

1

1 1

1 S. striolatus 2 S. whytii

2

2

1,2

1 1 S. gularis 2 S. reichardi

2

2 1

Serinus canicollis superspecies map on p. 460.

457

Plate 29 

Yellow-fronted Canary (p. 479) Serinus mozambicus

S. m. caniceps

S. m. samaliyae







S. m. mozambicus


S. m. barbatus Lemon-breasted Canary (p. 477) Serinus citrinipectus 






Yellow-throated Seedeater (p. 484) Serinus flavigula Salvadori’s Seedeater (p. 486) Serinus xantholaemus

White-bellied Canary (p. 483) Serinus dorsostriatus maculicollis

Ad. Southern Grosbeak-Canary (p. 488) Serinus buchanani


S. s. sulphuratus

S. s. sharpei



Northern Grosbeak-Canary (p. 487) Serinus donaldsoni






Brimstone Canary (p. 489) Serinus sulphuratus

S. f. flaviventris 

Ad. Imm.

Yellow Canary (p. 491) Serinus flaviventris.

S. a. albogularis

S. a. crocopygia

Ad.




S. f. damarensis 

White-throated Canary (p. 494) Serinus albogularis

458

6 in 15 cm

Plate 30

Thick-billed Seedeater (p. 498) Serinus burtoni

Streaky Seedeater (p. 495) Serinus striolatus S. s. graueri

S. b. kilimensis

S. b. burtoni

Tanzania Seedeater (p. 499) Serinus melanochrous

S. s. striolatus

S. r. thomensis Yellow-browed Seedeater (p. 497) Serinus whytii

Príncipe Seedeater (p. 500) Serinus rufobrunneus

Black-eared Seedeater (p. 504) Serinus mennelli

S. r. rufobrunneus




Protea Canary (p. 502) Serinus leucopterus

Streaky-headed Seedeater (p. 506) Serinus gularis S. g. humilis

S. g. benguellensis

S. g. gularis S. g. canicapilla







Yellow-crowned Canary (p. 511) Serinus flavivertex

S. r. reichardi

Juv. Cape Canary (p. 513) Serinus canicollis





Brown-rumped Seedeater (p. 510) Serinus tristriatus

S. r. striatipectus

Reichard’s Seedeater (p. 509) Serinus reichardi Ad.

459 6 in 15 cm

460

FRINGILLIDAE

Serinus canicollis superspecies

1 S. flavivertex 2 S. canicollis

1

2

Subgenus Dendrospiza Roberts

Small, olive and yellow, streaky serins; sexes alike or not; X builds nest, seldom accompanied by Y, displays using Headup and Body-swaying postures, nest rim kept clean, nestling period 18 days (i.e. Serinus (Dendrospiza) scotops; others not known: van den Elzen 1985). Distributed as numerous allopatric populations (some parapatric, others sympatric but only when occupying very different habitats), most looking rather alike. Differences are that YY have forehead, lores, chin and face olive or dusky or black, with or without yellow on forehead or supercilium; YY are slightly or heavily streaked; sexes dimorphic or YY X-plumaged; also differences in bill stoutness and in voice. Courtship postures include Head-up display and Body-swaying. Nestlings defaecate over (not on) rim of nest, which remains clean. There is broad agreement that scotops (South Africa, evergreen forest and thickets) is a good, independent species; capistratus (W-central Africa, forest clearings, riparian thickets) has generally been treated as an independent species too, although it is so similar to frontalis (differing mainly in bill form, robust in capistratus, quite slender in frontalis, hypostictus being intermediate) that we treat it as a 4th allospecies in the citrinelloides/frontalis/hypostictus complex. The papyrus-dwelling koliensis (papyrus-edged lakes from Kivu to Kyogo and Kisumu) was described as a race of capistratus but nowadays is always accorded species status; Sibley and Monroe (1990) make it an allospecies of S. scotops but we keep it independent. The S. citrinelloides superspecies inhabits forest edge, grassland and bracken-briar in eastern African mountains. Pending better knowledge, we follow Sibley and Monroe in treating them as 4 allospecies, notwithstanding that the 2 subspecies of S. citrinelloides and the 2 of S. hypostictus are geographically interdigitated (see superspecies map). Endemic. 6 species.

Plate 28

Serinus citrinelloides Ru ¨ ppell. African Citril. Serin d’Abyssinie.

(Opp. p. 411)

Serinus citrinelloides Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 95, pl. 34, fig. 1; Simen, Abyssinia. Forms a superspecies with S. frontalis, S. hypostictus and S. capistratus. Range and Status. Endemic resident, Eritrea, Ethiopia and E Africa. Eritrea, locally common on central plateau at

2200–2300 m (Zinner 2001), south to near Guna Guna at c. 2450 m. Ethiopia, locally common in W and SE Highlands,

460

FRINGILLIDAE

Serinus canicollis superspecies

1 S. flavivertex 2 S. canicollis

1

2

Subgenus Dendrospiza Roberts

Small, olive and yellow, streaky serins; sexes alike or not; X builds nest, seldom accompanied by Y, displays using Headup and Body-swaying postures, nest rim kept clean, nestling period 18 days (i.e. Serinus (Dendrospiza) scotops; others not known: van den Elzen 1985). Distributed as numerous allopatric populations (some parapatric, others sympatric but only when occupying very different habitats), most looking rather alike. Differences are that YY have forehead, lores, chin and face olive or dusky or black, with or without yellow on forehead or supercilium; YY are slightly or heavily streaked; sexes dimorphic or YY X-plumaged; also differences in bill stoutness and in voice. Courtship postures include Head-up display and Body-swaying. Nestlings defaecate over (not on) rim of nest, which remains clean. There is broad agreement that scotops (South Africa, evergreen forest and thickets) is a good, independent species; capistratus (W-central Africa, forest clearings, riparian thickets) has generally been treated as an independent species too, although it is so similar to frontalis (differing mainly in bill form, robust in capistratus, quite slender in frontalis, hypostictus being intermediate) that we treat it as a 4th allospecies in the citrinelloides/frontalis/hypostictus complex. The papyrus-dwelling koliensis (papyrus-edged lakes from Kivu to Kyogo and Kisumu) was described as a race of capistratus but nowadays is always accorded species status; Sibley and Monroe (1990) make it an allospecies of S. scotops but we keep it independent. The S. citrinelloides superspecies inhabits forest edge, grassland and bracken-briar in eastern African mountains. Pending better knowledge, we follow Sibley and Monroe in treating them as 4 allospecies, notwithstanding that the 2 subspecies of S. citrinelloides and the 2 of S. hypostictus are geographically interdigitated (see superspecies map). Endemic. 6 species.

Plate 28

Serinus citrinelloides Ru ¨ ppell. African Citril. Serin d’Abyssinie.

(Opp. p. 411)

Serinus citrinelloides Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 95, pl. 34, fig. 1; Simen, Abyssinia. Forms a superspecies with S. frontalis, S. hypostictus and S. capistratus. Range and Status. Endemic resident, Eritrea, Ethiopia and E Africa. Eritrea, locally common on central plateau at

2200–2300 m (Zinner 2001), south to near Guna Guna at c. 2450 m. Ethiopia, locally common in W and SE Highlands,

460

FRINGILLIDAE

Serinus canicollis superspecies

1 S. flavivertex 2 S. canicollis

1

2

Subgenus Dendrospiza Roberts

Small, olive and yellow, streaky serins; sexes alike or not; X builds nest, seldom accompanied by Y, displays using Headup and Body-swaying postures, nest rim kept clean, nestling period 18 days (i.e. Serinus (Dendrospiza) scotops; others not known: van den Elzen 1985). Distributed as numerous allopatric populations (some parapatric, others sympatric but only when occupying very different habitats), most looking rather alike. Differences are that YY have forehead, lores, chin and face olive or dusky or black, with or without yellow on forehead or supercilium; YY are slightly or heavily streaked; sexes dimorphic or YY X-plumaged; also differences in bill stoutness and in voice. Courtship postures include Head-up display and Body-swaying. Nestlings defaecate over (not on) rim of nest, which remains clean. There is broad agreement that scotops (South Africa, evergreen forest and thickets) is a good, independent species; capistratus (W-central Africa, forest clearings, riparian thickets) has generally been treated as an independent species too, although it is so similar to frontalis (differing mainly in bill form, robust in capistratus, quite slender in frontalis, hypostictus being intermediate) that we treat it as a 4th allospecies in the citrinelloides/frontalis/hypostictus complex. The papyrus-dwelling koliensis (papyrus-edged lakes from Kivu to Kyogo and Kisumu) was described as a race of capistratus but nowadays is always accorded species status; Sibley and Monroe (1990) make it an allospecies of S. scotops but we keep it independent. The S. citrinelloides superspecies inhabits forest edge, grassland and bracken-briar in eastern African mountains. Pending better knowledge, we follow Sibley and Monroe in treating them as 4 allospecies, notwithstanding that the 2 subspecies of S. citrinelloides and the 2 of S. hypostictus are geographically interdigitated (see superspecies map). Endemic. 6 species.

Plate 28

Serinus citrinelloides Ru ¨ ppell. African Citril. Serin d’Abyssinie.

(Opp. p. 411)

Serinus citrinelloides Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 95, pl. 34, fig. 1; Simen, Abyssinia. Forms a superspecies with S. frontalis, S. hypostictus and S. capistratus. Range and Status. Endemic resident, Eritrea, Ethiopia and E Africa. Eritrea, locally common on central plateau at

2200–2300 m (Zinner 2001), south to near Guna Guna at c. 2450 m. Ethiopia, locally common in W and SE Highlands,

Serinus citrinelloides

Serinus citrinelloides

Rift Valley and in W and S Ethiopia; common in Addis Ababa gardens; commoner in higher than lower elevations; in Yavello and Alghe areas, common in evergreenforested highlands and in drier, juniper-clad highlands; in NW, occurs down to 2050 m and common at 2150–2750 m in Simen Mts, Waibela, Big Abbai, Axum, Tigrai and Dangila. Kenya, frequent to common in highlands from Meru District and Mt Kenya to Nairobi, west to Molo, Kericho, Nandi, Sotik, Rapogi, Kakamega and Siaya; commonest wherever some Aspilia and Cosmos are plentiful. Description. S. c. citrinelloides Ru ¨ ppell: highlands of Eritrea and Ethiopia. ADULT Y: variable, some being black-faced with sparse streaking on underparts, and others grey-faced with heavier streaking on underparts (very like Y and X S. hypostictus hypostictus, q.v.). The situation remains to be clarified. Grey-faced YY are certainly post-juv.; they could be immature, but occur at all times of year. Black-faced Y: forehead black, rest of upperparts bright yellowish olive-green, yellower on rump, crown to hindneck, mantle, back and scapulars with fine blackish streaks. Tail feathers blackish, narrowly edged greenish yellow. Narrow yellow superciliary stripe above and behind eye; lores to behind and below eye, forecheeks and chin black, merging with dark olive-green hindcheeks and ear-coverts; sides of neck olive-green. Throat and rest of underparts bright yellow with faint olive tinge, sides of breast suffused olive-green, and flanks with a few fine dusky streaks. Remiges blackish brown, primaries and secondaries narrowly edged greenish yellow with tips fringed pale buff, tertials edged and tipped greenish yellow; primary coverts and alula blackish, finely fringed greenish yellow; greater and median coverts blackish brown, edged and tipped yellow to form 2 wingbars; lesser coverts broadly tipped yellowish olive-green. Underwing-coverts and axillaries greenish yellow. Bill moderately long (A), dark horn above, pale horn, yellowish or flesh below; eyes brown; legs light brown or pinkish brown. (Greyfaced Y): differs in having lores to behind eye, cheeks and chin dark grey; upperparts slightly duller, more prominently streaked;

breast and flanks more streaked. ADULT X: top of head and upperparts olive-green, streaked blackish brown, more yellowish with fewer streaks on rump and uppertail-coverts. Poorly marked, narrow yellowish superciliary stripe; lores to behind eye dark olive-green, merging with paler dull olive-green cheeks and earcoverts. Underparts pale greenish yellow, whiter on chin, paler on belly and undertail-coverts, chin to breast and flanks streaked blackish brown. Tail and wings as ad. Y but feathers browner; yellowish edges of tertials and yellow tips of greater, median and lesser coverts paler and narrower. SIZE (10 black-faced YY, 10 XX): wing, Y 66–72 (684), X 63–68 (666); tail, Y 46–48 (468), X 44–50 (463); bill, Y 12–14 (131), X 12–13 (125); tarsus, Y 15–16 (152), X 14–16 (149). Wing of 9 grey-faced YY: 66–72 (688). WEIGHT: (Ethiopia) Y (n ¼ 26) 110–147 (129), X (n ¼ 15) 116– 134 (125). IMMATURE: Y very like ad. Y and X S. h. hypostictus, q.v. Juvenile buffish brown above with broad dark brown streaks; sides of head buffish brown with a few dusky streaks; underparts pale buff, tinged yellow from breast to undertail-coverts, chin to breast and flanks with broad dark brown streaks. Wings and tail browner than ad. X, flight feathers narrowly edged olive-yellow, edges of tertials and tips of greater and median coverts buff. NESTLING: at 3 days, skin pinkish brown, quite long tufts of down on head, middle of back, base of wings, and pelvis. S. c. kikuyuensis (Neumann): Kenya highlands. Y has broader, conspicuous yellow superciliary stripe, sometimes crossing forehead behind black frontal band; upperparts greener than in nominate race, and more heavily streaked. X slightly browner (less green) than nominate race, more broadly streaked above and below. Juvenile more olive above and yellow below (less buffish) than nominate race. WEIGHT: Y (n ¼ 31) 117–151 (135), X (n ¼ 36) 117–158 (136); unsexed (n ¼ 324) av. 132 (SD 241, SE 067); YY and XX lightest at onset of moulting and significantly heavier at onset of breeding, when they lay down 15 g of fat (Gichuki 1988). TAXONOMIC NOTE: see under S. hypostictus.

Field Characters. Length 11–12 cm. Y with black forehead, face and chin and plain bright yellow underparts is the only black-faced canary in its range; yellow supercilium thin in Ethiopia, broad in Kenya, sometimes meeting in narrow band above black forehead. Streaked X extremely similar to race brittoni of Southern Citril S. hypostictus, which it meets in W Kenya; rather browner above, with poorly marked yellow supercilium, dark lores and eyeline grading into olive-green cheeks and ear-coverts. X Ethiopian Siskin S. nigriceps has sooty face and forehead, strongly marked black and yellow wing. Voice. Tape-recorded (104, KEI). Song, 4 sweet, pure piping notes, ‘way-pee-woo-pee’, repeated at intervals of 3–4 s. Song often followed by or interwoven with longer complex series; dawn song a faster, high-pitched twittering; calls include rising ‘t’tweee’ and soft chittering in flight (Zimmerman et al. 1996). General Habits. Highlands, inhabiting scrub on mountain slopes and edges of wet woodland (Eritrea), wooded

461

462

FRINGILLIDAE

valleys, rank growth along streams, rivers and lakeshores, moist secondary bush, clearings in forest and bamboo, eucalyptus plantations, gardens, shrubby regrowth, and cultivated land especially with mature oil-seed crops (Kenya). Occurs in small flocks, often of 10–30 birds, including some still breeding (Lynes 1934). Sings from tops of tall trees, and of shrubs surrounding boulder outcrops. Forages on ground, eating seeds, and commonly takes seeds from dried flowers of Aspilia and from ripe heads of Cosmos which have bent over; sometimes pursues and catches flying termites. Often feeds amongst garden flowers and wayside weeds, and takes fruits from shrubs and herbs. Very skilful in extracting kernels; takes a seed in bill and passes it rapidly from side to side until husk drops away. Feeding flock put to flight soon returns to ground or to patch of flowers and continues to forage. Resident but wanders; irregular visitor in Kerio Valley, Kenya, from higher altitudes in early rains (Wilson and Wilson 1994). Moults whole plumage once in 11–13 months, body moult starting and finishing earlier than primary moult; moult cycle takes 16 weeks, primary moult 14–15 weeks, finishing near onset of breeding; YY moult rather earlier than XX (Gichuki 1988). Food. Seeds, particularly of Aspilia and Cosmos (van Someren 1956) and Amaranthus hybridus, Nicandra physalloides, Erucastrum arabicum and Acacia spp., also rapeseed Brassica and sunflower Helianthus annus; Salvia. Ripe fruits, mainly berries, of Cordia spp., Grewia similis, Lantana trifolia, L. camela, Solanum nigrum, Schinus molle, also bits of strawberry Fragaria, raspberry Rubus and grape Vitis; buds and leaves of Rumex abyssinicus and cultivated legumes (Gichuki 1988). Insects, including ants, beetles, grasshopper nymphs, lepidopteran larvae, moths, and termite alates; taken throughout year. S. citrinelloides and/ or Southern Citril S. hypostictus brittoni fond of seeds of Bidens pilosus, sunflowers, thistles and grasses (Clement et al. 1993). Seeds make up 390% by weight of diet, fruits 324% leafy shoots, fruit buds and flower buds 105%, and insects 114% (remaining 67% unidentified: Gichuki 1988). Breeding Habits. Solitary nester. Double-brooded (in Kenya in the long rains); one pair nested again a month after 1st brood fledged. NEST: a deep or shallow cup made of neatly woven fine grass stems and rootlets (B), thickly lined with yellowbrown plant down; rim of nest bound with cobweb; outside may be decorated with lichen and cocoons, attached with cobweb. Ext. diam. c. 70, int. diam. 48 and c. 45, int. depth c. 35. Sited toward tip of upright branch 2– 45 m up in Rhus or Acacia tree (Kenya) or, quite often (in NW Ethiopia) in tall thistle-like plants. Built by both sexes, X doing most of the work, Y often perching nearby. EGGS: 2–4 (av., Kenya, 24, Gichuki 1988). Elongate; white or very pale blue, with sparse, irregular, small, reddish or dark brown spots, mostly at large end. SIZE: (n ¼ 2) 17  125, 175  115. LAYING DATES: Ethiopia, Sept–Dec (and sings well into July); S Sudan, Oct–Mar; Kenya, Nakuru district, in

period July 1983 to Oct 1984, 54 clutches and broods were in all months except Feb and Sept 1984, mainly in Oct– Dec 1983 (especially Dec) and May–July 1984 (especially May), but not apparent why breeding is intermittent (Gichuki 1988); E Africa, Region A, Oct, Dec, May (some records may refer to S. h. brittoni), Region B, Mar–July and Nov–Dec, Region D (including some Arusha, Tanzania, records of S. h. hypostictus) Mar–Aug and Nov–Jan (mainly Apr–May, 15 out of 34 records; breeds mainly in long rains (Brown and Britton 1980) but see Kenya records above). INCUBATION: almost entirely by X. She leaves nest readily during 1st few days; later, she sits very close and is off nest only for very short spells. Y often feeds her on nest, X begging with quivering wings and taking offered food from his bill a small portion at a time. Period 12–13 days (Kenya). DEVELOPMENT AND CARE OF YOUNG: feather quills show from 4th or 5th day; feathers show green by 7th day. Young closely brooded by X for first 3 days, when Y feeds her and she passes some food to the young. In her absence, Y feeds chicks and broods them. Brood of 3 fills nest at day 10; to shelter them from sun, X stands on nest rim, fluffs out breast feathers, lowers body and partly opens wings. Y and X utter faint ‘seepsiip’ call just before arriving at nest and just before leaving. Young fed by regurgitation from parent’s crop into its bill, mouthful being delivered in rotation, with small portions. Later, X makes 6–8 regurgitations per visit. Nestling period: once 15 days, the 3 chicks leaving nest within short time of each other and joining one or other parent in branches below; both parents feed chicks as soon as they flutter down from nest, and, later, marshall the 3 together. One brood remained in nest tree for a week after leaving nest. Key References Clement, P. et al. (1993), Gichuki, N.N. (1988), Lynes, H. (1934), van Someren, V.G.L. (1956).

Serinus frontalis

463

Serinus frontalis Reichenow. Western Citril. Serin a ` diade`me.

Plate 28

Serinus citrinelloides frontalis Reichenow, 1904. Vo ¨gel Afrikas, 3, p. 275; Victoria-Niansa area (Butumbi, Bukoba, Karevia, Kafuro); Nandi, ‘‘Chiroko’’, L. Nyasa; Kenia and L. Kivu; type from L. Kivu, fide W.L. Sclater, 1930, Syst, Avium Aethiopicarum, p. 826.

(Opp. p. 411)

Forms a superspecies with S. citrinelloides, S. hypostictus and S. capistratus. Range and Status. Endemic resident, highlands of E Zaı¨re and from Uganda to N Zambia. Zaı¨re, common on Lendu Plateau, up to 2070 m on Mt Aboro; in Rwenzori Mts up to 2130 m; near Butembo and Luofu and low and mid levels on Kivu Volcanoes (once at Kabara, 3350 m); highlands around L. Kivu; Itombwe, very common, at 1070–2060 m; Marungu, at 1590–2040 m (common around Pepa; also at Kasiki, Lyapenda, Mukuli, Ketendwe, Sambwe and Kasiki); record at Mukumbi in S (R.J. Dowsett, pers. comm.). Uganda, uncommon in Kalinzu, Malabigambo, Namalala, Kasyoha-Kitomi, Itwara and Bugoma Forests, east to Sezibwa R., Entebbe and L. Kyoga, and around N end of L. Albert, south to Masindi. Rwanda and Burundi, widespread east to about 30
E; at all altitudes in Nyungwe Forest (Rwanda) and Bururi Forest (Burundi). W Tanzania, known from 4 areas: Bukoba on W shores of L. Victoria; between Biharamulo and Burundi border; Matengo highlands (Myangayanga, Ugano, Mtindi, Lipumba; treated as S. (c.) hypostictus by Sassi and Zimmer 1941); and Ufipa Plateau, where widespread (N. E. Baker, pers. comm.). Zambia, only in Northern Prov., at Mbala, with sight record from Old Fife. Description. ADULT Y: usually a narrow black frontal band; behind this a broad yellow band across forehead extending back as superciliary stripe to behind eye. Rest of upperparts yellowish olive-green, yellower on rump and uppertail-coverts, crown to hindneck, mantle, back and scapulars with narrow blackish streaks. Tail feathers blackish brown, outer webs edged yellow, T1 with broad yellow-tinged border on both webs. Lores to behind and below eye, forecheeks and chin black, merging with dark olive green ear-coverts and hindcheeks; sides of neck lighter olive-green. Throat and rest of underparts bright yellow, sides of breast washed olive-green. Remiges blackish brown, primaries and secondaries narrowly edged greenish yellow with tips fringed pale grey-buff, tertials broadly edged and tipped yellowish green, primary coverts and alula blackish, finely fringed yellowish green; greater coverts blackish, edged green and broadly tipped yellow; median coverts blackish, broadly tipped yellowish green; lesser coverts yellowish green. Underwing-coverts and axillaries greenish yellow. Bill rather long and pointed, slenderer than bills of S. citrinipectus and S. hypostictus, upper mandible horn brown, lower pale yellowish flesh; eyes brown; legs light brown or brownish flesh. ADULT X: forehead yellow, extending back as superciliary stripe; crown to mantle, back and scapulars light olive-green, sharply streaked blackish brown; rump and uppertailcoverts more yellowish, unstreaked. Lores, cheeks and ear-coverts dark olive-green; sides of neck paler green. Underparts pale yellow, tinged green, with a few diffuse dusky streaks on flanks and sometimes on breast; breast sides washed olive-green. Tail and wings as ad. Y, but green edges to tertials and pale yellowish tips to greater and median coverts narrower. SIZE (10 YY, 10 XX): wing, Y 62–67 (641), X 60–67 (636); tail, Y 45–47 (459), X 43– 51 (458); bill, Y 13–14 (137), X 125–14 (133); tarsus, Y 14–15 (145), X 135–15 (145). WEIGHT: 1 Y 12, 1 X 12, 1 unsexed 13. IMMATURE: juv. has yellow forehead; rest of upperparts olivebrown with broad dark brown streaks; underparts olive-yellow

Serinus frontalis

with a few short narrow brown streaks on breast and flanks; flight feathers edged olive-yellow; edges of tertials and tips to greater and median coverts yellowish buff. TAXONOMIC NOTE: see under S. hypostictus.

Field Characters. Length 11–12 cm. Y has narrow black frontal band and broad yellow band above it, extending back as supercilium, face and chin black; very similar Y African Citril S. citrinelloides has narrow yellow frontal band, but the 2 do not quite meet (Zimmerman et al. 1996, contra Clement et al. 1993). X distinguished from all other canary XX in being almost identical to Y, with largely unstreaked yellow underparts, black face with olive wash. Closely approaches Black-faced Canary S. capistratus in NE Zambia; for comparison, see that species. Voice. Tape-recorded (C, KAE P). Song very different from African Citril, a high-pitched jingle at moderate speed, with short sweet notes mixed with thin ‘sit-sit’ and frequent buzzy ‘zwee’. Call, ‘chit-chit’. General Habits. Frequents open grassy and weedy places, often near farmland, bracken-briar, secondary forest and fallow ground (Zaı¨re); marshes, heath, and large forest clearings (Nyungwe, Rwanda). Occurs in parties of 4–8 birds. Forages at flowers in bracken-briar; takes seeds from ground and pulls seeds from heads of thistles; often at mid

464

FRINGILLIDAE

levels in moist forest trees, and sometimes joins mixedspecies foraging flocks which pass through diverse habitats (Prigogine 1971). Food. Seeds. Fond of seeds of Senecio montuosum (Nyungwe, Rwanda) and ‘may take small fruit on the margin of forest (Urera) and seeds of Hagenia’ (DowsettLemaire 1990). Eats small seeds amid tall grasses and patches of Bidens, Galinsoga and tall thistles (Chapin 1954). Insects and small seeds in a stomach. Breeding Habits. Solitary nester, evidently monogamous and territorial.

NEST: a small cup made of grass and plant fibres; once largely of white plant down, with dried leaves on outside and a lining of fine, brownish plant fibres; some nests have lichen fixed to outer surface with spider web. Sited 2–5 m up in fork of bush or tree, or in growing bunch of bananas. EGGS: not known to differ significantly from those of S. citrinelloides. LAYING DATES: Zaı ¨re, Rwenzori (Kilongi), (nestlings, Nov; ads with large gonads, Dec), Kivu, (gonads enlarged June; bird just fledged in mid June), Marungu, (breeding condition Feb); Uganda, Mar–July, Nov. Nothing further known.

Key Reference Chapin, J.P. (1954).

Plate 28

Serinus hypostictus (Reichenow). Southern Citril. Serin est-africain.

(Opp. p. 411)

Spinus citrinelloides hypostictus Reichenow, 1904. Vo ¨gel Afrikas, 3, p. 275; Kilimandscharo, Grossaruscha, Ngomingi, Tembo, Rupira, Somba; type from Moshi, Kilimanjaro, fide W.L. Sclater, 1930. Syst. Avium Aethiopicarum, p. 826. Forms a superspecies with S. citrinelloides, S. frontalis and S. capistratus. Range and Status. Endemic resident with 2 populations, brittoni in SW Sudan and W Kenya and hypostictus from S ˆ i. Kenya to S Malaw Sudan, very common up to 2400 m in Imatong, Dongotona and Didinga Mts, moving to lower altitudes in wet season. Kenya, brittoni frequent to common at 1200– 2800 m from Mt Elgon, Siaya, Kitale, Kapenguria and Kongelai to Kakamega; hypostictus from Lolgorien, NW Mara Game Res., and Loita and Nguruman Hills, also Chyulu Hills. Tanzania, Loliondo to Crater and Mbulu Highlands, Arusha Nat. Park, Mt Lolkissale, Mt Kilimanjaro at 1700–2300 m on W slopes and below 1600 m on S slopes (Cordeiro 1994), Pare Mts at 1400–1800 m, Usambaras, Ukagurus, Ulugurus (down to 400 m), Udzungwa, Iringa, Njombe and Mbeya highlands west to about Mbozi, Mt Mahenge, and hills near Lipumba and Liuli in extreme SW. Zambia, Nyika Plateau in NE and Chipata in SE (Bull. ˆ i, widespread but not Afr. Bird Club 3, 1996, 140). Malaw common: 910 m up to 2130 m on Nyika Plateau, 1830 m on Misuku Hills, Mafinga and Viphya Mts, 1670 m at Dedza; ˆi Chintheche, Mulanje, Mitongwe, Namlenga and Malaw Hill, at c. 560–670 m; Namwera. Mozambique, old record(s) ˆ i border (Hall from highlands about 15
150 S near Malaw and Moreau 1970). ˆ i. Description. S. h. hypostictus (Reichenow): SE Kenya to Malaw ADULT Y: top of head and upperparts olive-green, narrowly streaked blackish, brighter more yellowish green on rump. Tail feathers blackish, narrowly edged olive-green. Superciliary stripe indistinct or absent; lores to behind eye, forecheeks and chin dark grey, merging with lighter grey hind cheeks and ear-coverts; sides of neck greyish olive, streaked dusky. Underparts yellow, tinged greenish, with dull olive-green suffusion on sides of breast and flanks; throat and upper breast with short blackish streaks, sides of breast and flanks with longer dusky shaft streaks. Remiges blackish brown, edged olive-green, tertials tipped yellowish; primary coverts and alula blackish brown, fringed olive-green; greater coverts blackish brown, edged olive-green

Serinus hypostictus

and tipped yellowish; median coverts blackish, tipped yellowish olive-green; lesser coverts olive-green. Underwing coverts and axillaries dull olive-green. Bill horn brown above with darker culmen, whitish or pale horn below; eyes dark brown; legs dull brown. ADULT X: similar to ad. Y, but breast and flanks usually more heavily streaked. SIZE (10 YY, 10 XX): wing, Y 63–67 (656), X 61–67 (642); tail, Y 43–49 (468), X 42–48 (454); bill, Y 125– 14 (132), X 125–135 (130); tarsus, Y 14–155 (149), X 14–15 (144). WEIGHT: (S Tanzania) Y (n ¼ 10) 115–14 (131), X (n ¼ 12) 10–155 (129). IMMATURE: juv. olive-brown above, streaked blackish brown, tinged greener on rump; inconspicuous superciliary stripe buffish

464

FRINGILLIDAE

levels in moist forest trees, and sometimes joins mixedspecies foraging flocks which pass through diverse habitats (Prigogine 1971). Food. Seeds. Fond of seeds of Senecio montuosum (Nyungwe, Rwanda) and ‘may take small fruit on the margin of forest (Urera) and seeds of Hagenia’ (DowsettLemaire 1990). Eats small seeds amid tall grasses and patches of Bidens, Galinsoga and tall thistles (Chapin 1954). Insects and small seeds in a stomach. Breeding Habits. Solitary nester, evidently monogamous and territorial.

NEST: a small cup made of grass and plant fibres; once largely of white plant down, with dried leaves on outside and a lining of fine, brownish plant fibres; some nests have lichen fixed to outer surface with spider web. Sited 2–5 m up in fork of bush or tree, or in growing bunch of bananas. EGGS: not known to differ significantly from those of S. citrinelloides. LAYING DATES: Zaı ¨re, Rwenzori (Kilongi), (nestlings, Nov; ads with large gonads, Dec), Kivu, (gonads enlarged June; bird just fledged in mid June), Marungu, (breeding condition Feb); Uganda, Mar–July, Nov. Nothing further known.

Key Reference Chapin, J.P. (1954).

Plate 28

Serinus hypostictus (Reichenow). Southern Citril. Serin est-africain.

(Opp. p. 411)

Spinus citrinelloides hypostictus Reichenow, 1904. Vo ¨gel Afrikas, 3, p. 275; Kilimandscharo, Grossaruscha, Ngomingi, Tembo, Rupira, Somba; type from Moshi, Kilimanjaro, fide W.L. Sclater, 1930. Syst. Avium Aethiopicarum, p. 826. Forms a superspecies with S. citrinelloides, S. frontalis and S. capistratus. Range and Status. Endemic resident with 2 populations, brittoni in SW Sudan and W Kenya and hypostictus from S ˆ i. Kenya to S Malaw Sudan, very common up to 2400 m in Imatong, Dongotona and Didinga Mts, moving to lower altitudes in wet season. Kenya, brittoni frequent to common at 1200– 2800 m from Mt Elgon, Siaya, Kitale, Kapenguria and Kongelai to Kakamega; hypostictus from Lolgorien, NW Mara Game Res., and Loita and Nguruman Hills, also Chyulu Hills. Tanzania, Loliondo to Crater and Mbulu Highlands, Arusha Nat. Park, Mt Lolkissale, Mt Kilimanjaro at 1700–2300 m on W slopes and below 1600 m on S slopes (Cordeiro 1994), Pare Mts at 1400–1800 m, Usambaras, Ukagurus, Ulugurus (down to 400 m), Udzungwa, Iringa, Njombe and Mbeya highlands west to about Mbozi, Mt Mahenge, and hills near Lipumba and Liuli in extreme SW. Zambia, Nyika Plateau in NE and Chipata in SE (Bull. ˆ i, widespread but not Afr. Bird Club 3, 1996, 140). Malaw common: 910 m up to 2130 m on Nyika Plateau, 1830 m on Misuku Hills, Mafinga and Viphya Mts, 1670 m at Dedza; ˆi Chintheche, Mulanje, Mitongwe, Namlenga and Malaw Hill, at c. 560–670 m; Namwera. Mozambique, old record(s) ˆ i border (Hall from highlands about 15
150 S near Malaw and Moreau 1970). ˆ i. Description. S. h. hypostictus (Reichenow): SE Kenya to Malaw ADULT Y: top of head and upperparts olive-green, narrowly streaked blackish, brighter more yellowish green on rump. Tail feathers blackish, narrowly edged olive-green. Superciliary stripe indistinct or absent; lores to behind eye, forecheeks and chin dark grey, merging with lighter grey hind cheeks and ear-coverts; sides of neck greyish olive, streaked dusky. Underparts yellow, tinged greenish, with dull olive-green suffusion on sides of breast and flanks; throat and upper breast with short blackish streaks, sides of breast and flanks with longer dusky shaft streaks. Remiges blackish brown, edged olive-green, tertials tipped yellowish; primary coverts and alula blackish brown, fringed olive-green; greater coverts blackish brown, edged olive-green

Serinus hypostictus

and tipped yellowish; median coverts blackish, tipped yellowish olive-green; lesser coverts olive-green. Underwing coverts and axillaries dull olive-green. Bill horn brown above with darker culmen, whitish or pale horn below; eyes dark brown; legs dull brown. ADULT X: similar to ad. Y, but breast and flanks usually more heavily streaked. SIZE (10 YY, 10 XX): wing, Y 63–67 (656), X 61–67 (642); tail, Y 43–49 (468), X 42–48 (454); bill, Y 125– 14 (132), X 125–135 (130); tarsus, Y 14–155 (149), X 14–15 (144). WEIGHT: (S Tanzania) Y (n ¼ 10) 115–14 (131), X (n ¼ 12) 10–155 (129). IMMATURE: juv. olive-brown above, streaked blackish brown, tinged greener on rump; inconspicuous superciliary stripe buffish

Serinus capistratus yellow; sides of head to chin grey-brown; throat, breast and flanks yellowish buff with dusky streaks; belly pale yellow; tail and wings as ad., but flight feathers edged pale olive; edges of tertials and tips to greater and median coverts yellowish or tawny-buff. S. h. brittoni Traylor: S Sudan and W Kenya. Differs from nominate race in having distinct narrow yellow superciliary stripe; forehead feathers edged yellow; forecheeks more olivegrey, hind cheeks greener; chin pale yellow, streaked blackish like throat; upperparts rather brighter olive-green; underparts, including flanks, more heavily streaked. Wing, Y (n ¼ 3) 68–69 (683), X (n ¼ 5) 65–68 (667). WEIGHT: (S Sudan) unsexed (n ¼ 51) 12–15 (132). TAXONOMIC NOTE: hypostictus and frontalis have often been regarded as subspecies of S. citrinelloides, but were treated as separate species by Wolters (1979–1982), van den Elzen (1985) and Sibley and Monroe (1990). Dowsett and Dowsett-Lemaire (1993b) found no significant differences between the calls (series of 3–4 (–6) clear whistles, rising in pitch; not the songs) of ˆ i, with frontalis (in Rwanda) and hypostictus (throughout Malaw some dialectical variations), so they concluded that frontalis cannot be maintained as a species distinct from citrinelloides, ‘and the same applies to hypostictus’. S. h. brittoni marginally overlaps S. c. kikuyuensis in Siaya area, Kenya, where both birds have been recorded singing in the same garden in Nandi. As in the closely related S. koliensis, YY of S. hypostictus (particularly S. h. brittoni) are X-plumaged. For these reasons we treat S. hypostictus and S. citrinelloides as separate species. We raise frontalis to species level too, on the grounds of bill profile and facial appearance which differ sharply from those of citrinelloides.

Field Characters. Length 11–12 cm. Y differs from other black-faced canaries in having streaked underparts and forehead (no black or yellow); nominate race (S Kenyaˆ i) has grey chin and face with only a hint of a yellow Malaw supercilium; brittoni (W Kenya) overlaps Papyrus Canary S. koliensis, distinguished by yellow chin, pale lores, greenish face, and yellow supercilium.

465

Voice. Tape-recorded (86, 102, 104, B, C, LEM). Rapid tinkling song like Yellow-crowned Canary S. flavivertex but includes buzzy ‘zwee’ and liquid ‘woy’ notes. Another song, 3–4 sweet notes, ‘see-si-see-woo’ (last note lower), or ascending ‘way-pee-tee’, like African Citril S. citrinelloides but higher-pitched. Alarm, soft ‘t’t’tee’; soft twittering in flight. General Habits. Inhabits forest clearings, secondary growth and cultivation (S Sudan), edges of rank growth ˆ i), and also along streams and bracken-briar (Malaw Eucalyptus, Cupressus and Pinus caribea plantations in S and areas with food-plant Cosmos. Frequently forages amongst banana plants (W Kenya, Zimmerman et al. 1996). Having previously been regarded as a race of the better-known S. citrinelloides, few of its habits have been recorded separately, but they are thought to be much the same in both species. ˆ i). S. h. Food. Seeds, including those of Cosmos (Malaw brittoni and/or African Citril S. citrinelloides fond of seeds of Bidens pilosus, sunflowers, thistles and grasses (Clement et al. 1993). Breeding Habits. Many nests have been found, but breeding habits have not been studied separately from those of S. citrinelloides. It nests amongst banana Musa fruits (Zimmerman et al. 1996). LAYING DATES: E Africa, Region D (including some Nairobi, Kenya, records of S. citrinelloides), Mar–Aug and Nov–Jan (mainly Apr–May, 15 out of 34 records; breeds mainly in long rains); Tanzania, Feb, Apr; Zambia, ˆ i, Sept, Nov, Dec, Feb (gonads active Apr, May); Malaw and Mar. Key Reference Zimmerman, D.A. et al. (1996).

Serinus capistratus (Finsch and Hartlaub). Black-faced Canary. Serin a` face noire.

Plate 28

Crithagra capistrata Finsch and Hartlaub, 1870. Vo ¨g. Ost.-Afr., p. 458; Golungo Alto, 1800–2800 ft, Angola.

(Opp. p. 411)

Forms a superspecies with S. citrinelloides, S. frontalis and S. hypostictus. Range and Status. Endemic resident, W-central Africa. Probably far more widespread than shown. Gabon, record from near Port-Gentil and several in SW; frequent in Re´serve de la Lope´, along Lope´ R., Brazza L., and in Le´koni area. Congo, records from Mayombe and Brazzaville. Zaı¨re, in extreme W, adjacent to Cabinda, and along lower Congo R. upstream to Manyanga, Temvo, Boma (where quite common) and Kinshasa; records near Angola border in Bandundu Prov.; several records in E Kasai Occidental and in Kasai Oriental Prov.; numerous in Kananga region; known in E Haut Zaı¨re Prov. from Baraka, Kivu; and widespread but rather uncommon in Katanga up to c. 1350 m (e.g. Kambove, upper Lufira R., Bunkeya R., Kansenia, Mulungwishi, Upemba Nat. Park, Lubumbashi). Burundi, known from Kajeke, Usumbura, between

Bujumbura and Minago, and Rusizi. ‘The common forest canary in Angola’ (Dean 2000), in Cabinda around Ganda Sundi and to south; locally common from NW Huı´la along escarpment to Gabela and Quitondo, into N Bie´, W Malanje, Cuanza Norte, Uı´ge, NE Moxico, and Lunda Norte; another subspecies in Huambo and SE Benguela. Zambia, fairly common at Chiengi on L. Mweru; elsewhere sparse: Mporokoso, Kawambwa District; Kitwe; Solwezi, Mwombezhi R., Mundwiji Plain, around Mwinilunga town; Ngalulu, Kasempa and Kabompo. Description. S. c. capistratus (Finsch and Hartlaub): range of species except W-central Angola. ADULT Y: forehead black; behind this a yellow band across forecrown which extends back above eye and ear-coverts as broad superciliary stripe; hindcrown to uppertail-coverts and scapulars bright olive-green, crown with

Serinus capistratus yellow; sides of head to chin grey-brown; throat, breast and flanks yellowish buff with dusky streaks; belly pale yellow; tail and wings as ad., but flight feathers edged pale olive; edges of tertials and tips to greater and median coverts yellowish or tawny-buff. S. h. brittoni Traylor: S Sudan and W Kenya. Differs from nominate race in having distinct narrow yellow superciliary stripe; forehead feathers edged yellow; forecheeks more olivegrey, hind cheeks greener; chin pale yellow, streaked blackish like throat; upperparts rather brighter olive-green; underparts, including flanks, more heavily streaked. Wing, Y (n ¼ 3) 68–69 (683), X (n ¼ 5) 65–68 (667). WEIGHT: (S Sudan) unsexed (n ¼ 51) 12–15 (132). TAXONOMIC NOTE: hypostictus and frontalis have often been regarded as subspecies of S. citrinelloides, but were treated as separate species by Wolters (1979–1982), van den Elzen (1985) and Sibley and Monroe (1990). Dowsett and Dowsett-Lemaire (1993b) found no significant differences between the calls (series of 3–4 (–6) clear whistles, rising in pitch; not the songs) of ˆ i, with frontalis (in Rwanda) and hypostictus (throughout Malaw some dialectical variations), so they concluded that frontalis cannot be maintained as a species distinct from citrinelloides, ‘and the same applies to hypostictus’. S. h. brittoni marginally overlaps S. c. kikuyuensis in Siaya area, Kenya, where both birds have been recorded singing in the same garden in Nandi. As in the closely related S. koliensis, YY of S. hypostictus (particularly S. h. brittoni) are X-plumaged. For these reasons we treat S. hypostictus and S. citrinelloides as separate species. We raise frontalis to species level too, on the grounds of bill profile and facial appearance which differ sharply from those of citrinelloides.

Field Characters. Length 11–12 cm. Y differs from other black-faced canaries in having streaked underparts and forehead (no black or yellow); nominate race (S Kenyaˆ i) has grey chin and face with only a hint of a yellow Malaw supercilium; brittoni (W Kenya) overlaps Papyrus Canary S. koliensis, distinguished by yellow chin, pale lores, greenish face, and yellow supercilium.

465

Voice. Tape-recorded (86, 102, 104, B, C, LEM). Rapid tinkling song like Yellow-crowned Canary S. flavivertex but includes buzzy ‘zwee’ and liquid ‘woy’ notes. Another song, 3–4 sweet notes, ‘see-si-see-woo’ (last note lower), or ascending ‘way-pee-tee’, like African Citril S. citrinelloides but higher-pitched. Alarm, soft ‘t’t’tee’; soft twittering in flight. General Habits. Inhabits forest clearings, secondary growth and cultivation (S Sudan), edges of rank growth ˆ i), and also along streams and bracken-briar (Malaw Eucalyptus, Cupressus and Pinus caribea plantations in S and areas with food-plant Cosmos. Frequently forages amongst banana plants (W Kenya, Zimmerman et al. 1996). Having previously been regarded as a race of the better-known S. citrinelloides, few of its habits have been recorded separately, but they are thought to be much the same in both species. ˆ i). S. h. Food. Seeds, including those of Cosmos (Malaw brittoni and/or African Citril S. citrinelloides fond of seeds of Bidens pilosus, sunflowers, thistles and grasses (Clement et al. 1993). Breeding Habits. Many nests have been found, but breeding habits have not been studied separately from those of S. citrinelloides. It nests amongst banana Musa fruits (Zimmerman et al. 1996). LAYING DATES: E Africa, Region D (including some Nairobi, Kenya, records of S. citrinelloides), Mar–Aug and Nov–Jan (mainly Apr–May, 15 out of 34 records; breeds mainly in long rains); Tanzania, Feb, Apr; Zambia, ˆ i, Sept, Nov, Dec, Feb (gonads active Apr, May); Malaw and Mar. Key Reference Zimmerman, D.A. et al. (1996).

Serinus capistratus (Finsch and Hartlaub). Black-faced Canary. Serin a` face noire.

Plate 28

Crithagra capistrata Finsch and Hartlaub, 1870. Vo ¨g. Ost.-Afr., p. 458; Golungo Alto, 1800–2800 ft, Angola.

(Opp. p. 411)

Forms a superspecies with S. citrinelloides, S. frontalis and S. hypostictus. Range and Status. Endemic resident, W-central Africa. Probably far more widespread than shown. Gabon, record from near Port-Gentil and several in SW; frequent in Re´serve de la Lope´, along Lope´ R., Brazza L., and in Le´koni area. Congo, records from Mayombe and Brazzaville. Zaı¨re, in extreme W, adjacent to Cabinda, and along lower Congo R. upstream to Manyanga, Temvo, Boma (where quite common) and Kinshasa; records near Angola border in Bandundu Prov.; several records in E Kasai Occidental and in Kasai Oriental Prov.; numerous in Kananga region; known in E Haut Zaı¨re Prov. from Baraka, Kivu; and widespread but rather uncommon in Katanga up to c. 1350 m (e.g. Kambove, upper Lufira R., Bunkeya R., Kansenia, Mulungwishi, Upemba Nat. Park, Lubumbashi). Burundi, known from Kajeke, Usumbura, between

Bujumbura and Minago, and Rusizi. ‘The common forest canary in Angola’ (Dean 2000), in Cabinda around Ganda Sundi and to south; locally common from NW Huı´la along escarpment to Gabela and Quitondo, into N Bie´, W Malanje, Cuanza Norte, Uı´ge, NE Moxico, and Lunda Norte; another subspecies in Huambo and SE Benguela. Zambia, fairly common at Chiengi on L. Mweru; elsewhere sparse: Mporokoso, Kawambwa District; Kitwe; Solwezi, Mwombezhi R., Mundwiji Plain, around Mwinilunga town; Ngalulu, Kasempa and Kabompo. Description. S. c. capistratus (Finsch and Hartlaub): range of species except W-central Angola. ADULT Y: forehead black; behind this a yellow band across forecrown which extends back above eye and ear-coverts as broad superciliary stripe; hindcrown to uppertail-coverts and scapulars bright olive-green, crown with

466

FRINGILLIDAE

Serinus capistratus

face and yellow supercilium readily distinguished from X Western Citril, which looks like Y, with black face and plain yellow underparts. Voice. Tape-recorded (104, B, F, CART, McVIC). Song very different from Western Citril, a mixture of whistles, rattles and buzzy notes, e.g. ‘tee-ti-tyew, trrrrrr, zhwayzhwee, trrrrrr’; short (for a canary) phrases, with pauses in between, not the continuum typical of other species. Call described as ‘swee swee’ with minor variations (Aspinwall and Beal 1998) or a variety of ‘chissick’ notes (Clement et al. 1993).

dusky shaft streaks, mantle, back and scapulars with more distinct narrow blackish streaks. Tail feathers blackish brown, edged yellowish olive-green. Lores, forecheeks and chin, and narrowly above eye to front of ear-coverts, black, forming large face patch continuous with black frontal band. Rear of ear-coverts and hind cheeks olive-green. Throat and rest of underparts bright yellow, washed greenish on sides of breast and flanks. Remiges blackish brown, primaries and secondaries narrowly edged light olive-green with tips fringed greyish white, tertials broadly edged and tipped bright olive-green; alula and upperwing-coverts blackish brown, greater coverts edged olive-green, these and median coverts broadly tipped yellowish green, lesser coverts tipped yellow. Underwing-coverts and axillaries pale yellow. Bill short and rather stout with curved culmen, pinkish white (A); eyes dark brown; legs brown or pale brown. ADULT X: forehead and forecrown olive-green with a few short blackish streaks and often a yellowish frontal band; rest of upperparts olive-green, narrowly streaked blackish brown except on rump and uppertailcoverts. Supercilium broad, yellow with fine greenish flecking; lores, cheeks and ear-coverts dark olive-green. Underparts (including chin) yellow, slightly duller than in Y, sides of breast and flanks washed olive-green; chin, throat and upper breast with short narrow dusky streaks, lower breast and flanks with faint dusky shaft streaks. Tail and wings as ad. Y. SIZE (10 YY, 7 XX): wing, Y 61–66 (634), X 58–63 (607); tail, Y 40–45 (421), X 39– 42 (404); bill, Y 115–125 (119), X 11–125 (117); tarsus, Y 135–15 (139), X 135–145 (139). IMMATURE: juv. similar to ad., but underparts more buffy (less yellow), more heavily streaked. S. c. hildegardae Rand and Traylor: central Angola (Mts Moco and Soque). Y duller olive-green above than in nominate race, frontal band and superciliary stripe washed greenish.

Field Characters. Length 115 cm. Black-faced Y very similar to Y Western Citril S. frontalis, which it approaches but does not quite meet in NE Zambia; face more extensively black, bill short and stubby (fine and pointed in Western Citril), broad greenish yellow edges to tertials make wing appear less black. X with streaked throat, green

General Habits. Inhabits shrubland and grassland adjacent to forest, edges of roads and clearings in primary and secondary forest and coffee forest (Angola: Dean 2000), also forest around bases of inselbergs; bushy savanna, grassy road verges, forest edges and gallery woods (Gabon); near marshes, on floodplains and in fields bordering lakes (Burundi); open ground with dense, long grass and scattered small trees and dense growth along streams (S Katanga, Zaı¨re); edges of moist evergreen forest and thickets near water (Zambia). Occurs in pairs and flocks, e.g. of a dozen and c. 50 (Burundi) but ‘never more than half a dozen’ in Lower Congo (Chapin 1954). Forages on ground and takes seeds in heads of tall grasses; often flocks mix with estrildines, including Bronze Mannikins Spermestes cucullatus. Y sings for long periods from regularly used song posts, 3–12 m above ground. Frequently caged as singing birds in W Zaı¨re. Food. Seeds of tall grasses and tall weeds. Breeding Habits. NEST: one, not described before being destroyed, probably by a mouse, was 25 m up on a horizontal branch in a cluster of leafy twigs, in a small tree close to a stream; built by X, collecting bents from 200 m away and bringing them to nest site every 3 min; Y accompanied her on collecting trips. EGGS: unknown. LAYING DATES: Gabon, (Jan–Feb and Apr–June); Zaı ¨re, Boma, (breeding season starts in mid Jan), Kananga, Nov (i.e. fledglings early Dec, juvs late Dec; also a very young bird, May), upper Katanga, (nest being built in mid Jan); Angola (locality?), (breeding condition, Mar); Zambia, (gonads active Dec, Feb). Nothing further known. Key References Clement, P. et al. (1993), Dean, W.R.J. (2000), Vincent, A.W. (1949).

Serinus koliensis

467

Serinus koliensis Grant and Mackworth-Praed. Papyrus Canary. Serin du Koli.

Plate 28

Serinus capistratus koliensis Grant and Mackworth-Praed, 1952. Bull. Br. Orn. Club, 72, p. 1; Onyulu’s, Koli River, Lango, Uganda.

(Opp. p. 411)

Range and Status. Endemic resident, papyrus swamps at 900–1600 m in S and SW Uganda, W Kenya, E Zaı¨re, Rwanda, Burundi and NW Tanzania. Uganda, locally common, N shore of L. Victoria from Masaka to Kenyan border; southwest to Ankole and Kigezi and north to c. 2
N in L. Kyogo basin, north to Lango; scarce in Kampala area, rare at Namulonge. Kenya, fairly common from Port Victoria, L. Kanyaboli and Usengi to Kisumu and Kendu Bay on S of Kavirondo (Winam) Gulf, where frequent; isolated areas of papyrus along streams in Siaya, north to Ukwala. Tanzania, near Katera (1
200 S, 30
400 E), 2 km from Rwandan border (Baker and Baker 1994) and along Kagera R. (N. E. Baker, pers. comm.). E Zaı¨re, known only from south and north of L. Kivu, at 1700 m in papyrus marshes on Kivu volcanoes and at 2000 m near Mokoto lakes (Lippens and Wille 1976). Rwanda and Burundi, locally common to abundant as mapped (40 locations, including Rugezi, Mulindi, Luhondo, Bulera, Akanyaru, Nyabarongo, Kibaya, Kagogo; Nyamabuye, Nyakijanda, Nyamushwaga, Ndurumu, Ruvubu, Kayongozi and Malagarazi: Vande weghe 1981). Description. ADULT Y: top of head and upperparts olive-green, yellower on rump and uppertail-coverts, all except uppertailcoverts streaked blackish brown. Tail feathers blackish brown, narrowly edged olive-green. Poorly marked yellowish olive-green superciliary stripe behind eye; lores dusky olive; ear-coverts, cheeks and sides of neck dull olive-green. Underparts yellow, chin to breast and flanks washed olive-green; short blackish streaks from chin to breast, longer narrow dusky streaks on lower breast and flanks. Remiges blackish brown, flight feathers narrowly edged olive-green, tertials more broadly so; alula and larger upperwing-coverts blackish brown, edges of greater coverts broadly olive-green, tips of greater and median coverts yellowish olive-green forming poorly marked wing-bars; lesser coverts olive-green. Underwing-coverts and axillaries pale yellow. Bill stout with curved culmen, horn or flesh above, whitish below; eyes blackish brown; legs flesh. ADULT X: similar to ad. Y, but duller olive-green above, especially on crown; sides of head greyer; blackish streaks bolder from chin to breast and present also on flanks. SIZE (3 YY, 1 X): wing, Y 64–66 (647), X 64; tail, Y 43–48 (453), X 48; bill, Y 13–14 (135), X 14; tarsus, Y 145– 165 (153), X 155. WEIGHT: W Kenya, Y (n ¼ 13) 112–148 (133), X (n ¼ 7) 124–161 (144), unsexed (n ¼ 13) 113–15 (131). IMMATURE: unknown.

Field Characters. Length 105–11 cm. Confined to papyrus and nearby farmland. Small dusky (Y) or grey (X) face mask not extending behind eye or onto chin, heavy streaks on breast and flanks; very similar to partly sympatric race brittoni of Southern Citril S. hypostictus but lacks yellow supercilium, has short stubby bill with curved culmen, fainter wing-bars. Voice also different. Voice. Tape-recorded (104, B, FISP, LEM, McVIC). Song of measured, high-pitched whistled and sibilant notes, lasting 3–6 s, e.g. ‘sit, syoo, tsipee, tsitsit, tsisi-pee-tsit’; dry rattle sometimes included. Another song described as a

Serinus koliensis

rapid series of short chippering notes ending with (or including) a rising slurred ‘surrreet’ (Zimmerman et al. 1996, q.v. for further transcriptions). Calls include whistled ‘wee-titi-woy’ and hard ‘wee-chachachacha’. Also said to have a variety of soft wheezing and slurring notes (Clement et al. 1993). General Habits. Inhabits papyrus Cyperus papyrus swamps; often forages in cleared areas and temporary cultivation nearby, returning to papyrus beds to roost. Has been seen among banana plants. Roosts in papyrus throughout year. Forages on ground and takes seeds in situ by clinging to heads of herbs and cereals. Solitary or, usually, in pairs, forms small flocks of up to 15 birds when foraging. Food. Seeds, principally of papyrus, also of sorghum and maize. Breeding Habits. NEST: deep open cup, rather scantily made, entirely from loosely woven, dry pieces of papyrus heads, without lining; placed in centre of papyrus head, 1–2 m (av. of 4, 15 m) above water or mud, at edge of swamp. Built by X (only?); once, 2 nests 3 m apart, 1 possibly not occupied. EGGS: 1–2 (i.e. once 1, perhaps clutch incomplete, twice 2). Dirty white, with small beige and rich brown blotches unevenly distributed all over, more concentrated at wide end, or buff, similarly with pale reddish brown blotches. SIZE: (n ¼ 5) 160–190  125–130 (176  128). WEIGHT: 05

468

FRINGILLIDAE

and 06 g, but 2 larger eggs both 12 g (P. Britton, pers. comm.). LAYING DATES: E Africa, Region B, Apr–Aug; may breed also in Nov, when sings. INCUBATION: X sits very tight.

BREEDING

SUCCESS/SURVIVAL:

some

nests

evidently

destroyed by heavy rain. Key References Britton, P.L. (1971)., Clement, P. et al. (1993), Vande weghe, J.P. (1981), Zimmerman, D.A. et al. (1996).

Plate 28

Serinus scotops (Sundevall). Forest Canary. Serin forestier.

(Opp. p. 411)

¨ fv. K. Sv. Vet.-Akad. Fo Crithagra scotops Sundevall, 1850. O ¨rh., 7, p. 98; lower Caffraria; type from Pietermaritzburg, Natal, fide W.L. Sclater, 1930, Syst. Avium Aethiopicarum, p. 816. Range and Status. Endemic resident, E South Africa and just entering Swaziland and Lesotho; locally frequent, from Soutspansberg, along Drakensberg escarpment to East London, Great Winter Berg, Grahamstown, and west along S coast to Baviaanskloofberge, Kougaberge, George and Swellendam, with several records in Caledon district and near Stellenbosch and Cape Town. In Transvaal restricted to Escarpment region from Kangwane to Entabeni, and in relict patches of forest in E Highveld; in Free State restricted to a few localities near E borders, where locally common. In Swaziland, only in far N; in Lesotho, a few records along NE borders. Description. S. s. scotops (Sundevall): coastal E Cape, Natal below 920 m. ADULT Y: narrow blackish frontal band at base of upper mandible; rest of upperparts olive-green, top of head, mantle, back and scapulars with bold blackish streaks; rump and uppertail-coverts yellower, plain or almost so, long uppertail coverts with blackish shaft streaks. Tail feathers blackish brown, outer edges olive-green. Narrow yellow superciliary stripe from above lores to side of crown. Lores and around eye to forecheeks and chin dark olive-grey, connecting with frontal band and merging with olive-green hindcheeks, ear-coverts and sides of neck. Upper throat yellow; lower throat and upper breast olivegreen, usually with a few fine dusky streaks; rest of underparts yellow, side of lower breast and flanks streaked blackish. Remiges blackish brown, primaries and secondaries edged yellowish olivegreen, tertials broadly edged and tipped yellowish olive-green; upperwing-coverts blackish brown, primary coverts and alula narrowly edged yellowish olive-green, greater coverts with olivegreen outer edges and broad yellowish olive-green tips, median coverts broadly tipped yellowish olive-green; lesser coverts broadly tipped olive-green. Underwing coverts and axillaries yellow. Bill greyish brown above, brownish white or flesh below; eyes hazel or dark brown; legs flesh-brown or grey-brown. ADULT X: differs from ad. Y in having frontal band, lores, around eye and forecheeks less dark; chin and upper throat yellow, streaked dusky olive, lower throat and upper breast with more prominent short dark streaks. SIZE (7 YY, 7 XX): wing, Y 67–74 (699), X 67–70 (680); tail, Y 51–55 (522), X 49–54 (506); bill, Y 125– 135 (130), X 125–14 (132); tarsus, Y 17–175 (171), X 165– 175 (171). WEIGHT (all races): Y (n ¼ 3) 148–156 (153), X (n ¼ 3) 146–16 (151), unsexed (n ¼ 21) 138–18 (158) (Maclean 1993). IMMATURE: juv. similar to ad. X, but slightly more olive (less green) above, paler yellow below. C. c. umbrosus Clancey: coastal S Cape, thence inland at higher levels to E Cape, Griqualand East, Natal above 1070 m and SE Transvaal. Upperparts slightly duller, darker green than in nominate race, rump usually with some streaks; slightly smaller: wing, Y (n ¼ 4) 65–69 (663). S. c. transvaalensis Roberts: E and N Transvaal. Lower throat and upper breast of Y distinctly spotted; sides and flanks more

Serinus scotops

heavily streaked in both sexes than in nominate race, upperparts slightly more heavily streaked.

Field Characters. Length 125–13 cm. A forest canary, darker and more heavily streaked than any in its range, with blackish face and chin outlined by short yellow supercilium and yellow bar across throat; pale base to bill contrasts with dark face. Unstreaked yellow-green rump offsets dark back. X and juv. similar but face and chin grey or dusky, yellow supercilium indistinct, throat streaked. Voice. Tape-recorded (72–75, 88, 99, B, X, LEM). Song high-pitched and thin, with silvery tinkling quality, sibilant notes mixing with clear sweet down-slurred whistles and a few trills, lasting 11–16 s. Calls include thin ‘tsik’, ‘tsip-tsip’ or ‘tsit-ititit’, and plaintive ‘pee-pee-pee-pyoow . . . tseet’, also rendered ‘tseetoo-tswee-ee’, with central syllable accented (Skead 1960). General Habits. Inhabits interior and edges of montane evergreen forests, dense kloofs and dense bushveld,

468

FRINGILLIDAE

and 06 g, but 2 larger eggs both 12 g (P. Britton, pers. comm.). LAYING DATES: E Africa, Region B, Apr–Aug; may breed also in Nov, when sings. INCUBATION: X sits very tight.

BREEDING

SUCCESS/SURVIVAL:

some

nests

evidently

destroyed by heavy rain. Key References Britton, P.L. (1971)., Clement, P. et al. (1993), Vande weghe, J.P. (1981), Zimmerman, D.A. et al. (1996).

Plate 28

Serinus scotops (Sundevall). Forest Canary. Serin forestier.

(Opp. p. 411)

¨ fv. K. Sv. Vet.-Akad. Fo Crithagra scotops Sundevall, 1850. O ¨rh., 7, p. 98; lower Caffraria; type from Pietermaritzburg, Natal, fide W.L. Sclater, 1930, Syst. Avium Aethiopicarum, p. 816. Range and Status. Endemic resident, E South Africa and just entering Swaziland and Lesotho; locally frequent, from Soutspansberg, along Drakensberg escarpment to East London, Great Winter Berg, Grahamstown, and west along S coast to Baviaanskloofberge, Kougaberge, George and Swellendam, with several records in Caledon district and near Stellenbosch and Cape Town. In Transvaal restricted to Escarpment region from Kangwane to Entabeni, and in relict patches of forest in E Highveld; in Free State restricted to a few localities near E borders, where locally common. In Swaziland, only in far N; in Lesotho, a few records along NE borders. Description. S. s. scotops (Sundevall): coastal E Cape, Natal below 920 m. ADULT Y: narrow blackish frontal band at base of upper mandible; rest of upperparts olive-green, top of head, mantle, back and scapulars with bold blackish streaks; rump and uppertail-coverts yellower, plain or almost so, long uppertail coverts with blackish shaft streaks. Tail feathers blackish brown, outer edges olive-green. Narrow yellow superciliary stripe from above lores to side of crown. Lores and around eye to forecheeks and chin dark olive-grey, connecting with frontal band and merging with olive-green hindcheeks, ear-coverts and sides of neck. Upper throat yellow; lower throat and upper breast olivegreen, usually with a few fine dusky streaks; rest of underparts yellow, side of lower breast and flanks streaked blackish. Remiges blackish brown, primaries and secondaries edged yellowish olivegreen, tertials broadly edged and tipped yellowish olive-green; upperwing-coverts blackish brown, primary coverts and alula narrowly edged yellowish olive-green, greater coverts with olivegreen outer edges and broad yellowish olive-green tips, median coverts broadly tipped yellowish olive-green; lesser coverts broadly tipped olive-green. Underwing coverts and axillaries yellow. Bill greyish brown above, brownish white or flesh below; eyes hazel or dark brown; legs flesh-brown or grey-brown. ADULT X: differs from ad. Y in having frontal band, lores, around eye and forecheeks less dark; chin and upper throat yellow, streaked dusky olive, lower throat and upper breast with more prominent short dark streaks. SIZE (7 YY, 7 XX): wing, Y 67–74 (699), X 67–70 (680); tail, Y 51–55 (522), X 49–54 (506); bill, Y 125– 135 (130), X 125–14 (132); tarsus, Y 17–175 (171), X 165– 175 (171). WEIGHT (all races): Y (n ¼ 3) 148–156 (153), X (n ¼ 3) 146–16 (151), unsexed (n ¼ 21) 138–18 (158) (Maclean 1993). IMMATURE: juv. similar to ad. X, but slightly more olive (less green) above, paler yellow below. C. c. umbrosus Clancey: coastal S Cape, thence inland at higher levels to E Cape, Griqualand East, Natal above 1070 m and SE Transvaal. Upperparts slightly duller, darker green than in nominate race, rump usually with some streaks; slightly smaller: wing, Y (n ¼ 4) 65–69 (663). S. c. transvaalensis Roberts: E and N Transvaal. Lower throat and upper breast of Y distinctly spotted; sides and flanks more

Serinus scotops

heavily streaked in both sexes than in nominate race, upperparts slightly more heavily streaked.

Field Characters. Length 125–13 cm. A forest canary, darker and more heavily streaked than any in its range, with blackish face and chin outlined by short yellow supercilium and yellow bar across throat; pale base to bill contrasts with dark face. Unstreaked yellow-green rump offsets dark back. X and juv. similar but face and chin grey or dusky, yellow supercilium indistinct, throat streaked. Voice. Tape-recorded (72–75, 88, 99, B, X, LEM). Song high-pitched and thin, with silvery tinkling quality, sibilant notes mixing with clear sweet down-slurred whistles and a few trills, lasting 11–16 s. Calls include thin ‘tsik’, ‘tsip-tsip’ or ‘tsit-ititit’, and plaintive ‘pee-pee-pee-pyoow . . . tseet’, also rendered ‘tseetoo-tswee-ee’, with central syllable accented (Skead 1960). General Habits. Inhabits interior and edges of montane evergreen forests, dense kloofs and dense bushveld,

Ochrospiza keeping mainly to canopy and mid levels of forest; in E Cape Prov. dry, Euphorbia-dominated communities on south-facing slopes of valleys; readily enters gardens, orchards, scrub, tree plantations and maize fields (Harrison et al. 1997). Singly or in pairs, sometimes in flocks of up to c. 12 birds. Occurs with Cape Canaries S. canicollis in plantations of exotic trees. Active; forages in mid strata and tops of forest trees, members of pair or flock keeping in contact with quiet calls; but generally rather silent and easily overlooked. Keeps mainly well within leafy cover, high in forest canopy. Sometimes feeds on ground, at least in gardens, where comes readily to bird tables. Forages for seeds and small fruits in situ in weeds or shrubs, or for fallen seeds on ground. Once seen ‘to tear off the newly grown leaves of a sneezewood tree, and chew at the ends of the petioles with their beaks, after letting the rest of the leaf fall to the ground’ (Skead 1960). Singing Y perches prominently on tree top or sometimes remains within dense vegetation. Sedentary around Grahamstown (as shown by ringing study); resident in East London area but moves locally in winter; tends to be attracted to plants with unpredictable flowering and fruiting seasons (C.J. Vernon in Ginn et al. 1989).

Food. Seeds of shrubs and weeds, small fruits including ripe figs Ficus sp. and Anthospermum sp. (Rubiaceae); parts of sneezewood Ptaeroxylon obliquum (see above). In captivity eats termites and seeds of Alternanthera pungens, Senecio tamoides and S. polyanthemoides, and very fond of unripe seeds of Bidens pilosa (Brickell and Green 1986).

Breeding Habits. Solitary nester; thought to be monogamous and territorial. NEST: a small, neat, open cup made of stringy moss and very fine herb stems, warmly lined with fibrous material and lichen; sited 1–5 m above ground, in fork amongst branches at side of tree or bush (thorn tree or Schotia bush), amongst foliage and well hidden from view. One nest, ext. diam. 102, int. diam. 58; noticeably larger than nest of e.g. Yellow-fronted Canary S. mozambicus. 2 nests were only 2 m apart, in conifer in garden near edge of forest, East London, one with eggs, the other with chicks. Built by X; both sexes bring material. Double-brooded in some years. EGGS: 2–4, usually 3 or 4. White or pale bluish white, speckled sparingly or sometimes quite heavily with greys, reds and browns, mainly at broad end, not in a ring. SIZE: (n ¼ 6) 169–183  122–133 (173  126). LAYING DATES: South Africa, Transvaal, Jan, (and fledglings June); E Cape, Nov, Mar (and nest-building, Oct, juvs, Jan); in East London area breeds whenever conditions are favourable (C.J. Vernon in Ginn et al. 1989); KwaZulu-Natal, (breeds Oct–Jan: Maclean 1993). INCUBATION: by X only. Period in captivity, 14 days (n ¼ 2). DEVELOPMENT AND CARE OF YOUNG: in captivity fed by X only; Y brings food to give to X on nest. Nestling period (n ¼ 4) 15–19 (175) days. Both parents feed the young after they have left nest. BREEDING SUCCESS/SURVIVAL: no information. Key References Clancey, P.A. (1964), Clement, P. et. al. (1993), Ginn, P.J. et al. (1989), Harrison, J.A. et al. (1997), Skead, C.J. (1960), Maclean, G.L. (1993).

Subgenus Ochrospiza Roberts

10 species of mostly rather small, stubby-billed serins, 9 African, 1 Arabian, of moist or arid or subdesert scrubland and savannas; plumage colours and patterns rather diverse. Diagnosed by van den Elzen (1985) (who studied all species, in captivity, except S. (O.) flavigula) on the basis of 4 behavioural characters: (1) in highest intensity courtship, YY finish with body held upright and wings half-opened and raised in a heraldic posture (most pronounced in yellow-rumped grey atrogularis, reichenowi and xanthopygius; posture less upright in yellow-rumped and yellow-faced mozambicus and dorsostriatus; the white-rumped grey leucopygius fluffs rump feathers instead); (2) Y helps X build nest; (3) nest rim becomes sullied with thick layer of nestling faeces; (4) threat display like those of subgenus Crithagra, i.e. with head held high and tail flicking; also a morphological character: gapes of fledglings red, with blue bill corners. Ancestral leucopygius is thought to have been yellow-rumped (van den Elzen 1985). Ethiopian S. (O.) flavigula and xantholaemus form a superspecies; Arabian rothschildi may belong to that superspecies or to the following one. S. (O.) atrogularis (southern Africa), reichenowi (E Africa) and xanthopygius (N Ethiopia) compose a superspecies; their differences in plumage and facial appearance are slight and they were regarded as a single species by Hall and Moreau (1970), but were treated as specifically separate by van den Elzen (1985, 1999) and Sibley and Monroe (1990). Dowsett and DowsettLemaire (1993a) treated reichenowi as conspecific with atrogularis, contra Irwin (1964), but kept xanthopygius specifically separate on account of its proportions, strikingly different from those of atrogularis (Erard 1974). Other authorities have been equally divided. S. (O.) atrogularis and S. (O.) mozambicus can hybridize in captivity (van den Elzen 1981). Endemic. 9 species, 5 in 2 superspecies, 4 independent.

469

Ochrospiza keeping mainly to canopy and mid levels of forest; in E Cape Prov. dry, Euphorbia-dominated communities on south-facing slopes of valleys; readily enters gardens, orchards, scrub, tree plantations and maize fields (Harrison et al. 1997). Singly or in pairs, sometimes in flocks of up to c. 12 birds. Occurs with Cape Canaries S. canicollis in plantations of exotic trees. Active; forages in mid strata and tops of forest trees, members of pair or flock keeping in contact with quiet calls; but generally rather silent and easily overlooked. Keeps mainly well within leafy cover, high in forest canopy. Sometimes feeds on ground, at least in gardens, where comes readily to bird tables. Forages for seeds and small fruits in situ in weeds or shrubs, or for fallen seeds on ground. Once seen ‘to tear off the newly grown leaves of a sneezewood tree, and chew at the ends of the petioles with their beaks, after letting the rest of the leaf fall to the ground’ (Skead 1960). Singing Y perches prominently on tree top or sometimes remains within dense vegetation. Sedentary around Grahamstown (as shown by ringing study); resident in East London area but moves locally in winter; tends to be attracted to plants with unpredictable flowering and fruiting seasons (C.J. Vernon in Ginn et al. 1989).

Food. Seeds of shrubs and weeds, small fruits including ripe figs Ficus sp. and Anthospermum sp. (Rubiaceae); parts of sneezewood Ptaeroxylon obliquum (see above). In captivity eats termites and seeds of Alternanthera pungens, Senecio tamoides and S. polyanthemoides, and very fond of unripe seeds of Bidens pilosa (Brickell and Green 1986).

Breeding Habits. Solitary nester; thought to be monogamous and territorial. NEST: a small, neat, open cup made of stringy moss and very fine herb stems, warmly lined with fibrous material and lichen; sited 1–5 m above ground, in fork amongst branches at side of tree or bush (thorn tree or Schotia bush), amongst foliage and well hidden from view. One nest, ext. diam. 102, int. diam. 58; noticeably larger than nest of e.g. Yellow-fronted Canary S. mozambicus. 2 nests were only 2 m apart, in conifer in garden near edge of forest, East London, one with eggs, the other with chicks. Built by X; both sexes bring material. Double-brooded in some years. EGGS: 2–4, usually 3 or 4. White or pale bluish white, speckled sparingly or sometimes quite heavily with greys, reds and browns, mainly at broad end, not in a ring. SIZE: (n ¼ 6) 169–183  122–133 (173  126). LAYING DATES: South Africa, Transvaal, Jan, (and fledglings June); E Cape, Nov, Mar (and nest-building, Oct, juvs, Jan); in East London area breeds whenever conditions are favourable (C.J. Vernon in Ginn et al. 1989); KwaZulu-Natal, (breeds Oct–Jan: Maclean 1993). INCUBATION: by X only. Period in captivity, 14 days (n ¼ 2). DEVELOPMENT AND CARE OF YOUNG: in captivity fed by X only; Y brings food to give to X on nest. Nestling period (n ¼ 4) 15–19 (175) days. Both parents feed the young after they have left nest. BREEDING SUCCESS/SURVIVAL: no information. Key References Clancey, P.A. (1964), Clement, P. et. al. (1993), Ginn, P.J. et al. (1989), Harrison, J.A. et al. (1997), Skead, C.J. (1960), Maclean, G.L. (1993).

Subgenus Ochrospiza Roberts

10 species of mostly rather small, stubby-billed serins, 9 African, 1 Arabian, of moist or arid or subdesert scrubland and savannas; plumage colours and patterns rather diverse. Diagnosed by van den Elzen (1985) (who studied all species, in captivity, except S. (O.) flavigula) on the basis of 4 behavioural characters: (1) in highest intensity courtship, YY finish with body held upright and wings half-opened and raised in a heraldic posture (most pronounced in yellow-rumped grey atrogularis, reichenowi and xanthopygius; posture less upright in yellow-rumped and yellow-faced mozambicus and dorsostriatus; the white-rumped grey leucopygius fluffs rump feathers instead); (2) Y helps X build nest; (3) nest rim becomes sullied with thick layer of nestling faeces; (4) threat display like those of subgenus Crithagra, i.e. with head held high and tail flicking; also a morphological character: gapes of fledglings red, with blue bill corners. Ancestral leucopygius is thought to have been yellow-rumped (van den Elzen 1985). Ethiopian S. (O.) flavigula and xantholaemus form a superspecies; Arabian rothschildi may belong to that superspecies or to the following one. S. (O.) atrogularis (southern Africa), reichenowi (E Africa) and xanthopygius (N Ethiopia) compose a superspecies; their differences in plumage and facial appearance are slight and they were regarded as a single species by Hall and Moreau (1970), but were treated as specifically separate by van den Elzen (1985, 1999) and Sibley and Monroe (1990). Dowsett and DowsettLemaire (1993a) treated reichenowi as conspecific with atrogularis, contra Irwin (1964), but kept xanthopygius specifically separate on account of its proportions, strikingly different from those of atrogularis (Erard 1974). Other authorities have been equally divided. S. (O.) atrogularis and S. (O.) mozambicus can hybridize in captivity (van den Elzen 1981). Endemic. 9 species, 5 in 2 superspecies, 4 independent.

469

470

Plate 28 (Opp. p. 411)

FRINGILLIDAE

Serinus leucopygius (Sundevall). White-rumped Seedeater; White-rumped Serin; Grey Canary. Serin a ` croupion blanc. ¨ fv. K. Sv. Vet.-Akad. Fo Crithagra leucopygia Sundevall, 1850. O ¨rh., 7, p. 127; Sennar. Range and Status. Endemic resident, sahel and soudanian zone of W Africa, east to Eritrea, south to NW Uganda. Mauritania, frequent north to 17
N; common in lower Senegal R. valley and Nouakchott area, at least in Jan–Aug (Gee 1984). Senegal, frequent and widespread north of Gambia; uncommon in Djoudj Nat. Park; Casamance. Gambia, uncommon on coast and in lower river valley, locally frequent in central and upper river valley. Mali, as mapped (Lamarche 1993); resident in lower delta area (e.g. Niafunke´) but subject to seasonal movements both north and south of there. Burkina Faso, frequent resident in N Yatenga area, north of c. 13
450 N; uncommon south to Ouaga and Koubri (30 km southeast of Ouagadougou), and occurs south to 12
200 N; not south of 12
N. Ghana, uncommon resident in extreme NE, regular at Vea Dam and Bolgatanga; flocks of c. 30 in Mar–Apr; singing in July. Togo, rare in extreme NW: 1–3 birds at Barkoissi, Kpani and Mango, Mar–Apr, and 1 singing Cinkanse´, Aug. Niger, uncommon resident in ‘W’ Nat. Park area, rare in Niger valley further north (Ayorou, Niamey), numerous in Aı¨r Massif in N, uncommon in remainder of country (Zinder, Takoukout, Filinge´, Damergu, Asben, Tahoua, west of Tahoua, near Maradi, Nguigmi). Nigeria, locally frequent south to Sokoto, Zaria and Yankari Nat. Park; common around Katsina, Kano and in Bornu Province. Cameroon, uncommon, south to Rei Buba and Ngaounde´re´. Chad, common in sahel and soudanian zones and Ennedi Massif; abundant at Abe´che´, where 500 caught in one mistnet during one year; frequent in Ouadi Rime-Ouadi Achim Faunal Res. in areas of permanent settlement where cereals are grown, e.g. S Goz Arian, Subbou and Arada. Central African Republic, occasional in Manovo-GoundaSaint Floris Nat. Park and Vakaga area (June). Sudan, frequent to common but local; common and widespread in W Kordofan in S, in Nile Valley at Gogrial, Rumbek and Torit; extending its range with new irrigation schemes (Nikolaus 1987, 1989). Eritrea, very common below 1000 m in W. Uganda, uncommon, along Albert Nile and in Murchison Falls Nat. Park at 600–1000 m. Zaı¨re, known only from N end of L. Albert. Ethiopia, common in W. Unidentified white-rumped serins have been seen in and near Ethiopia’s Awash/Rift Valley, at Jerbasri near Agere Maryam (5
130 , 39
200 E), L. Langano and – another species?, with c. 10 sightings in 20 years – at foot of Mt Fantalle (9
000 N, 39
540 E), and in acacias between Awash and Gewane (10
120 N, 40
400 E) (Ash 1979, Atkins and Harvey 1996). Large numbers of S. leucopygius (even more of Yellowfronted Canary S. mozambicus) used to be caught in Senegambia and exported for the songbird trade, and perhaps still are.

scapulars and back; rump and short uppertail-coverts white; long uppertail-coverts dull brown, fringed greyish. Tail feathers dark brown, finely edged pale buff. Lores, cheeks and ear-coverts greybrown, darker on lores and behind eye, forming indistinct eyestripe; sides of neck paler grey-brown with faint dusky streaks. Chin to upper breast pale grey-brown with fine short dark brown shaft marks; rest of underparts white, tinged grey-brown, lower breast and flanks with a few dusky streaks. Remiges dark brown, flight feathers finely edged, and secondary tips finely fringed pale grey-buff, tertials edged and tipped pale grey-buff; alula and larger upperwing-coverts dark brown, greater and median coverts narrowly edged and tipped pale grey-buff to form 2 distinct narrow wing-bars. Underwing-coverts and axillaries brownish white. Bill horn or pinkish brown with dusky tip and cutting edge and pale or whitish base; eyes light brown or hazel; legs pale fleshy pink to pale horn. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 63–68 (661), X 63–70 (654); tail, Y 39–43 (404), X 38–46 (408); bill, Y 11–115 (112), X 105–115 (108); tarsus, Y 13– 145 (137), X 13–14 (137). WEIGHT: (Ethiopia, unsexed, n ¼ 14) 94–127 (108). IMMATURE: juv. has upperparts more buffish than ad., more streaked with dark brown; chin to breast and flanks washed buff and spotted dark brown. S. l. riggenbachi Neumann (including ‘pallens’): Senegal to W Sudan. Slightly greyer than nominate race; chin to breast whitish with distinct dark brown spots across upper breast; top of head more strongly spotted. WEIGHT: (Chad, unsexed, n ¼ 100) 8–16 (105).

Description. S. l. leucopygius: central and S Sudan to W Eritrea, W Ethiopia and NW Uganda. ADULT Y: forehead to back and scapulars pale grey-brown, dark brown feather centres forming spotting from forehead to hindneck, broader mottling on mantle,

Field Characters. Length 10–12 cm. The only seedeater with a white rump; otherwise pale grey-brown, streaked and mottled on upperparts and breast, with pale wing-bar.

Serinus leucopygius

?

X's in Ethiopia: see text.

Serinus atrogularis On the ground told from other seedeaters and X indigobirds Vidua by small stubby pale bill and uniform appearance, without supercilium or other facial markings. Voice. Tape-recorded (104, B, CHA, MOR, PAY). Rapid song of clear sweet notes, with some liquid trills. Calls, rising ‘twee’ or ‘chuwee’ and nasal ‘jwer’. General Habits. Inhabits arid country with sandy soil, scattered bushes and trees, light woodland with much open space, Acacia steppe, surrounds of towns and around small rural settlements in W African sahel zone, cattle watering points, agricultural land, millet fields, and large gardens. In pairs and small flocks; forages on ground, often with waxbills, indigobirds and Yellow-fronted Canary. To take seeds in fruiting head of composite Tridax procumbens, bird standing on ground reaches up as high as it can, seizes stem in bill and bends it to ground, treads on it proximal to point where it is held in bill, and walks rapidly crabwise along prone stem to the fruiting head; bird spends a few s plucking and eating seeds before moving off (when stem springs upright again); up to 3 birds feed on adjacent stems in this manner (Fry 1975). Tends to become stained yellow with mimosa pollen (Chad, Oct). Food. Seeds, some buds. Partial to seeds of Tridax procumbens (Nigeria, where Tridax introduced in 1930s). In captivity eats seeds of Stellaria media, Taraxacum officinale and Senecio consanguineus and buds of Galinsoga parviflora. Breeding Habits. Solitary nester. 8 pairs of S. leucopygius and S. mozambicus nested in 1 ha patch of soft-foliaged trees

471

in Kulme, Sudan (southwest of Jebel Marra, Darfur). One nest (with 3 eggs) was built only a few cm from one collected, and incubating X shot, only 14 days earlier (Kano, Nigeria, Shuel 1938). NEST: a compact, rounded cup, strongly made of plant stems, fibres, leaf spines, vegetable down and small fragments of leaf, held together with silk, lined with a well-defined layer of rootlets. Ext. diam. 60, int. diam. 43– 45, ext. depth 52, int. depth 27. Placed over and between 2 slender twigs, 15–5 m above ground, in extremities of tree branches, often acacias; once near end of a 50 mm thick branch 5 m up in a baobab. Nest built in 60 in Goda Massif (Welch and Welch 1998). Ethiopia, frequent in SE Highlands, E Rift Valley and S region, marginally into NE region; occurs in Bahar Dar-L. Tana, Jemma and Jara Valleys, and Tiro Boter-Becho Forest IBAs (Tilahun et al. 1996). Sudan, uncommon, restricted to extreme SE (mainly southeast of Kapoeta; once Tambura). Uganda, Moroto and Kidepo Valley Nat. Park. Somalia, frequent in extreme S, fairly common but very local in lower and mid Jubba R. and Shabeelle R. valleys. Kenya and Tanzania, fairly common south of 1
N from coast and lowlands up to 2000 m, mainly east of 35
E, from Moyale and Wajir to Dar es Salaam,

hinterland of Kilwa Masoko, Iringa highlands, and Chiwanda area and lower Ruvuma Valley in extreme S. In very arid N Kenya, known from Marsabit, Moyale and Wajir; seasonal in Nguuni, Mombasa, where flocks, sometimes large, occur mainly in Nov–Apr, outside rainy periods (Ryall 1991). In Tanzania as mapped; several records of yellow-rumped serins along SL. Victoria shores between Emin Pasha Gulf and Mwanza Gulf (N. Baker, pers. comm.), not mapped, probably refer to S. atrogularis. Description. ADULT Y: frontal band buffish white, continuing back to above ear-coverts as broad superciliary stripe; otherwise, forehead to back and scapulars buffish brown, streaked dark brown; rump bright lemon yellow; uppertail-coverts dull brown,

476

FRINGILLIDAE

Serinus reichenowi

IMMATURE: juv. more buffish than ad.; more extensively spotted on upper breast and more heavily streaked on lower breast and flanks.

Field Characters. Length 10–11 cm. The common yellowrumped brown seedeater of eastern Africa, smaller and shorter-tailed than Streaky Seedeater S. striolatus and less heavily streaked below. Differs from Black-throated Canary S. atrogularis, which it meets in W Kenya, in paler upperparts, pronounced white supercilium and unspotted white throat; underparts whiter, breast spotted brown. Voice. Tape-recorded (109). Calls, dry ‘chit’ or ‘chit-chee’, sometimes lengthened into whistled phrase ‘chit-cheechiddly-oo’; also a clear, rising ‘tweee’. Song described as a continuous rapid finch-like warbling with rich trills and whistles, sweet and canary-like (Zimmerman et al. 1996).

tipped greyish white. Tail feathers dark brown, finely fringed buff, inner webs of T6 and T5 tipped white. Broad brown band through lores and across upper cheek to ear-coverts, within which a narrow buffish white bar below eye; lower cheek buffish white, bordered by prominent dark brown malar stripe. Underparts white, washed with buff, upper breast with gorget of short brown streaks or spots, sides of breast and edge of flanks tinged brownish and streaked light brown. Remiges dark brown, primaries finely edged yellowish, secondaries narrowly edged pale buffish brown, tertials broadly edged buffish brown and tipped greyish white; alula and upperwing-coverts dark brown, greater coverts edged buff, these and median coverts tipped whitish to form distinct narrow wing-bars, lesser coverts tipped buffish brown. Underwing-coverts and axillaries buffish white. Bill brownish horn, base of lower mandible pale flesh or whitish; eyes brown; legs light brown or brownish flesh. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 64–69 (670), X 65–70 (673); tail, Y 36–44 (412), X 40– 44 (420); bill, Y 105–125 (112), X 10–115 (107); tarsus, Y 13– 14 (133), X 13–14 (135). WEIGHT: (Ethiopia, unsexed, n ¼ 42) 107–127 (104); (W Kenya, n ¼ 41, unsexed) 94–132 (109).

Plate 28 (Opp. p. 411)

General Habits. Inhabits arid areas with scrub, bush, open woodland and cultivation. Occurs in pairs and flocks of 10– 15 birds; occasionally up to 50 congregate at water to drink. Forages on ground and amongst roadside weeds, where unobtrusive until flock takes wing; feeds at seeding heads of herbs. Singing and displaying birds near Konso (S Ethiopia: see next species) did not interact with Yellowrumped Serins S. xanthopygins which were also in full display there at the time (N. Gardner, pers. comm.). Most populations are resident, but species prone to irregular wandering. Food. Not known; presumably seeds. Breeding Habits. NEST: a small cup made of rootlets and grass stems, bound with cobwebs, lined with plant down, placed on branch in bush or tree. EGGS: 3. Bluish white, often with dull mauve undermarkings, spotted with brown and black. SIZE: c. 16  12. LAYING DATES: Ethiopia, May; Somalia, Dec (and fledglings, Feb); E Africa, Region C, Jan–Mar, May, Region D, Dec–Jan, Mar–July, mainly in long rains, i.e. Mar–June (18 out of 22 clutches). Nothing further known.

Serinus xanthopygius Ru ¨ ppell. Yellow-rumped Seedeater; White-throated Serin. Serin a` croupion jaune. Serinus xanthopygius Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 96; Schoada Valley, Simen, Abyssinia. Forms a superspecies with S. atrogularis and S. reichenowi. Range and Status. Endemic resident, Eritrea and N Ethiopia. Eritrea, fairly common on central plateau at 1000–2500 m. Ethiopia, widespread at 900–2100 m in W Highlands, from Bahar Dar (where common at Tissisat Falls) east to escarpment at 40
E, marginally in W Rift Valley, probably widespread also in far N, adjoining Eritrean range; c. 5 birds in full display, 7 km southwest

of Konso, 5
170 N, 37
310 E, Nov 1996 (N. Gardner, pers. comm.). Description. ADULT Y: forehead to back and scapulars drab brown with narrow darker brown streaks, almost uniform in worn plumage; rump lemon yellow; uppertail-coverts drab brown, tinged yellow. Tail feathers dull dark brown. No distinct

476

FRINGILLIDAE

Serinus reichenowi

IMMATURE: juv. more buffish than ad.; more extensively spotted on upper breast and more heavily streaked on lower breast and flanks.

Field Characters. Length 10–11 cm. The common yellowrumped brown seedeater of eastern Africa, smaller and shorter-tailed than Streaky Seedeater S. striolatus and less heavily streaked below. Differs from Black-throated Canary S. atrogularis, which it meets in W Kenya, in paler upperparts, pronounced white supercilium and unspotted white throat; underparts whiter, breast spotted brown. Voice. Tape-recorded (109). Calls, dry ‘chit’ or ‘chit-chee’, sometimes lengthened into whistled phrase ‘chit-cheechiddly-oo’; also a clear, rising ‘tweee’. Song described as a continuous rapid finch-like warbling with rich trills and whistles, sweet and canary-like (Zimmerman et al. 1996).

tipped greyish white. Tail feathers dark brown, finely fringed buff, inner webs of T6 and T5 tipped white. Broad brown band through lores and across upper cheek to ear-coverts, within which a narrow buffish white bar below eye; lower cheek buffish white, bordered by prominent dark brown malar stripe. Underparts white, washed with buff, upper breast with gorget of short brown streaks or spots, sides of breast and edge of flanks tinged brownish and streaked light brown. Remiges dark brown, primaries finely edged yellowish, secondaries narrowly edged pale buffish brown, tertials broadly edged buffish brown and tipped greyish white; alula and upperwing-coverts dark brown, greater coverts edged buff, these and median coverts tipped whitish to form distinct narrow wing-bars, lesser coverts tipped buffish brown. Underwing-coverts and axillaries buffish white. Bill brownish horn, base of lower mandible pale flesh or whitish; eyes brown; legs light brown or brownish flesh. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 64–69 (670), X 65–70 (673); tail, Y 36–44 (412), X 40– 44 (420); bill, Y 105–125 (112), X 10–115 (107); tarsus, Y 13– 14 (133), X 13–14 (135). WEIGHT: (Ethiopia, unsexed, n ¼ 42) 107–127 (104); (W Kenya, n ¼ 41, unsexed) 94–132 (109).

Plate 28 (Opp. p. 411)

General Habits. Inhabits arid areas with scrub, bush, open woodland and cultivation. Occurs in pairs and flocks of 10– 15 birds; occasionally up to 50 congregate at water to drink. Forages on ground and amongst roadside weeds, where unobtrusive until flock takes wing; feeds at seeding heads of herbs. Singing and displaying birds near Konso (S Ethiopia: see next species) did not interact with Yellowrumped Serins S. xanthopygins which were also in full display there at the time (N. Gardner, pers. comm.). Most populations are resident, but species prone to irregular wandering. Food. Not known; presumably seeds. Breeding Habits. NEST: a small cup made of rootlets and grass stems, bound with cobwebs, lined with plant down, placed on branch in bush or tree. EGGS: 3. Bluish white, often with dull mauve undermarkings, spotted with brown and black. SIZE: c. 16  12. LAYING DATES: Ethiopia, May; Somalia, Dec (and fledglings, Feb); E Africa, Region C, Jan–Mar, May, Region D, Dec–Jan, Mar–July, mainly in long rains, i.e. Mar–June (18 out of 22 clutches). Nothing further known.

Serinus xanthopygius Ru ¨ ppell. Yellow-rumped Seedeater; White-throated Serin. Serin a` croupion jaune. Serinus xanthopygius Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 96; Schoada Valley, Simen, Abyssinia. Forms a superspecies with S. atrogularis and S. reichenowi. Range and Status. Endemic resident, Eritrea and N Ethiopia. Eritrea, fairly common on central plateau at 1000–2500 m. Ethiopia, widespread at 900–2100 m in W Highlands, from Bahar Dar (where common at Tissisat Falls) east to escarpment at 40
E, marginally in W Rift Valley, probably widespread also in far N, adjoining Eritrean range; c. 5 birds in full display, 7 km southwest

of Konso, 5
170 N, 37
310 E, Nov 1996 (N. Gardner, pers. comm.). Description. ADULT Y: forehead to back and scapulars drab brown with narrow darker brown streaks, almost uniform in worn plumage; rump lemon yellow; uppertail-coverts drab brown, tinged yellow. Tail feathers dull dark brown. No distinct

Serinus citrinipectus

Serinus xanthopygius

? ?

477

edged pale brown; greater and median coverts tipped pale brown, forming inconspicuous wing-bars. Underwing-coverts and axillaries brownish white. Bill dark brown; eyes brown; legs dark brown. Sexes alike. SIZE (4 YY, 1 X): wing, Y 67–70 (683), X 70; tail, Y 48–51 (493), X 49; bill, Y 12–135 (128), X 12; tarsus, Y 14–15 (145), X 145. Also, unsexed (n ¼ 12) wing, 66–71 (688), tail, 48–52 (503) (Erard 1974). IMMATURE: juv. similar to ad., but with stronger grey-brown wash on breast and flanks; may show small yellow throat patch (Erard 1974). TAXONOMIC NOTE: see subgeneric diagnosis.

Field Characters. Length 11–12 cm. Northernmost of the small yellow-rumped brown seedeaters; may meet Reichenow’s Seedeater S. reichenowi in central and S Ethiopia. Greyer above than Reichenow’s, pale supercilium indistinct; throat white, rest of underparts pale grey, almost plain (streaks reduced to narrow band across upper breast). Voice. Tape-recorded (B, GARD). Song of short phrases, high-pitched and rather tuneless for a canary, ‘see-seechewit’, ‘see-tsi-chew-sit-tsuwee’; very different from Reichenow’s Seedeater. Calls, sibilant ‘tseet’ and ‘tseesu’ and a thin, high-pitched chatter.

superciliary stripe. Lores, cheeks and ear-coverts greyish brown, with rather darker malar area; sides of neck greyish brown. Chin and throat brownish white; rest of underparts pale grey-brown, upper breast with band of poorly defined dusky mottling, flanks with faint dusky streaks. Upperwing feathers dull dark brown; narrow edges of primaries and secondaries tinged yellow; tertials

Habits. Very little on record. Inhabits upland scrub (Eritrea), where flocks often use telegraph wires; absent from dense woodland. Singing and displaying birds near Konso (S Ethiopia: see above) did not interact with Reichenow’s Serins which were also in full display there at the time (N. Gardner, pers. comm.). Details of nesting not described.

Serinus citrinipectus Clancey and Lawson. Lemon-breasted Canary. Serin a ` poitrine citron.

Plate 29

Serinus citrinipectus Clancey and Lawson, 1960. Durban. Mus. Novit., 6, p. 62; near Panda, Inhambane, Sul do Save, Portuguese East Africa.

(Opp. p. 458)

Range and Status. Endemic resident, S Mozambique and ˆ i, SE Zimbabwe, E Transvaal and NE adjacently in S Malaw KwaZulu-Natal. Rare to uncommon or locally common. Said to be a single record in Zambia (A. Berruti in Harrison ˆ i, uncommon: Mpatamanga Gorge, et al. 1997). Malaw Lengwe Nat. Park, Chiromo, Nchalo. Zimbabwe, record from Mozambique border at about 32
450 E; in SE, sparse in lowveld below 600 m from Chiredzi, along Lundi R. to confluences of Mkwasene and Sabi rivers and Sangwe Tribal Trust territory; also from c. 20 km north of Beitbridge to lower Bubye and Nuanetsi rivers; 2 flocks of c. 500 birds in total at confluence of Maose and Mwenezi rivers and records from Chikwarakwara (Hustler 1991). Mozambique, resident in arid woodland, broad-leaved littoral savanna and cultivated country, patchily distributed as mapped, generally uncommon, but locally numerous; population thought to exceed 50,000 birds. Transvaal, no substantiated record (Tarboton et al. 1987), but a few seen in recent years (Harrison et al. 1997). KwaZulu-Natal, resident, with quite large flocks wandering in regions of Hluhluwe, Mtubatuba, Pelindaba, Mkuzi and Umfolozi Game Reserves.

Seems to be strongly associated with Ilala palms Hyphaene natalensis (used for nesting) and its survival prospects could be tied with those of the palm, which is threatened in Zimbabwe by cane furniture manufacture (Hustler 1991). Commercial tapping for palm wine making could affect birds’ breeding success and status in the future (Chittenden 1998). From Mozambique up to 2000 are exported annually as cage birds; ‘the effect of the trade on the wild population is not known and must be of some concern’ (Parker 1999). Description. ADULT Y: forehead to hindneck pale greyish brown with sharp narrow blackish brown streaks; mantle, scapulars and back light buffy brown with broad dark brown streaks; rump lemon yellow; uppertail-coverts dark brown, tipped pale yellow. Tail feathers dark brown, outer webs with narrow buff edges, tinged olive-green basally; inner webs edged and tipped white, those of T4–T5 with broad white tips, and of T6 with distal half to two-thirds white. Supraloral spot white, tinged yellow; no superciliary stripe. Lores, cheeks and ear-coverts greyish brown, with pale yellow or buffy yellow patch on forecheek bordered below by broad dusky brown moustachial/malar stripe; sides of neck light greyish brown with diffuse darker streaking. Chin, throat and centre of upper breast dull lemon yellow, upper breast

Serinus citrinipectus

Serinus xanthopygius

? ?

477

edged pale brown; greater and median coverts tipped pale brown, forming inconspicuous wing-bars. Underwing-coverts and axillaries brownish white. Bill dark brown; eyes brown; legs dark brown. Sexes alike. SIZE (4 YY, 1 X): wing, Y 67–70 (683), X 70; tail, Y 48–51 (493), X 49; bill, Y 12–135 (128), X 12; tarsus, Y 14–15 (145), X 145. Also, unsexed (n ¼ 12) wing, 66–71 (688), tail, 48–52 (503) (Erard 1974). IMMATURE: juv. similar to ad., but with stronger grey-brown wash on breast and flanks; may show small yellow throat patch (Erard 1974). TAXONOMIC NOTE: see subgeneric diagnosis.

Field Characters. Length 11–12 cm. Northernmost of the small yellow-rumped brown seedeaters; may meet Reichenow’s Seedeater S. reichenowi in central and S Ethiopia. Greyer above than Reichenow’s, pale supercilium indistinct; throat white, rest of underparts pale grey, almost plain (streaks reduced to narrow band across upper breast). Voice. Tape-recorded (B, GARD). Song of short phrases, high-pitched and rather tuneless for a canary, ‘see-seechewit’, ‘see-tsi-chew-sit-tsuwee’; very different from Reichenow’s Seedeater. Calls, sibilant ‘tseet’ and ‘tseesu’ and a thin, high-pitched chatter.

superciliary stripe. Lores, cheeks and ear-coverts greyish brown, with rather darker malar area; sides of neck greyish brown. Chin and throat brownish white; rest of underparts pale grey-brown, upper breast with band of poorly defined dusky mottling, flanks with faint dusky streaks. Upperwing feathers dull dark brown; narrow edges of primaries and secondaries tinged yellow; tertials

Habits. Very little on record. Inhabits upland scrub (Eritrea), where flocks often use telegraph wires; absent from dense woodland. Singing and displaying birds near Konso (S Ethiopia: see above) did not interact with Reichenow’s Serins which were also in full display there at the time (N. Gardner, pers. comm.). Details of nesting not described.

Serinus citrinipectus Clancey and Lawson. Lemon-breasted Canary. Serin a ` poitrine citron.

Plate 29

Serinus citrinipectus Clancey and Lawson, 1960. Durban. Mus. Novit., 6, p. 62; near Panda, Inhambane, Sul do Save, Portuguese East Africa.

(Opp. p. 458)

Range and Status. Endemic resident, S Mozambique and ˆ i, SE Zimbabwe, E Transvaal and NE adjacently in S Malaw KwaZulu-Natal. Rare to uncommon or locally common. Said to be a single record in Zambia (A. Berruti in Harrison ˆ i, uncommon: Mpatamanga Gorge, et al. 1997). Malaw Lengwe Nat. Park, Chiromo, Nchalo. Zimbabwe, record from Mozambique border at about 32
450 E; in SE, sparse in lowveld below 600 m from Chiredzi, along Lundi R. to confluences of Mkwasene and Sabi rivers and Sangwe Tribal Trust territory; also from c. 20 km north of Beitbridge to lower Bubye and Nuanetsi rivers; 2 flocks of c. 500 birds in total at confluence of Maose and Mwenezi rivers and records from Chikwarakwara (Hustler 1991). Mozambique, resident in arid woodland, broad-leaved littoral savanna and cultivated country, patchily distributed as mapped, generally uncommon, but locally numerous; population thought to exceed 50,000 birds. Transvaal, no substantiated record (Tarboton et al. 1987), but a few seen in recent years (Harrison et al. 1997). KwaZulu-Natal, resident, with quite large flocks wandering in regions of Hluhluwe, Mtubatuba, Pelindaba, Mkuzi and Umfolozi Game Reserves.

Seems to be strongly associated with Ilala palms Hyphaene natalensis (used for nesting) and its survival prospects could be tied with those of the palm, which is threatened in Zimbabwe by cane furniture manufacture (Hustler 1991). Commercial tapping for palm wine making could affect birds’ breeding success and status in the future (Chittenden 1998). From Mozambique up to 2000 are exported annually as cage birds; ‘the effect of the trade on the wild population is not known and must be of some concern’ (Parker 1999). Description. ADULT Y: forehead to hindneck pale greyish brown with sharp narrow blackish brown streaks; mantle, scapulars and back light buffy brown with broad dark brown streaks; rump lemon yellow; uppertail-coverts dark brown, tipped pale yellow. Tail feathers dark brown, outer webs with narrow buff edges, tinged olive-green basally; inner webs edged and tipped white, those of T4–T5 with broad white tips, and of T6 with distal half to two-thirds white. Supraloral spot white, tinged yellow; no superciliary stripe. Lores, cheeks and ear-coverts greyish brown, with pale yellow or buffy yellow patch on forecheek bordered below by broad dusky brown moustachial/malar stripe; sides of neck light greyish brown with diffuse darker streaking. Chin, throat and centre of upper breast dull lemon yellow, upper breast

478

FRINGILLIDAE

Serinus citrinipectus

Canary S. atrogularis, but distinguished by dark malar stripe, no black on throat. Voice. Tape-recorded (99, GIB). Short rapid song rather tuneless and monotonous for a canary, with slightly plaintive tone like white-eye Zosterops; faster and less sweet than Yellow-fronted Canary, less variable and ringing than Black-throated Canary. Call ‘trreet’, with same tone as song.

with some dusky grey mottling; sides of breast light greyish or cinnamon-buff with diffuse darker streaks; lower breast to undertail-coverts white, washed with cinnamon; flanks cinnamonbuff with faint dusky grey streaks. Remiges blackish brown, primaries finely edged olive-green, secondaries and tertials narrowly edged and tipped buff; alula and primary coverts blackish brown, narrowly edged and tipped buff; rest of upperwing-coverts dark brown, greaters edged buff and these and medians broadly tipped buffy white to form 2 conspicuous wing-bars, lesser coverts broadly fringed buff. Underwing-coverts and axillaries pale cinnamon-buff. Bill dark flesh to dark horn, lower mandible paler flesh posteriorly; eyes dark brown; legs light brown. ADULT X: differs from Y in having chin and throat pale cinnamon-buff (not yellow) and upper breast light cinnamon brown with dusky mottling; supraloral spot and cheek patch pale buff; pale tips to upperwing-coverts tinged more cinnamon. SIZE wing, Y (n ¼ 25) 62–68 (650), X (n ¼ 7) 615–67 (640); tail, Y (n ¼ 25) 36–415 (381), X (n ¼ 7) 35–40 (374); bill, Y (n ¼ 9) 11–12 (115); tarsus, Y (n ¼ 9) 13–14 (135). Also, wing, Y (n ¼ 12) 63–69 (653), X (n ¼ 13) 62–67 (648); tail, Y (n ¼ 11) 36–39 (374), X (n ¼ 12) 34–39 (365). WEIGHT: Y (n ¼ 12) 96–127 (114), X (n ¼ 13) 105–130 (114), imm. (n ¼ 13) 104–122 (114) (Hanmer 2002). IMMATURE: juv. like ad. X but top of head buffier (less greyish); lower underparts whiter, without cinnamon wash; wing bars whiter. Wing bars of fledgling even more conspicuous (A). NESTLING: skin pale pink; sparse whitish down; bill pale yellowish white, mouth bright yellow (Brickell 1997).

Field Characters. Length 11–12 cm. A small canary with lemon-yellow throat, breast and rump; separated from Yellow-fronted Canary S. mozambicus, with which it associates, by distinctive face pattern (white spot above dark lores, dark malar stripe separating throat from white cheeks), and white underparts below breast. X similar but underparts buff, narrow white supercilium; drab colouration and contrasting yellow rump suggest Black-throated

General Habits. Inhabits clearings in dry woodland, thornveld/palm savanna with Syzygium cordatum trees, grassland, coastal flats, Brachystegia scrub, and cultivated land below 750 m; along roadsides and often around human habitation (Zimbabwe). Associated with Ilala palms Hyphaene natalensis, nesting in them and making nests from palm materials in KwaZulu-Natal (Chittenden 1998) although that may not be the case in S Mozambique (Parker 1999); in SE Zimbabwe ‘Ilala palms were not immediately obvious in the vicinity [of large flocks] but ˆi were numerous not too far away’ (Hustler 1991). In Malaw occurs in gardens and open woodland near water and associated with reedbeds and vegetation fringing them (Hanmer 1983); sometimes in dry areas away from water. Occurs in pairs, small parties, and flocks of up to 100, 160 and 250 or more birds. Forages on ground, in winter frequently with Yellow-fronted Canaries S. mozambicus (KwaZulu-Natal, Zimbabwe, Mozambique). Food. Seeds of herbs and grasses. In captivity eats seeds of Bidens pilosa, Sonchus oleraceus, Poa trivialis and Digitaria sanguinalis; buds and leaves of Flaveria bidentis, Stellaria sp. and Galinsoga parviflora. Captive birds readily take Trinervitermes termites.

Breeding Habits. Solitary nester; seems to be monogamous and territorial. In presence of X, Y sings at high intensity with neck stretched and wings held away from body (B). NEST: open cup, made mainly of dried flower petals of Ilala palm Hyphaene natalensis taken from old inflorescences and already conveniently matted together with spider web; cup lined with long hair-like fibres stripped from leaves of the same palm (4 nests: Chittenden 1998). Nest materials collected by both sexes. All sited in narrow axils of Ilala palm fronds; from 18 to 6 m above ground;

Serinus mozambicus

479

chewed bark bound with silk from caterpillar tubes’; cup diam. 45–55, lined with long thin fibres peeled from Ilala palm fronds. A nest of captive pair had its base made of grass stems, inflorescence tops, roots and large feathers, with cup made entirely of coir; cup diam. 40 and depth 30. EGGS: 3–4 (2 wild and 2 captive clutches). Laid at 24 h intervals. Subelliptical, smooth, glossy; immaculate white, or white sparsely marked with small red-brown spots mainly in belt around broad end, or white with sparse, inconspicuous, irregularly shaped spots, and short, pale rusty brown streaks (captivity: Lawson 1970, Robson 1990). SIZE: (n ¼ 1) 16  12. LAYING DATES: Zimbabwe, Jan (and courtship feeding ˆ i, (various indications, and enlarged gonads, Nov); Malaw Dec–May with peak in Jan–Feb: Hanmer 2002); KwaZuluNatal, late Dec, Jan. INCUBATION: by X only; eggs left unattended when X departs to feed; period 12–14 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: nestling period in captivity 14–16 days. well camouflaged, totally obscured from view except from narrow slit in leaf axil, immediately above nest. Another Ilala palm nest (Robson 1990) was sited not in frond axil but on midrib in deep V-shaped through between leaflet rows; nest base 60  65, made of short fibres from dead creeper stems, bits of a downy leaf, and ‘fine material of

Key References Brickell, N. (1997), Brickell, N. and Konigkramer, T. (1997), Chittenden, H. (1998), Clancey, P.A. and Lawson, W.J. (1960), Clement, P. et al. (1993), Hanmer, D.B. (2002), Harrison, J.A. et al. (1997), Hustler, K. (1991), Irwin, M.P.S. (1971, 1979).

Serinus mozambicus (S. Mu ¨ ller). Yellow-fronted Canary; Yellow-eyed Canary. Serin du Plate 29 Mozambique. (Opp. p. 458) Fringilla mozambica S. Mu ¨ ller, 1776. Natursyst., suppl., p. 163; Mozambique. Range and Status. Common endemic resident and seasonal wanderer, subsaharan Africa except rainforest and arid zones. Introduced into Sa˜o Tome´ I., Mafia I., Mauritius, Re´union, Rodrigues (Showler 2002), Assumption I., Hawaii and Puerto Rico. Extinct in Amirante Is. Mauritania, frequent southeast of Boghe´, in upper Senegal Valley and south of Guidimaka; in rainy season, occurs north to 16
300 N. Senegal, frequent almost throughout, common in Parc Nat. des Oiseaux du Djoudj but rare elsewhere in far N. Gambia, frequent on coast, common inland, especially in forest reserves in lower river district; thought to have expanded since 1970s. Guinea, occurs in Gaoual and Koundara areas and east to Doko, common in Haut Niger Nat. Park, widespread in coastal and Kindia areas. Sierra Leone, frequent in Outamba area; records from Tembikunda and Kundewarakaro and probably very common throughout the north (Field 1974); a common cage-bird in Freetown. Liberia, locally frequent along N borders; became commoner on Nimba from mid 1960s; occurs around Monrovia. Mali, very common in Boucle du Baoule´ Biosphere Res., abundant in Bani Valley and in gardens around Bamako, elsewhere quite common south of 15
N. Ivory Coast, common throughout, north of forest zone. Burkina Faso, common north at least to 12
N, frequent in Ouagadougou area and in S of Nord-Yatenga region. Ghana, common throughout, except for forest

zone; occurs south to Kumasi (escapees?) and Accra Plain; commonly kept as cage-birds in villages. Togo, common throughout savanna zones, south to Notse´. Benin, common at least in Be´te´rou area, frequent in Pendjari Nat. Park. Niger, frequent resident in ‘W’ Nat. Park; occurs north to Gaya and Tapoa (12
290 N, 2
250 E). Nigeria, absent from sahel zone in NE, occasional as far north as Kazaure, Katsina and Kano in N, frequent around Falgore and Bagauda, further south a locally common and widespread resident, south to Ibadan, Enugu and Afikpo; locally a popular cagebird and numbers may be diminishing because of collecting for the trade (Elgood et al. 1994); records at Ife and Lagos may be of escapees; frequent resident at Warri, perhaps also escapees, but it is likely that species is extending southwards in W Africa as forests are degraded. Cameroon, widespread outside forest zone, north to L. Chad; from 350 m (on Mt Kupe´, up to 900 m), 550 m (Ntui) to 800–950 m (around Yaounde´) and at 1400– 1550 m (Mt Manengouba). Chad, widespread in SW at least near L. Chad and in Chari region; uncommon in soudanian zone (Salamat, Am Timam, Zakouma) and, as throughout most of its range, decidedly ‘anthropophile’ (Salvan 1968–1969). Central African Republic, frequent to common in all open country north to Bamingui-Bangoran Nat. Park; uncommon in N Vakaga region; south to Lobaye Pre´fecture. Sudan, barbatus widespread in Jebel

Serinus mozambicus

479

chewed bark bound with silk from caterpillar tubes’; cup diam. 45–55, lined with long thin fibres peeled from Ilala palm fronds. A nest of captive pair had its base made of grass stems, inflorescence tops, roots and large feathers, with cup made entirely of coir; cup diam. 40 and depth 30. EGGS: 3–4 (2 wild and 2 captive clutches). Laid at 24 h intervals. Subelliptical, smooth, glossy; immaculate white, or white sparsely marked with small red-brown spots mainly in belt around broad end, or white with sparse, inconspicuous, irregularly shaped spots, and short, pale rusty brown streaks (captivity: Lawson 1970, Robson 1990). SIZE: (n ¼ 1) 16  12. LAYING DATES: Zimbabwe, Jan (and courtship feeding ˆ i, (various indications, and enlarged gonads, Nov); Malaw Dec–May with peak in Jan–Feb: Hanmer 2002); KwaZuluNatal, late Dec, Jan. INCUBATION: by X only; eggs left unattended when X departs to feed; period 12–14 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: nestling period in captivity 14–16 days. well camouflaged, totally obscured from view except from narrow slit in leaf axil, immediately above nest. Another Ilala palm nest (Robson 1990) was sited not in frond axil but on midrib in deep V-shaped through between leaflet rows; nest base 60  65, made of short fibres from dead creeper stems, bits of a downy leaf, and ‘fine material of

Key References Brickell, N. (1997), Brickell, N. and Konigkramer, T. (1997), Chittenden, H. (1998), Clancey, P.A. and Lawson, W.J. (1960), Clement, P. et al. (1993), Hanmer, D.B. (2002), Harrison, J.A. et al. (1997), Hustler, K. (1991), Irwin, M.P.S. (1971, 1979).

Serinus mozambicus (S. Mu ¨ ller). Yellow-fronted Canary; Yellow-eyed Canary. Serin du Plate 29 Mozambique. (Opp. p. 458) Fringilla mozambica S. Mu ¨ ller, 1776. Natursyst., suppl., p. 163; Mozambique. Range and Status. Common endemic resident and seasonal wanderer, subsaharan Africa except rainforest and arid zones. Introduced into Sa˜o Tome´ I., Mafia I., Mauritius, Re´union, Rodrigues (Showler 2002), Assumption I., Hawaii and Puerto Rico. Extinct in Amirante Is. Mauritania, frequent southeast of Boghe´, in upper Senegal Valley and south of Guidimaka; in rainy season, occurs north to 16
300 N. Senegal, frequent almost throughout, common in Parc Nat. des Oiseaux du Djoudj but rare elsewhere in far N. Gambia, frequent on coast, common inland, especially in forest reserves in lower river district; thought to have expanded since 1970s. Guinea, occurs in Gaoual and Koundara areas and east to Doko, common in Haut Niger Nat. Park, widespread in coastal and Kindia areas. Sierra Leone, frequent in Outamba area; records from Tembikunda and Kundewarakaro and probably very common throughout the north (Field 1974); a common cage-bird in Freetown. Liberia, locally frequent along N borders; became commoner on Nimba from mid 1960s; occurs around Monrovia. Mali, very common in Boucle du Baoule´ Biosphere Res., abundant in Bani Valley and in gardens around Bamako, elsewhere quite common south of 15
N. Ivory Coast, common throughout, north of forest zone. Burkina Faso, common north at least to 12
N, frequent in Ouagadougou area and in S of Nord-Yatenga region. Ghana, common throughout, except for forest

zone; occurs south to Kumasi (escapees?) and Accra Plain; commonly kept as cage-birds in villages. Togo, common throughout savanna zones, south to Notse´. Benin, common at least in Be´te´rou area, frequent in Pendjari Nat. Park. Niger, frequent resident in ‘W’ Nat. Park; occurs north to Gaya and Tapoa (12
290 N, 2
250 E). Nigeria, absent from sahel zone in NE, occasional as far north as Kazaure, Katsina and Kano in N, frequent around Falgore and Bagauda, further south a locally common and widespread resident, south to Ibadan, Enugu and Afikpo; locally a popular cagebird and numbers may be diminishing because of collecting for the trade (Elgood et al. 1994); records at Ife and Lagos may be of escapees; frequent resident at Warri, perhaps also escapees, but it is likely that species is extending southwards in W Africa as forests are degraded. Cameroon, widespread outside forest zone, north to L. Chad; from 350 m (on Mt Kupe´, up to 900 m), 550 m (Ntui) to 800–950 m (around Yaounde´) and at 1400– 1550 m (Mt Manengouba). Chad, widespread in SW at least near L. Chad and in Chari region; uncommon in soudanian zone (Salamat, Am Timam, Zakouma) and, as throughout most of its range, decidedly ‘anthropophile’ (Salvan 1968–1969). Central African Republic, frequent to common in all open country north to Bamingui-Bangoran Nat. Park; uncommon in N Vakaga region; south to Lobaye Pre´fecture. Sudan, barbatus widespread in Jebel

480

FRINGILLIDAE

Serinus mozambicus

?

?

Marra and in S Kordofan (populations possibly isolated from those further south); frequent to common south of 9
N, but seemingly absent from Shilluk, Sobat, Sudd and most of Jonglei regions; grotei occurs in SE Blue Nile and Boma districts, near Ethiopian border. Eritrea, locally common at 1650–2300 m in central plateau and Mareb R. catchment (Zinner 2001); Eritrea and N Ethiopia, very local in Tacazze and Mareb Valleys below 1500 m and around Chenafena at 1675 m; Ethiopia south of 12
N, frequent to abundant in W lowlands and highlands east to about 37
300 E in N and 37
E in S: in W Gojam, Welega, Ilubabor, Kefa and Gamo Gafo Provs, absent from Shoa and Sidamo Provs, also from Nechisar Nat. Park, and rare in Omo Valley. Sa˜o Tome´, introduced in late 19th century, common, widespread in N west to Ponta Figo, inland to Nova Moca, frequent in lowlands in S. Gabon, known from Port Gentil; common around Le´koni, Moukalaba and Tchibanga (Sargeant 1993); in Re´s. de la Lope´? (Christy and Clarke 1994). Congo, widespread in S Mayombe and along lower Congo R., otherwise uncommon: 3 records as mapped. Zaı¨re, lower Congo R.; frequent and widespread in Kasai Occidental and S Kasai Oriental, common in Katanga and S Kivu Provs; occurs in clearings near eastern borders; absent from Volcanoes area, uncommon in Rutshuru Valley; in NE, common around Medje and Bafwabaka and abundant in Uele grasslands. Uganda, common throughout, at all elevations up to 2300 m. Kenya, fairly common resident and wanderer, up to 2200 m, from Mt Elgon and Saiwa Nat. Park to Busia, Mumias, Kakamega, Nandi, L. Victoria shores, Sotik, Lolgorien and Mara Game Res.; also in coastal lowlands north to lower Tana R. (Baomo) and Lamu and Manda Is, south to Shimba Hills. Somalia, single record near mouth of Jubba R. (Hall and Moreau 1970). Rwanda, frequent to common, except perhaps in

NE. Burundi, very common throughout, up to 2000 m. Tanzania, uncommon to common, widespread as mapped (N. Baker, pers. comm.); Tabora (barbatus), Katavi Nat. Park (race?), Ufipa Plateau (samaliyae); Mafia I., introduced in late 19th century (tando), several old records in N and evidently still there since local people ‘know the bird, trap and cage it but state that it is uncommon in coral-rag bush country where millet is grown’ (Pakenham 1979). Angola, common: tando in Cabinda and from Huambo and N Huı´la to Uı´ge, Cuanza Norte, Malanje, Bie´, Lunda Norte and Lunda Sul, samaliyae in N and E Moxico, vansoni from S and central Huı´la and E Namibe to Cunene ˆ i, and Cuando Cubango. Throughout Zambia, Malaw Mozambique (granti south of Inkomati R. floodplain, mozambicus to the north), Zimbabwe, Transvaal (except W highveld; commonest in Escarpment region), KwaZuluNatal and E Cape Province, at all altitudes up to c. 1600 m. Free State, rare to uncommon, regular between Golden Gate and Harrismith and in Vaal Valley between Parys and Sasolburg. Botswana, uncommon to locally common in N, E and SE; very sparse in Okavango Delta. Lesotho, widespread in lower country. Ranges of most subspecies are entirely contiguous. Widely kept and traded as a cagebird over much of its range; amongst the people of N Mozambique, for instance, a ‘universal pet’ 70 years ago (Vincent 1936) and doubtless today too; in South Africa ‘almost every house in some native villages has its canary in a rough cage . . . where road workers carried their birds to work with them, a row of cages bedecked the roadside’ (Skead 1960). Easily caught using live-baited double traps, often in towns; also by birdliming (Inhaca I., Mozambique). Up to 10,000 birds exported annually under permit in S Mozambique (Sul do Save), whilst ‘unknown numbers are captured and sold illegally . . . unlikely that the trade has a significant effect on the wild population, which probably exceeds two million birds’ (Parker 1999). Commercial trapping quotas are also allocated in Botswana. Density in Sul do Save of 10 birds per 100 ha in acacia, 13 in mopane, 21 in miombo and 10 in other broad-leaved woodlands (ibid.). Density in Acacia savannas, Swaziland, 10, 36, 105 and 1280 birds per 100 ha (Monadjem 2002). Description. S. m. barbatus Heuglin (includes ‘pseudobarbatus’): S Chad to Sudan, west of Nile, and NE Zaı¨re to W Kenya and central Tanzania. ADULT Y: forehead bright golden yellow; crown to back and scapulars yellowish olive-green, crown to hindneck with narrow dusky streaks, mantle, scapulars and upper back with short broader blackish streaks; rump golden yellow; uppertail-coverts olive-green, tinged yellow. Tail feathers blackish brown, outer webs edged olive-green; tip of T1 and tips of outer webs of T2–T6 broadly fringed greenish white. Crown bordered by narrow bright yellow superciliary stripe; below this, a dark olive-grey stripe through lores and broader olive-grey band through ear-coverts; cheeks bright yellow; sides of neck yellowish olive-green; broad blackish malar stripe. Sides of breast olivegreen and lateral flanks tinged green; underparts otherwise bright golden yellow. Remiges blackish brown, primaries and secondaries edged yellowish olive-green, tertials broadly edged and tipped olive-green; alula and primary coverts blackish brown, narrowly fringed olive-green; rest of upperwing-coverts blackish brown, greaters edged olive-green, and these and medians tipped greenish yellow to form 2 narrow and inconspicuous wingbars,

Serinus mozambicus

lesser coverts tipped olive-green. Underwing coverts and axillaries pale yellow. Bill short, conical, robust (A), dark brown above, greyish brown below; eyes dark brown; legs dark brown. ADULT X: similar to ad. Y, but duller olive-green above; forehead, cheeks and underparts paler yellow; lores and malar stripe dark olive-green rather than blackish. SIZE (10 YY, 10 XX): wing, Y 61–71 (665), X 64–71 (662); tail, Y 36–42 (393), X 38–43 (402); bill, Y 115–125 (118), X 11–12 (117); tarsus, Y 135–145 (141), X 135–145 (145). Birds of W Kenya, Burundi and Tanzania (‘pseudobarbatus’) average larger: wing, Y (n ¼ 7) 71–73 (720). WEIGHT: W Uganda, (n ¼ 16, unsexed) 112–132 (120); W Kenya, Y (n ¼ 8) 115–134 (129), X (n ¼ 10) 12–14 (129), unsexed (n ¼ 53) 11–171 (130). IMMATURE: juv. browner above than ad., more heavily streaked; underparts paler yellow, chin to breast tinged buff, sides of breast and flanks suffused olive-brown, upper breast with dark brown spots and flanks with narrow streaks; edges of tail feathers, secondaries and tertials, and edges and tips of greater and median coverts buffy yellow. NESTLING (nominate mozambicus): at hatching, skin pink, greybrown down on crown, occiput, humerals, secondaries, back, rump, crurals and belly, legs orange, bill orange with brown tip, egg-tooth off-white, mouth scarlet, eyes closed. S. m. caniceps (d’Orbigny): Senegal to Cameroon south to Benue plain. Top of head, ear-coverts and sides of neck grey (rather than green); mantle and scapulars paler, duller olive-green than in barbatus; frontal band narrower, yellow of supercilium, cheeks, chin and throat less intense. X has throat spotted blackish. Wing, Y (n ¼ 9) 61–70 (643). WEIGHT: Ghana, Y (n ¼ 40) av. 108, X (n ¼ 31) av. 104, unsexed (n ¼ 45) 70–130 (102). S. m. punctigula Reichenow: Cameroon, north to Toukte, Grand Capitaine and Koum (8
200 –8
350 N). Similar to barbatus, but upperparts darker olive-green (less yellowish); ear-coverts darker; frontal band narrower; crown greener than in caniceps; X has blackish throat spotting like caniceps. Slightly larger than typical barbatus: wing, Y (n ¼ 10) 68–73 (699). WEIGHT: (n ¼ 8, unsexed) 13–15 (135). S. m. gommaensis Mackworth-Praed and Grant: W Ethiopia (L. Tana to Gomma). Olive-green upperparts darker, tinged browner than in barbatus. Sides and flanks darker than in grotei, and frontal band narrow. WEIGHT: unsexed (n ¼ 43) 98–124 (109). S. m. grotei Sclater and Mackworth-Praed: Sudan east of Nile to Eritrea and W Ethiopia (Baro R.). Yellowish green above like barbatus, but paler and tinged browner; eyestripe and malar stripe paler, more greyish; slightly paler below. Frontal band broad. ˆ i and central S. m. mozambicus (Mu ¨ ller): coastal Kenya to Malaw Mozambique (west in Tanzania to Moshi, Kilosa and Songea), and Zambia (except N and SW) to Zimbabwe, E and SE Botswana, N and W Transvaal and Free State. Dull olive-green above with dark brown streaking, heavier than in barbatus, crown, neck and rear of ear-coverts tinged greyish; yellow frontal band and superciliary stripe broad; eye-stripe dark olive-brown, malar stripe black; underparts golden yellow with sides of breast brownish olive (less green). Wing, Y (n ¼ 8) 66–70 (680). WEIGHT: Zimbabwe, Y (n ¼ 11) 93–128 (114), X (n ¼ 17) 100– 133 (116); monthly means of ads vary from 112 (Feb) to 120 (Apr–May) and imms from 111 (July–Aug) to 119 (Oct–Dec) (Hanmer 2002). S. m. granti Clancey: S Mozambique (south of Inkomati R.), SE Transvaal and E Swaziland to KwaZulu-Natal and E Cape Prov.

Similar to nominate race, but deeper green above, especially on crown, streaking slightly darker and heavier; ear-coverts very dark olive-green. WEIGHT (all South African races): Y (n ¼ 97) 93–162 (118), X (n ¼ 45) 100–136 (116), unsexed (n ¼ 110) 85–162 (133) (Maclean 1993). S. m. vansoni Roberts: extreme SE Angola and adjacent Namibia to SW Zambia (Barotseland north to Balovale) and N Botswana. Rather paler than nominate race, more greyish green (less brownish), streaking on head less heavy. S. m. samaliyae White: SE Zaı¨re (Katanga and Lomami), SW Tanzania (Ufipa and Rukwa) and adjacent Zambia (Mwinilunga and Kabompo to Luapala region and L. Mweru). Bright olivegreen above like barbatus, but mantle and scapulars strongly streaked as in nominate race; ear-coverts dark olive-green; underparts golden yellow. Large: wing, Y (n ¼ 5) 70–71 (706). S. m. tando Sclater and Mackworth-Praed: Gabon, Congo, W and S Zaı¨re (Kasai) and Angola (except extreme SE); introduced to Sa˜o Tome´ (population formerly named ‘santhome’) and Mafia I. Intergrades in Zaı¨re with samaliyae. Similar to samaliyae, but darker green above, sides of breast and flanks greener.

Field Characters. Length 10–13 cm. A common and familiar small canary, with bright yellow underparts and rump and short, white-tipped tail; face pattern diagnostic: broad yellow forehead, supercilium and face offset by black eyeline and malar stripe. X less bright, face stripes grey rather than black; juv. duller and greyer above, buffy yellow below, but still with vestigial face pattern; breast streaked, well-marked buff wing-bar. Some races of larger Yellow Canary S. flaviventris have similar face pattern but streaks are olive and broader, and rump green, not yellow. Separated from Lemon-breasted Canary S. citrinipectus by yellow supercilium, green back and entirely yellow underparts. Confusion possible with nominate race of Whitebellied Canary S. dorsostriatus (Zimmerman et al. 1996); for differences, see that species. Voice. Tape-recorded (86, 88, 99, 104, B, C, F, LEM, PAY). Variety of sweet whistled calls, ‘see-woo’, ‘way-tseeway’, ‘way-tsrrr’, ‘toway-towee’, ‘toway-pu-tseep’, often strung together in leisurely manner to form song of separate short phrases; much variation, but individual tends to repeat a theme; one bird introduced most songs with ‘see-were-chi-wu’ and went on to add 3–4 other phrases, another repeatedly used ‘see-see-suwayo-tyoop, suwayo-tyoop’ as part of longer song. Some songs have liquid trills and twitters, others none at all. Less often sings in a continuum, but more slowly than other canaries. ‘Ordinary’ calls, quiet, unmusical ‘chirrip’, ‘chrup’ or ‘zizzit’; chicks in nest make quiet churring; fledglings maintain husky and tinny ‘kizzkizzkizz’ or rapid, husky hunger call ‘chachacha . . . ’ (Skead 1960). General Habits. Inhabits a wide variety of open woodlands, tree savanna and open countryside: miombo and other dense woods, clearings, plantations, acacias and other thorn trees, rough grassland with scattered trees and shrubs, riverside bushes, parks, arable land especially with pearl millet fields and standing pollarded trees, pasture, around farmsteads and rural hamlets, in gardens, on muddy paths and sandy verges, amongst coarse grasses in marshes, and in seaside scrub and sand-dunes. On Inhaca

481

482

FRINGILLIDAE

I., Mozambique, in forest, swamps, fields, whistling-pines and often in mangrove. Freely uses elevated perches, e.g. telephone wires, fences, trees and tall grasses; enters cowsheds and farm outbuildings for grain, and feeds at bird-tables. Solitary or usually in pairs or threes; often in parties of 5–10 birds, occasionally 20–100. Active, attractive and ‘playful’. Forages mainly but not exclusively on ground; gait a hop, but can also walk. Commonly forages with other serin spp. and with viduines and waxbills. To take seeds in situ at tip of tall grass stem, bird alights half-way up so that it droops under bird’s weight, sidesteps towards seedhead until it is on ground and then pecks at seeds; often stem springs up and the bending process has to be repeated (Skead 1960). Removes Casuarina seeds from small cones. Feeds at head of Sorghum by stripping sheath away and eating only a part of the grain exposed. Takes aphids on fruit trees; catches termites on the wing, returning to perch with one and rubbing it against perch to remove its wings. Flight bounding and erratic. Birds in flock are very vocal and often dash at and chase each other, posture and gape; on breaking away from such activities, 2 birds sometimes flutter towards their perches rather slowly, on rapidly beating wings; 2 birds in flight often flutter down together breast to breast; birds flying in flock utter ‘sweeeet’ notes. Roosts in flocks of up to 10, in trees; much singing and twittering before birds settle down; flock does not use regular roost site. Drinks and bathes at puddles communally. Choruses of several singing birds happen at any time of day over a period of several months. When singing at an elevated perch, bird holds head high, slightly fans tail, and slowly sways body from side to side. Resident, but wanders seasonally and in some places almost a migrant, e.g. in Nguuni, Mombasa, Kenya, occurs regularly in Dec–Mar, irregularly in Nov and Apr-July, absent in July–Oct. Little seasonal variation in reporting rates in southern Africa, but wanders seasonally in Botswana, and vagrants sometimes appear in E Karoo. Food. Small seeds; also petals, fruits, small leaves, nectar and insects. Favours soft achenes of composites, e.g. Ursinia and Vernonia (South Africa). Small figs (Sierra Leone). Seeds of Eucalyptus, Helianthus annuus, Sorghum caffrorum, Panicum maximum, Casuarina equisetifolia; Kniphofia petals; leaves of Hibiscus rosasinensis. Probes for nectar at base of tubular corallas of Aloe marlothii, A. candelabrum, A. arborescens and Strychnos innocua. In captivity, takes seeds of Alternanthera pungens, Bidens pilosa, Sonchus oleraceus, Poa trivialis and Panicum laevifolium; buds, petals and leaves of Senecio tamides, Alternanthera ficoidea and Flaveria bidentis (Brickell and Konigkramer 1997). Eats fly larvae, aphids and small caterpillars. Termites and grasshoppers, as well as considerable mass of seeds, fed to young. Breeding Habits. Solitary nester or somewhat colonial; monogamous; territorial. ‘Colonies’ of 2–3 nests only a few m apart in same tree or adjacent trees are unusual but have been found many times. Courting Y does not have usual serin ‘butterfly’ flight. One chases another, both flying not fast but with rather jerky, airy, level flight, interspersed

with bouts of rapid wingbeats and with occasional singing. 4–5 birds hop through branches in pursuit of each other. NEST: very small, regular cup, very compactly built of fine grass and weed stems or thin soft petioles, soft plant fibres (stripped from herbs, once from a rope), white plant down, and wool, secured with cobweb, or with the outside plastered with cobweb and a few small dead leaves, looking silvery, and scantily lined with very fine plant fibres and a few hairs. SIZE: ext. diam. 60–70, int. diam. 40–44, ext. depth 30–34, int. depth 24–31 (Nigeria, Serle 1940); int. diam. (n ¼ 3) 50 (Zaı¨re); ext. diam. av. 68, int. diam. av. 47, ext. depth av. 50, int. depth av. 33 (n ¼ 4, South Africa). Placed 1–4 (usually 2) m above ground, in shrubs and trees, in a fork or where intersecting twigs and foliage give support and to which nest is firmly bound by cobweb; often near end of branch at side of small tree or towards its top, usually well screened, sometimes exposed to view; often in bushy top of pollarded sapling, or small Eucalyptus tree or Protea bush, Acacia thicket, street tree in town or tall pine; once in bunch of green bananas; on sloping frond of young oil-palm; in bunch of mistletoe. Built by X, the Y accompanying her on collecting trips and singing whilst she builds. But YY of 3 pairs seen also to gather nesting material (from old nest of Red-winged Starling Onychognathus morio, from an old rope on ground, and from tree bark fibres: Skead 1960). EGGS: 2–3, mainly 3 (Nigeria); 1–4, av. 27 (33 clutches, ˆ i); 2–5, av. 32 (86 clutches, southern Africa). Laid Malaw on successive days. Ovate, slightly glossy; pure white, or greyish white bluish white or pale cream, immaculate or sparingly speckled and spotted with fine orange-brown or dark brown spots and faint, pale lilac shell-marks, the latter confined to large end (Serle 1940). SIZE: (n ¼ 8, Nigeria) 150–166  118–123 (157  120); (n ¼ 9, Zimbabwe) 174–180  121–130 (165  127). LAYING DATES: Mauritania, July–Oct; Gambia, Nov; Liberia, (sings in Apr); Mali, July–Oct; Burkina Faso, Yatenga, (carrying nest material, Sept); N Ghana, (nestbuilding May, dependent juvs Aug, Nov); Niger, Oct; Nigeria, Wukari, July–Aug, Owerri, Aug–Oct, Kafanchan, Oct–Nov; Sudan, Sept; E Africa, Region A, Jan, Region B Jan, Feb, Apr–July, Oct, Dec, Region C, Nov–June, mainly Feb–Apr (15 out of 22 clutches), Region D, Jan, May, June, Region E, Apr, June-Aug; Zaı¨re, Katanga, (breeding condition, Feb), Uele, Aug–Oct; Angola, Apr; Zambia, ˆ i, Dec– Dec–Apr, Aug–Sept, mainly Feb and Aug; Malaw May, July, mainly Feb–May; Botswana, Dec; Zimbabwe, Nov–Apr (131 clutches: Dec 22%, Jan 30%, Feb 17%, Mar 21%); N Mozambique, Mar–May; Transvaal, Oct–Apr; KwaZulu-Natal, Sept–Apr; E Cape Prov., mainly Apr–May. INCUBATION: by X only, sitting closely, Y often singing from prominent perch nearby. Y feeds her on nest. Period: (n ¼ 4) 13–145 days (134). DEVELOPMENT AND CARE OF YOUNG: on day 5, primary quills showing; day 6, quills show in secondaries and primary and secondary coverts; eyes starting to open; day 7, back and breast quills showing; day 8, all other quills showing, feathers emerging on flanks, crurals, breast and belly, eyes half open; day 9–10, all feathers emerging except on chin; day 11, chin feathers emerging. At first,

Serinus dorsostriatus brooding X fed by Y; later, young fed by both parents. For first few days parents swallow nestling faeces, then they are deposited on nest rim and left there. Nestling period (n ¼ 7) 16–24 days (192). Young out of nest fed by both parents and keep with them for much of time, the family moving as a compact unit. BREEDING SUCCESS/SURVIVAL: survival rate of birds >6 months old, 65% per year, so life span after first 6 months

483

is 23 years, though some birds live to 6–7 years (Hanmer 2002). Greatest elapsed time of wild bird 85 years (Oschadleus 2000). A disabled ad. lived in captivity for 165 years (Liversidge 1971). Key References Clement, P. et al. (1993), Hanmer, D.B. (2002), Irwin, M.P.S. (1979), Serle, W. (1940), Skead, C.J. (1960), Maclean, G.L. (1993), Vincent, A.W. (1949), Vincent, J. (1936).

Serinus dorsostriatus (Reichenow). White-bellied Canary. Serin a` ventre blanc.

Plate 29

Crithagra dorsostriata Reichenow, 1887. J. Orn., 35, p. 72; Wembere, Tabora, Tanganyika fide W.L. Sclater, 1930, Syst. Avium Aethiop., p. 815.

(Opp. p. 458)

Range and Status. Endemic resident, Horn of Africa and E Africa. Ethiopia, common in Rift Valley, NE, S and SE districts; common at 1220–1370 m in Arero Forest, around Mega and in Yabello Sanctuary and Hills, frequent in Omo Nat. Park, Awash Nat. Park, Awash R. Valley and DawaWachile. Somalia, locally common in NW; in S, known mainly from middle Shabeelle Valley, with scattered records elsewhere. Sudan, only in Natoporoputh Hills (4
520 N, 34
000 E). Uganda, in E Karamoja and on NE shores of L. Victoria. Kenya, mainly at 400–1400 m, but up to 1800 m in Narok and 2650 m on Mt Nyiru; local in NW, south to foothills of highlands, and locally common in hot plains from Loita and Narok east to Amboseli and West and East Tsavo Nat. Parks. Tanzania, widespread from Loliondo to Serengeti Nat. Park, Crater and Mbulu Highlands, Lake Manyara Nat. Park, Tarangire Nat. Park, Masai Steppe, Arusha and Moshi Districts, Same, Mkomazi Game Res. and foot of Mt Meru, south to Tabora, Singida, Dodoma and Geiro (Kilosa) and west to Shinyanga, Sibiti R., Mwanza and Biharamulo; common near Dar es Salaam (population derived from escapees from the cagebird trade: N.E. Baker, pers. comm.); otherwise rare south of 5
300 S. Description. S. d. maculicollis Sharpe: Somalia, Ethiopia, Sudan, NE Uganda and N Kenya (south to Equator). (See Nomenclatural Note below.) Frontal band pale yellow, connecting with broad pale yellow superciliary stripe; crown to back pale dull olivegreen, streaked blackish brown, broadly so on mantle and scapulars; rump lemon yellow; uppertail-coverts olive-green with dusky shaft streaks. Tail feathers blackish brown, outer webs edged olive-yellow, tips of T5–T6 narrowly fringed white. Lores to ear-coverts, upper cheeks and sides of neck pale dull olive-green; lower cheeks pale yellow bordered below by broad olive-green malar stripe. Chin to breast yellow, sides of breast olive-green with fine dusky streaking; flanks white with short broad dark brown streaks; belly and undertail-coverts white. Remiges dark brown, primaries and secondaries narrowly edged yellowish olive-green, tertials edged pale olive-green; alula and upperwing-coverts dark brown, greater coverts edged olivegreen and tipped yellowish white, median and lesser coverts broadly tipped pale yellowish green. Underwing-coverts and axillaries pale yellow. Bill horn-brown or sepia above, pale horn or whitish below; eyes brown; legs dark grey-brown to black. ADULT X: similar to ad. Y, but cheeks, ear-coverts and ground colour of upperparts tinged browner; underparts paler yellow,

Serinus dorsostriatus

with chin and throat whitish and gorget of large blackish brown spots across upper breast. SIZE (10 YY, 10 XX): wing, Y 67–74 (709), X 67–74 (691); tail, Y 45–50 (480), X 45–49 (473); bill, Y 12–13 (124), X 115–125 (120); tarsus, Y 155–17 (162), X 15–17 (162). WEIGHT: unsexed (n ¼ 37) 120–173 (145). IMMATURE: juv. differs from ad. X in having crown to back and scapulars streaked buffish and dark brown; underparts buffy white, tinged brown on throat and breast, and spotted dark brown across upper breast and on sides of breast and flanks. NESTLING: downy, with very dark skin. S. d. taruensis van Someren: central, S and SE Kenya (south of Equator) to NE Tanzania (west to Arusha). Y brighter green above than maculicollis; yellow on underparts more intense, extending to upper belly; frontal band and superciliary stripe slightly broader. Larger: wing, Y (n ¼ 6) 71–76 (732). S. d. dorsostriatus Reichenow: SE Uganda (Kampala, Busoga), W Kenya (Kakamega, Nyanza) and NW Tanzania. Like taruensis but more yellowish green above; white below limited to vent area; belly and undertail-coverts pale yellow; edges of tail feathers and edges and tips of greater coverts pale olive-buff (not green).

Serinus dorsostriatus brooding X fed by Y; later, young fed by both parents. For first few days parents swallow nestling faeces, then they are deposited on nest rim and left there. Nestling period (n ¼ 7) 16–24 days (192). Young out of nest fed by both parents and keep with them for much of time, the family moving as a compact unit. BREEDING SUCCESS/SURVIVAL: survival rate of birds >6 months old, 65% per year, so life span after first 6 months

483

is 23 years, though some birds live to 6–7 years (Hanmer 2002). Greatest elapsed time of wild bird 85 years (Oschadleus 2000). A disabled ad. lived in captivity for 165 years (Liversidge 1971). Key References Clement, P. et al. (1993), Hanmer, D.B. (2002), Irwin, M.P.S. (1979), Serle, W. (1940), Skead, C.J. (1960), Maclean, G.L. (1993), Vincent, A.W. (1949), Vincent, J. (1936).

Serinus dorsostriatus (Reichenow). White-bellied Canary. Serin a` ventre blanc.

Plate 29

Crithagra dorsostriata Reichenow, 1887. J. Orn., 35, p. 72; Wembere, Tabora, Tanganyika fide W.L. Sclater, 1930, Syst. Avium Aethiop., p. 815.

(Opp. p. 458)

Range and Status. Endemic resident, Horn of Africa and E Africa. Ethiopia, common in Rift Valley, NE, S and SE districts; common at 1220–1370 m in Arero Forest, around Mega and in Yabello Sanctuary and Hills, frequent in Omo Nat. Park, Awash Nat. Park, Awash R. Valley and DawaWachile. Somalia, locally common in NW; in S, known mainly from middle Shabeelle Valley, with scattered records elsewhere. Sudan, only in Natoporoputh Hills (4
520 N, 34
000 E). Uganda, in E Karamoja and on NE shores of L. Victoria. Kenya, mainly at 400–1400 m, but up to 1800 m in Narok and 2650 m on Mt Nyiru; local in NW, south to foothills of highlands, and locally common in hot plains from Loita and Narok east to Amboseli and West and East Tsavo Nat. Parks. Tanzania, widespread from Loliondo to Serengeti Nat. Park, Crater and Mbulu Highlands, Lake Manyara Nat. Park, Tarangire Nat. Park, Masai Steppe, Arusha and Moshi Districts, Same, Mkomazi Game Res. and foot of Mt Meru, south to Tabora, Singida, Dodoma and Geiro (Kilosa) and west to Shinyanga, Sibiti R., Mwanza and Biharamulo; common near Dar es Salaam (population derived from escapees from the cagebird trade: N.E. Baker, pers. comm.); otherwise rare south of 5
300 S. Description. S. d. maculicollis Sharpe: Somalia, Ethiopia, Sudan, NE Uganda and N Kenya (south to Equator). (See Nomenclatural Note below.) Frontal band pale yellow, connecting with broad pale yellow superciliary stripe; crown to back pale dull olivegreen, streaked blackish brown, broadly so on mantle and scapulars; rump lemon yellow; uppertail-coverts olive-green with dusky shaft streaks. Tail feathers blackish brown, outer webs edged olive-yellow, tips of T5–T6 narrowly fringed white. Lores to ear-coverts, upper cheeks and sides of neck pale dull olive-green; lower cheeks pale yellow bordered below by broad olive-green malar stripe. Chin to breast yellow, sides of breast olive-green with fine dusky streaking; flanks white with short broad dark brown streaks; belly and undertail-coverts white. Remiges dark brown, primaries and secondaries narrowly edged yellowish olive-green, tertials edged pale olive-green; alula and upperwing-coverts dark brown, greater coverts edged olivegreen and tipped yellowish white, median and lesser coverts broadly tipped pale yellowish green. Underwing-coverts and axillaries pale yellow. Bill horn-brown or sepia above, pale horn or whitish below; eyes brown; legs dark grey-brown to black. ADULT X: similar to ad. Y, but cheeks, ear-coverts and ground colour of upperparts tinged browner; underparts paler yellow,

Serinus dorsostriatus

with chin and throat whitish and gorget of large blackish brown spots across upper breast. SIZE (10 YY, 10 XX): wing, Y 67–74 (709), X 67–74 (691); tail, Y 45–50 (480), X 45–49 (473); bill, Y 12–13 (124), X 115–125 (120); tarsus, Y 155–17 (162), X 15–17 (162). WEIGHT: unsexed (n ¼ 37) 120–173 (145). IMMATURE: juv. differs from ad. X in having crown to back and scapulars streaked buffish and dark brown; underparts buffy white, tinged brown on throat and breast, and spotted dark brown across upper breast and on sides of breast and flanks. NESTLING: downy, with very dark skin. S. d. taruensis van Someren: central, S and SE Kenya (south of Equator) to NE Tanzania (west to Arusha). Y brighter green above than maculicollis; yellow on underparts more intense, extending to upper belly; frontal band and superciliary stripe slightly broader. Larger: wing, Y (n ¼ 6) 71–76 (732). S. d. dorsostriatus Reichenow: SE Uganda (Kampala, Busoga), W Kenya (Kakamega, Nyanza) and NW Tanzania. Like taruensis but more yellowish green above; white below limited to vent area; belly and undertail-coverts pale yellow; edges of tail feathers and edges and tips of greater coverts pale olive-buff (not green).

484

FRINGILLIDAE

W Kenya, Y (n ¼ 31) 137–177 (159), X (n ¼ 11) 14–177 (155). TAXONOMIC NOTE: treated as a subspecies of S. (Crithagra) flaviventris by E. Mayr in Paynter 1968 and by White (1963). Hall and Moreau (1970) gave reasons for allying it superspecifically with S. (Ochrospiza) mozambicus. Van den Elzen and Baars (1980) and van den Elzen (1985) have shown that dorsostriatus belongs in subgenus Ochrospiza, where we regard it as a dry-country derivative of ancestral mozambicus, no longer consuperspecific with it. NOMENCLATURAL NOTE: the type of maculicollis Sharpe, 1895, Bull. Br. Orn. Club iv, 41, is actually a specimen of S. xantholaemus (J.S. Ash and D. Pearson, pers. comm., 2002); xantholaemus was described in 1896, so the name maculicollis takes precedence for it and a substitute name is required for the bird called maculicollis above (M. Walters, pers. comm.). WEIGHT:

Field Characters. Length 115–13 cm. A dry-country canary of E Africa. Olive eyeline, ear-coverts and malar stripe on yellow face suggest Yellow-fronted Canary S. mozambicus but are more diffuse, less sharp, especially malar stripe. Northern race maculicollis (south to central Kenya) distinctive with white belly, flanks and undertailcoverts, but S birds have white areas washed with yellow, and could be mistaken for Yellow-fronted Canary (Zimmerman et al. 1996); they differ (besides face patch) in longer tail, streaks on flanks (and on breast in X). Voice. Tape-recorded (B, GREG, McVIC, PAY, PEAR). Song both sibilant and wheezy, short phrases, often repeated, ‘chu-weez, chu-weez’, ‘weez-cha, weez, cha’, ‘chercher-weez-cha’, ‘chicher, chicha-weez-cha’; sometimes includes brief rattle. Other songs described as ‘several emphatic sharp notes ending with sharp rising trill’ (Zimmerman et al. 1996) and ‘a sweet series of Goldfinch-like notes or trills, interspersed with some harsher phrases, and quite loud’ (Clement et al. 1993). Buzzy calls, ‘zwee-weez’ and ‘zweess’.

General Habits. Inhabits dry Acacia thorn bush and grassland; often near impenetrable clumps of thorn scrub and often near water (Somalia, Ash and Miskell 1998); in E Africa in bushy and wooded grassland, open woodland and cultivation in low and medium rainfall areas. Solitary, in pairs, or up to 6 birds together. Forages mainly on ground, also in low shrubs. Resident but prone to wander; an irregular visitor to S Kerio Valley, Kenya, in Aug–Dec. Food. Said to be mainly grass seeds. Breeding Habits. Solitary nester. NEST: small, usually very neat, sometimes untidy, delicate open cup, made of lichens, fibres, cobwebs and bents or dry grass and rootlets or twiglets, lined with hair, feathers or plant down and fine grass; one was smooth and red-brown at bottom of cup and black at sides. Int. diam. 45, int. depth 35. Sited generally at 25–35 m above ground on horizontal fork near end of branch in tree, often on plain with isolated or scattered clumps of trees; one in topmost clump of leaves of vertical branch of Boscia coriacea; one only 1 m up in small Maerua cassifolia tree; one 8 m high in thick foliage. EGGS: 1–4, av. (Kenya, 19 clutches) 295. Very pale blue, whitish or greenish blue, usually immaculate, sometimes with sparse reddish brown or blackish spots. SIZE: (n ¼ 8) 165–1925  127–1375 (180  131). LAYING DATES: Ethiopia, Yavello (Yabelo) Mar (and gonads slightly enlarged June, July; imms in June–Aug); Somalia, (young just out of nest Dec and Mar, nestbuilding Nov); E Africa, Region B, Mar–July, Sept, Nov, Jan (mainly Mar–June), Region C, May, June, Region D, Apr–June, Sept, Jan. INCUBATION: by X, sitting tight. Nothing further known. Key References van den Elzen, R. (1985), van den Elzen, R. and Baars, W. (1980).

Plate 29

Serinus flavigula Salvadori. Yellow-throated Seedeater. Serin a ` gorge jaune.

(Opp. p. 458)

Serinus flavigula Salvadori, 1888. Ann. Civ. Mus. Genova, 26, p. 272; Ambokarra, Shoa Prov., Abyssinia. Forms a superspecies with S. xantholaemus. Range and Status. Endemic resident, Shoa Prov., Ethiopia. Map: see Salvadori’s Seedeater S. xantholaemus. 3 specimens taken at Kolla di Aigaber (9
360 N, 40
040 E) in 1880, at Ambokarra (9
310 N, 40
090 E) in 1885 and at Malca Jebdu Shet (i.e. stream near Malca-Ghebdu, 9
340 N, 39
530 E); rediscovered at Malca-Ghebdu at 1400–1500 m in 1989, when parties of 3 then 4 birds seen on one day. The 3 localities are close together near foot of eastern escarpment of W Highlands, Shoa Prov., c. 20 km east of Ankober (see Ankober Serin S. ankoberensis) and enclose an area of only c. 30 km2. 6–7 found at 1310–1610 m on crater rim of Fantalle Mt (9
000 N, 39
540 E) in Awash Nat. Park, c. 65 km south of Malca-Ghebdu, in 1993; one seen in Awash Nat. Park and family at Malca-Ghebdu in 1996, one or

more in Mt Fantalle crater in 1997 and one at Malca Ghebdu in 1998 (Bull. Afr. Bird Club 5, 1998, 144 and 7, 2000, 46). Unconfirmed report of pair at Lalibela (Bull. Afr. Bird Club 3, 1996, 62). Description. ADULT: top of head grey-brown, forehead and crown with fine, short darker streaks, fading on nape and hindneck; mantle darker, warmer brown, with diffuse narrow streaks formed by dark brown feather centres; lower back and rump pale dull yellow (Malca-Ghebdu, Ash and Gullick 1989) or bright canary yellow (Fantalle Mt, Atkins and Harvey 1996), extending to rump sides and short uppertail-coverts; long uppertail-coverts greenish yellow, browner toward tips. Tail feathers dark greyish brown, edged buffy brown. Indistinct whitish superciliary stripes to just behind eye, meeting above bill; lores dark grey-brown;

484

FRINGILLIDAE

W Kenya, Y (n ¼ 31) 137–177 (159), X (n ¼ 11) 14–177 (155). TAXONOMIC NOTE: treated as a subspecies of S. (Crithagra) flaviventris by E. Mayr in Paynter 1968 and by White (1963). Hall and Moreau (1970) gave reasons for allying it superspecifically with S. (Ochrospiza) mozambicus. Van den Elzen and Baars (1980) and van den Elzen (1985) have shown that dorsostriatus belongs in subgenus Ochrospiza, where we regard it as a dry-country derivative of ancestral mozambicus, no longer consuperspecific with it. NOMENCLATURAL NOTE: the type of maculicollis Sharpe, 1895, Bull. Br. Orn. Club iv, 41, is actually a specimen of S. xantholaemus (J.S. Ash and D. Pearson, pers. comm., 2002); xantholaemus was described in 1896, so the name maculicollis takes precedence for it and a substitute name is required for the bird called maculicollis above (M. Walters, pers. comm.). WEIGHT:

Field Characters. Length 115–13 cm. A dry-country canary of E Africa. Olive eyeline, ear-coverts and malar stripe on yellow face suggest Yellow-fronted Canary S. mozambicus but are more diffuse, less sharp, especially malar stripe. Northern race maculicollis (south to central Kenya) distinctive with white belly, flanks and undertailcoverts, but S birds have white areas washed with yellow, and could be mistaken for Yellow-fronted Canary (Zimmerman et al. 1996); they differ (besides face patch) in longer tail, streaks on flanks (and on breast in X). Voice. Tape-recorded (B, GREG, McVIC, PAY, PEAR). Song both sibilant and wheezy, short phrases, often repeated, ‘chu-weez, chu-weez’, ‘weez-cha, weez, cha’, ‘chercher-weez-cha’, ‘chicher, chicha-weez-cha’; sometimes includes brief rattle. Other songs described as ‘several emphatic sharp notes ending with sharp rising trill’ (Zimmerman et al. 1996) and ‘a sweet series of Goldfinch-like notes or trills, interspersed with some harsher phrases, and quite loud’ (Clement et al. 1993). Buzzy calls, ‘zwee-weez’ and ‘zweess’.

General Habits. Inhabits dry Acacia thorn bush and grassland; often near impenetrable clumps of thorn scrub and often near water (Somalia, Ash and Miskell 1998); in E Africa in bushy and wooded grassland, open woodland and cultivation in low and medium rainfall areas. Solitary, in pairs, or up to 6 birds together. Forages mainly on ground, also in low shrubs. Resident but prone to wander; an irregular visitor to S Kerio Valley, Kenya, in Aug–Dec. Food. Said to be mainly grass seeds. Breeding Habits. Solitary nester. NEST: small, usually very neat, sometimes untidy, delicate open cup, made of lichens, fibres, cobwebs and bents or dry grass and rootlets or twiglets, lined with hair, feathers or plant down and fine grass; one was smooth and red-brown at bottom of cup and black at sides. Int. diam. 45, int. depth 35. Sited generally at 25–35 m above ground on horizontal fork near end of branch in tree, often on plain with isolated or scattered clumps of trees; one in topmost clump of leaves of vertical branch of Boscia coriacea; one only 1 m up in small Maerua cassifolia tree; one 8 m high in thick foliage. EGGS: 1–4, av. (Kenya, 19 clutches) 295. Very pale blue, whitish or greenish blue, usually immaculate, sometimes with sparse reddish brown or blackish spots. SIZE: (n ¼ 8) 165–1925  127–1375 (180  131). LAYING DATES: Ethiopia, Yavello (Yabelo) Mar (and gonads slightly enlarged June, July; imms in June–Aug); Somalia, (young just out of nest Dec and Mar, nestbuilding Nov); E Africa, Region B, Mar–July, Sept, Nov, Jan (mainly Mar–June), Region C, May, June, Region D, Apr–June, Sept, Jan. INCUBATION: by X, sitting tight. Nothing further known. Key References van den Elzen, R. (1985), van den Elzen, R. and Baars, W. (1980).

Plate 29

Serinus flavigula Salvadori. Yellow-throated Seedeater. Serin a ` gorge jaune.

(Opp. p. 458)

Serinus flavigula Salvadori, 1888. Ann. Civ. Mus. Genova, 26, p. 272; Ambokarra, Shoa Prov., Abyssinia. Forms a superspecies with S. xantholaemus. Range and Status. Endemic resident, Shoa Prov., Ethiopia. Map: see Salvadori’s Seedeater S. xantholaemus. 3 specimens taken at Kolla di Aigaber (9
360 N, 40
040 E) in 1880, at Ambokarra (9
310 N, 40
090 E) in 1885 and at Malca Jebdu Shet (i.e. stream near Malca-Ghebdu, 9
340 N, 39
530 E); rediscovered at Malca-Ghebdu at 1400–1500 m in 1989, when parties of 3 then 4 birds seen on one day. The 3 localities are close together near foot of eastern escarpment of W Highlands, Shoa Prov., c. 20 km east of Ankober (see Ankober Serin S. ankoberensis) and enclose an area of only c. 30 km2. 6–7 found at 1310–1610 m on crater rim of Fantalle Mt (9
000 N, 39
540 E) in Awash Nat. Park, c. 65 km south of Malca-Ghebdu, in 1993; one seen in Awash Nat. Park and family at Malca-Ghebdu in 1996, one or

more in Mt Fantalle crater in 1997 and one at Malca Ghebdu in 1998 (Bull. Afr. Bird Club 5, 1998, 144 and 7, 2000, 46). Unconfirmed report of pair at Lalibela (Bull. Afr. Bird Club 3, 1996, 62). Description. ADULT: top of head grey-brown, forehead and crown with fine, short darker streaks, fading on nape and hindneck; mantle darker, warmer brown, with diffuse narrow streaks formed by dark brown feather centres; lower back and rump pale dull yellow (Malca-Ghebdu, Ash and Gullick 1989) or bright canary yellow (Fantalle Mt, Atkins and Harvey 1996), extending to rump sides and short uppertail-coverts; long uppertail-coverts greenish yellow, browner toward tips. Tail feathers dark greyish brown, edged buffy brown. Indistinct whitish superciliary stripes to just behind eye, meeting above bill; lores dark grey-brown;

Serinus flavigula richer and more extensive than on yellow-rumped seedeaters, extending onto uppertail-coverts and down sides of rump and so visible on stationary bird with folded wings. Diffuse pale yellow area on lower throat and upper breast, not obvious except when ruffled by wind (Salvadori’s Seedeater S. xantholaema has conspicuous yellow throat crossed by black bar). In fresh plumage pale green primary edges form panel on wing.

Serinus flavigula, S. xantholaemus

S. flavigula

S. xantholaemus

cheeks, ear-coverts, malar area and sides of neck grey-buff with fine brown streaks. Chin and throat whitish, centre of throat with yellow patch, most intense on lower throat; upper breast and flanks mouse-brown, the former with band of short, faint blackish streaks; lower breast, belly and undertail-coverts paler greybrown, the last washed pale yellow. Upperwing feathers dark greyish brown; inner primaries sometimes narrowly edged greenish, secondaries and tertials buffy brown; greater and median coverts narrowly edged and tipped buffy brown; lesser coverts sometimes with greenish wash. Underwing-coverts and axillaries pale mouse-brown, with suggestion of yellow near alula. Bill pale horn, slightly darker and pinker on culmen; eyes brown; legs pinkish brown. Sexes alike. SIZE: wing, 2 unsexed 66, 675, 2 XX 665, 67; tail, 2 unsexed 505, 49, 2 XX 495, 50; tarsus, 1 unsexed 16. TAXONOMIC NOTE: the taxonomy of S. flavigula (now 4 specimens) and S. xantholaemus (6 specimens) and related serins was discussed in detail by Erard (1974) and Atkins and Harvey (1996). They conclude that the 2 taxa, very similar as they are, are best treated provisionally as separate species, but point out that more field studies and laboratory analyses (e.g. of mtDNA) are needed before a firm conclusion as to relationships can be reached. The isolated Mt Fantalle population is intermediate in character between typical S. flavigula and S. xantholaemus but in C. Erard’s opinion is not a discrete taxon but belongs to S. flavigula (Atkins and Harvey 1996).

Field Characters. Length 10–115 cm. Rare and little known, restricted to central Ethiopia; this account adapted from Atkins and Harvey (1996). Appearance slim and linnet-like, with long wings, quite long tail; lacks the dumpy appearance of Streaky Seedeater S. striolatus or yellow-rumped seedeaters S. xanthopygius and S. reichenowi, superficially resembling a small Yellow-spotted Petronia P. pyrgita. Very plain, with greyish upperparts and pale underparts; most striking feature is bright yellow rump,

Voice. Tape-recorded (109). Canary-like ‘sip’ repeated 2–3 times in flight between food plants; song like call but more musical, ‘zeet ze ze zee’ with lower tone to middle 2 notes (Atkins and Harvey 1996). Chattering food-begging call given by young. General Habits. Malca Ghebdu: in Mar 1989, 3 birds alighted from heights of 20–30 m on bushes on steep, rocky slope by an almost dry rivulet and soon after 4 arrived; area was broken, arid country on rocky hillsides, along valley of a small stream, with scattered acacias and other trees 4–6 m tall, interspersed with patches of thick scrub and bush and a few patches of cereal stubble. Restless and intolerant of approach of observers; sat on outer twigs of trees and bushes and did not seek shade, nor drink at water trickles where there were other serins (Ash and Gullick 1990). Mt Fantalle: in Sept 1993, flock of 6–7 birds flew from top of roadside acacia and settled 80 m away on low Lavandula coronopifolia plants on edge of gully with sparse acacia trees. Not particularly shy but extremely restless, moving from plant to plant in an area of c. 500 m2. In Nov 1993 parties of 1, 2, 3 and 3 birds in same area, feeding on open ground under acacia trees or on long, dry grass clumps on rocky slopes; again very restless, flying readily, almost straight up into trees and often up to 1 km away. In Feb 1994 c. 30 birds on crater rim, some feeding on remnant Lavandula on steep inner slopes of crater, others flying high and strongly on narrow flight path in direction of the only available fresh water, Kassam R., 15 km away. In Sept 1994, 11 seen feeding on Lavandula on inner crater walls; they hardly left the crater, which contained water. Usually in groups of 2–3 but up to 7–11. Forages on ground and in low growth; perches on upper branches of acacias and other trees up to 6 m tall, and on branches of saplings and shrubs growing from rock faces; rarely perches on rocks. ‘Sip, sip’ call given in flight between food-plants. Sings from top of 4 m tall acacia. Food. Seeds, taken on ground (Nov) but taken mainly in situ from tall clumps of dry grass and particularly from Lavandula coronopifolia (Labiatae). Breeding Habits. Bird singing at Malca Ghebdu, early Dec, and on Mt Fantalle, Feb; pair feeding recently fledged young, same place, early Jan (Bull. Afr. Bird Club 2, 1995, 62; 3, 1996, 62; 7, 2000, 46). Nothing further known.

Key References Ash, J.S. and Gullick, T.M. (1990), Atkins, J.D. and Harvey, W.G. (1996), Clement, P. et al. (1993), Erard, C. (1974), Rand, A.L. (1968).

485

486

FRINGILLIDAE

Plate 29

Serinus xantholaemus Salvadori. Salvadori’s Seedeater. Serin de Salvadori.

(Opp. p. 458)

Serinus xantholaema Salvadori, 1896. Ann. Mus. Civ. St. Nat. Giacomo Doria, 2, 16, p. 43; between Wabi Shebelle and L. Stephanie, Abyssinia. Forms a superspecies with S. flavigula. Range and Status. Endemic resident, Ethiopia, map p. 485. Very rare. Known from c. 10 localities in Sidamo, NW Bale and NW Harar Provs (Hall and Moreau 1970, Erard 1974), including Yabelo, Arero, Filtu, Sheikh Hussein, Sof Omar on Weyb R., Karaju on Mane R., Tale Gara Mullasa and Milmil. Best known in Sof Omar, where 1, 2, 4 and 6 birds have been seen in recent years (Ash and Gullick 1989; Bull. Afr. Bird Club 2, 1995, 62 and 7, 2000, 46) and 4–5 pairs, the YY singing so probably breeding, in 1997 (Francis and Shirihai 1999). Description. ADULT: forehead to back and scapulars rather pale greyish brown, top of head with narrow, short darker streaks, mantle, back and scapulars with broader but diffuse grey-brown streaks; rump and short uppertail-coverts lemon yellow; long uppertail-coverts dark grey-brown with blackish central streak, edges tinged yellow. Tail dark grey-brown. Indistinct greyish white superciliary stripes back to eye, meeting above bill. Lores, cheeks and ear-coverts pale greyish brown, bordered below by darker malar area; sides of neck pale greyish brown. Chin white; throat yellow, bordered below by narrow blackish or sooty grey band, and below this a yellow bib on centre of upper breast; sides of breast and flanks pale greyish brown, the latter with a few dusky shaft streaks; lower breast, belly and undertail-coverts greyish white, the last sometimes tinged olive-yellow. Upperwing feathers dark greyish brown, primaries narrowly edged oliveyellow, secondaries and tertials more broadly edged pale buffy brown, greater and median coverts tipped buffy brown to form inconspicuous wing-bars. Underwing-coverts and axillaries pale greyish brown. Bill horn-brown above, pale horn below; eyes brown; legs brown or pale brown. Sexes alike. SIZE: wing, Y (n ¼ 4) 66–695 (674), X (n ¼ 2) 66, 695; tail, Y (n ¼ 4) 45–49 (470), X (n ¼ 1) 485 (Erard 1974); bill (1 X) 125; tarsus (1 X) 135. WEIGHT: Y (n ¼ 2) 12, 12, X (n ¼ 2) 11, 12. TAXONOMIC/NOMENCLATURAL NOTES: S. xantholaemus (including ‘collaris’ and ‘dimidiata’: Erard 1974) was formerly considered to be a hybrid between S. (atrogularis) xanthopygius and either S. mozambicus (Irwin 1960) or S. dorsostriatus (Hall and Moreau 1970), or to be a mutant of S. (a.) reichenowi (Rand 1968).

The type was named in 1896, but a bird described as S. (flaviventris) maculicollis Sharpe in 1895 is in fact S. xantholaemus (J.S. Ash and D. Pearson, pers. comm., 2002); by priority, S. xantholaemus will thus have to be called S. maculicollis. Further study is required. See also Taxonomic Notes under S. dorsostriatus and S. flavigula.

Field Characters. Length 11 cm. A rather plain grey seedeater with bright yellow rump and white underparts with yellow throat and upper breast; distinguished from similar Yellow-throated Seedeater S. flavigula by diagnostic black bar across lower throat, also by white frontal band and pale grey face; their ranges come close in central Ethiopia but they are not known to overlap.

Voice. Tape-recorded (B, GARD, MAC). Song, short phrases of sweet whistles, usually rising in pitch, ‘wayweeo-chip’, ‘way-sicho-wee’, ‘weeo-way-chichi’; phrase frequently repeated, and may be lengthened, ‘way-weeochip-ps-whichowee’, ‘weeo-way-woo-chichi-wee’; pause of several s between phrases. Call, a sharp ‘tsip’.

Habits. Several of the localities are limestone areas. At Sof Omar, inhabits mixed acacia and deciduous woodland along bottom and sides of a steep-sided valley (Francis and Shirihai 1999); this locality, described as a gorge so perhaps a few km away, had been laid almost bare by hundreds of resettled people and their livestock 8 years earlier; nonetheless 6 birds were found (Ash and Gullick 1989). YY sing from tops of tall trees (Sept, presumably in breeding season). Nothing further known. Key References (1974)

Ash, J.S. and Gullick, T.M. (1989), Erard, C.

Subgenus Crithagra Swainson Medium or large finches, YY either yellow/olive (most called canaries) or brown (most called seedeaters), most with sides of head patterned (pale eyebrow, cheek patch, chin and throat, dark band through eye and moustache); bill small, stubby and grey or large, thick and pink-horn (or relatively slender in rufobrunneus); Y canaries plain-breasted, XX greyish and stripe-breasted or Y-plumaged (Serinus flaviventris guillarmodi, S. sulphuratus, S. buchanani); seedeater sexes alike, stripebreasted. Courtship and threat displays by both sexes, with head held high and tail flicking; nest-building by X alone; nest rim does not accumulate crust of nestling faeces; gapes of fledglings red, with blue bill corners (van den Elzen 1985). Endemic. 11 species, 6 independent, 2 superspecies: donaldsoni/buchanani/sulphuratus and striolatus/whytii. We think that flaviventris is close to the sulphuratus superspecies, that melanochrous is closer to striolatus/whytii than to burtoni, and that leucopterus and burtoni may be closely allied.

486

FRINGILLIDAE

Plate 29

Serinus xantholaemus Salvadori. Salvadori’s Seedeater. Serin de Salvadori.

(Opp. p. 458)

Serinus xantholaema Salvadori, 1896. Ann. Mus. Civ. St. Nat. Giacomo Doria, 2, 16, p. 43; between Wabi Shebelle and L. Stephanie, Abyssinia. Forms a superspecies with S. flavigula. Range and Status. Endemic resident, Ethiopia, map p. 485. Very rare. Known from c. 10 localities in Sidamo, NW Bale and NW Harar Provs (Hall and Moreau 1970, Erard 1974), including Yabelo, Arero, Filtu, Sheikh Hussein, Sof Omar on Weyb R., Karaju on Mane R., Tale Gara Mullasa and Milmil. Best known in Sof Omar, where 1, 2, 4 and 6 birds have been seen in recent years (Ash and Gullick 1989; Bull. Afr. Bird Club 2, 1995, 62 and 7, 2000, 46) and 4–5 pairs, the YY singing so probably breeding, in 1997 (Francis and Shirihai 1999). Description. ADULT: forehead to back and scapulars rather pale greyish brown, top of head with narrow, short darker streaks, mantle, back and scapulars with broader but diffuse grey-brown streaks; rump and short uppertail-coverts lemon yellow; long uppertail-coverts dark grey-brown with blackish central streak, edges tinged yellow. Tail dark grey-brown. Indistinct greyish white superciliary stripes back to eye, meeting above bill. Lores, cheeks and ear-coverts pale greyish brown, bordered below by darker malar area; sides of neck pale greyish brown. Chin white; throat yellow, bordered below by narrow blackish or sooty grey band, and below this a yellow bib on centre of upper breast; sides of breast and flanks pale greyish brown, the latter with a few dusky shaft streaks; lower breast, belly and undertail-coverts greyish white, the last sometimes tinged olive-yellow. Upperwing feathers dark greyish brown, primaries narrowly edged oliveyellow, secondaries and tertials more broadly edged pale buffy brown, greater and median coverts tipped buffy brown to form inconspicuous wing-bars. Underwing-coverts and axillaries pale greyish brown. Bill horn-brown above, pale horn below; eyes brown; legs brown or pale brown. Sexes alike. SIZE: wing, Y (n ¼ 4) 66–695 (674), X (n ¼ 2) 66, 695; tail, Y (n ¼ 4) 45–49 (470), X (n ¼ 1) 485 (Erard 1974); bill (1 X) 125; tarsus (1 X) 135. WEIGHT: Y (n ¼ 2) 12, 12, X (n ¼ 2) 11, 12. TAXONOMIC/NOMENCLATURAL NOTES: S. xantholaemus (including ‘collaris’ and ‘dimidiata’: Erard 1974) was formerly considered to be a hybrid between S. (atrogularis) xanthopygius and either S. mozambicus (Irwin 1960) or S. dorsostriatus (Hall and Moreau 1970), or to be a mutant of S. (a.) reichenowi (Rand 1968).

The type was named in 1896, but a bird described as S. (flaviventris) maculicollis Sharpe in 1895 is in fact S. xantholaemus (J.S. Ash and D. Pearson, pers. comm., 2002); by priority, S. xantholaemus will thus have to be called S. maculicollis. Further study is required. See also Taxonomic Notes under S. dorsostriatus and S. flavigula.

Field Characters. Length 11 cm. A rather plain grey seedeater with bright yellow rump and white underparts with yellow throat and upper breast; distinguished from similar Yellow-throated Seedeater S. flavigula by diagnostic black bar across lower throat, also by white frontal band and pale grey face; their ranges come close in central Ethiopia but they are not known to overlap.

Voice. Tape-recorded (B, GARD, MAC). Song, short phrases of sweet whistles, usually rising in pitch, ‘wayweeo-chip’, ‘way-sicho-wee’, ‘weeo-way-chichi’; phrase frequently repeated, and may be lengthened, ‘way-weeochip-ps-whichowee’, ‘weeo-way-woo-chichi-wee’; pause of several s between phrases. Call, a sharp ‘tsip’.

Habits. Several of the localities are limestone areas. At Sof Omar, inhabits mixed acacia and deciduous woodland along bottom and sides of a steep-sided valley (Francis and Shirihai 1999); this locality, described as a gorge so perhaps a few km away, had been laid almost bare by hundreds of resettled people and their livestock 8 years earlier; nonetheless 6 birds were found (Ash and Gullick 1989). YY sing from tops of tall trees (Sept, presumably in breeding season). Nothing further known. Key References (1974)

Ash, J.S. and Gullick, T.M. (1989), Erard, C.

Subgenus Crithagra Swainson Medium or large finches, YY either yellow/olive (most called canaries) or brown (most called seedeaters), most with sides of head patterned (pale eyebrow, cheek patch, chin and throat, dark band through eye and moustache); bill small, stubby and grey or large, thick and pink-horn (or relatively slender in rufobrunneus); Y canaries plain-breasted, XX greyish and stripe-breasted or Y-plumaged (Serinus flaviventris guillarmodi, S. sulphuratus, S. buchanani); seedeater sexes alike, stripebreasted. Courtship and threat displays by both sexes, with head held high and tail flicking; nest-building by X alone; nest rim does not accumulate crust of nestling faeces; gapes of fledglings red, with blue bill corners (van den Elzen 1985). Endemic. 11 species, 6 independent, 2 superspecies: donaldsoni/buchanani/sulphuratus and striolatus/whytii. We think that flaviventris is close to the sulphuratus superspecies, that melanochrous is closer to striolatus/whytii than to burtoni, and that leucopterus and burtoni may be closely allied.

Serinus donaldsoni

487

Serinus donaldsoni Sharpe. Northern Grosbeak-Canary. Serin a ` gros bec.

Plate 29

Serinus donaldsoni Sharpe, 1895. Bull. Br. Orn. Club, 4, p. 41; Sheikh Hussein, Abyssinian Somaliland, fide W.L. Sclater, 1930, Syst. Avium Aethiopicarum, p. 817.

(Opp. p. 458)

Forms a superspecies with S. buchanani and S. sulphuratus. Range and Status. Endemic resident and wanderer, Horn of Africa and Kenya. Ethiopia, known from Awash Nat. Park and Fejeje (Farnsworth et al. 2000); frequent at 1220– 1370 m around Yavello and in headwaters of Webi Shabeelle. Somalia, uncommon; 15 old and 10 recent records as mapped (Ash and Miskell 1998); 7 YY found in 4 days between Garrero (Ain Valley), Nogal Valley and Bohotleh (Archer and Godman 1961); 3 birds separately on 1 day, Botot in NW. Kenya, scarce and local at 200–1600 m, mainly in dry acacia country at 400–1300 m (Britton 1980, referring also to S. buchanani); from Ethiopian border, E side of L. Turkana and Central Island, Marsabit, Ndotos, Nasolot to Wamba and Rumuruti Districts (east to Archers Post) and Baringo; also Garissa in Tana R. valley (Benson 1947); may be only an irregular non-breeding visitor south of c. 1
300 N. Description. ADULT Y: forehead to back and scapulars yellowish olive, streaked dark brown, narrowly on top of head, broadly on mantle, scapulars and back; rump yellowish olive at sides, bright yellow in centre; uppertail-coverts olive-yellow, feathers with blackish central streak. Tail feathers dark brown, outer webs edged pale olive. Prominent superciliary stripe and short stripe below eye bright yellow; lores, ear-coverts and upper cheeks dark olive, tinged yellow; yellow patch on lower cheeks bordered by broad dark olive malar stripe; sides of neck streaked olive-yellow and dark brown. Chin to upper belly and flanks bright golden yellow, sides of breast and flanks with narrow dark brown streaks; lower belly and vent white; undertail-coverts yellow. Remiges dark brown, primaries and secondaries narrowly edged olive-yellow, tertials edged olive-buff and tipped pale yellowish buff; rest of upperwing feathers dark brown, primary coverts and alula finely edged and tipped yellowish, greater coverts broadly edged oliveyellow, and these and median coverts tipped pale yellowish buff to form 2 narrow but distinct wingbars; lesser coverts tipped yellowish olive. Underwing-coverts buffy white with dusky bases; axillaries olive-yellow with dusky centres. Bill large and stout (A), pale brown or flesh-brown above, whitish or pink below with dusky tip; eyes brown; legs brown or sepia. ADULT X: top of head pale tawny-brown, streaked dark brown; mantle to back and scapulars greyish olive-brown streaked dark brown and buffish; rump yellowish olive at sides, yellow in centre; uppertail-coverts olive-yellow. Superciliary stripe and narrow stripe below eye whitish; lores, ear-coverts and upper cheek tawny brown; whitish patch on lower cheek bordered by tawny brown moustachial area. Underparts buffy white, breast and flanks boldly streaked dark brown, sides of chin and throat with a few small dark streaks. Tail, upperwing and bare parts as in ad. Y. Underwing-coverts and axillaries whitish with dusky bases. SIZE (6 YY, 5 XX): wing, Y 81–86 (828), X 75–79 (772); tail, Y 57–67 (620), X 59–61 (598); bill, Y 175–18 (178), X 16–17 (164), max. width (base of mandible) 116; tarsus, Y 17–18 (176), X 165–175 (169). WEIGHT: Ethiopia (unsexed, n ¼ 7) 160–217 (195); Kenya, 1 Y 235, 1 X 233, 1 unsexed 263. IMMATURE: juv. similar to ad. X, but superciliary stripe and underparts buffier; lacks cheek patch; often more extensively streaked below. TAXONOMIC NOTE: see S. buchanani.

Serinus donaldsoni

Field Characters. Length 15–155 cm. Stout, with large head and heavy pale pink bill. Y bright yellow below, with white belly, streaked flanks, heavily streaked dark olivegreen upperparts and contrasting bright yellow rump; further distinguished from Southern Grosbeak Canary S. buchanani by well-marked yellow and olive face pattern and more contrast between rump and upperparts; larger and darker above than Brimstone Canary S. sulphuratus, which has lightly streaked back, green rump, entirely yellow underparts without flank streaks, prominent olive malar stripe. X very different from Y and from XX of other species: grey-brown above, white below with heavily streaked breast; yellow confined to rump.

Voice. Tape-recorded (B, PEAR). Song a dry, monotonous, bunting-like trill at unvarying speed and pitch, lasting c. 3 s, given at intervals of c. 5 s. Another song of sweet yet piercing notes, many up-slurred and given in groups at

488

FRINGILLIDAE

3- or 4-s intervals, ‘sreeeet . . . wriseet . . . sew . . . sreet . . . wreet . . . ’ (Zimmerman et al. 1996, q.v. for further variations). Calls, ‘seep’, ‘pee-chu’ and loud ‘tweea’. General Habits. Inhabits park-like country with scattered thorn trees and scrub on arid plains (S Ethiopia), open acacia bush (Somalia), and open bush, acacia steppe and semi-desert scrub (N Kenya). Solitary or in pairs, once 3 together (Ethiopia) and sometimes in small flocks (Somalia). Usually rather silent; in breeding season Y sings from prominent tree-top perch. Food. Not known.

Breeding Habits. Solitary nester. NEST: several found but none described. EGGS: 2 (3 clutches, Kenya). LAYING DATES: 2 YY had probably bred in July, YY singing Mar, Nov (Benson 1946); Kenya, Mar (1), June (2), July (1), Sept (1). Nothing further known.

Key References Benson, C.W. (1947)., Clement, P. et al. (1993), Farnsworth, S.J. et al. (2000), Zimmerman, D.A. et al. (1996).

Plate 29

Serinus buchanani Hartert. Southern Grosbeak-Canary. Serin de Buchanan.

(Opp. p. 458)

Serinus buchanani Hartert, 1919. Bull. Br. Orn. Club 39, p. 50; Maktan (Voi), British East Africa. Forms a superspecies with S. donaldsoni and S. sulphuratus. Range and Status. Endemic resident and wanderer, S Kenya and NE Tanzania. Kenya, uncommon resident in Olorgesaillie area and southeast to Tsavo East Nat. Park; northeast to Mua Hills. Records on Tana R. in Meru and Garissa may refer to this species or to S. donaldsoni (Benson 1947). Tanzania, uncommon, as mapped: c. 34 records in c. 19 localities (N.E. Baker, pers. comm.); west to Mangola in L. Eyasi depression. Description. ADULT Y: upperparts dull olive-green, top of head narrowly streaked dusky brown, mantle, back and scapulars mottled dark brown, rump and uppertail-coverts brighter, more yellowish, plain apart from a few dusky shaft streaks. Tail feathers dark brown, outer webs narrowly edged olive-green. Superciliary stripe and short stripe below eye pale yellow, narrow and poorly marked. Lores, ear-coverts and cheeks to malar region dusky olive, tinged yellowish green. Underparts olive-yellow, washed dull greyish green on throat and breast, sides of breast and flanks with a few narrow dark brown streaks. Remiges dark brown, primaries and secondaries narrowly edged olive-green, tertials more broadly edged dull olive-green and tipped paler olive-green; primary coverts and alula dark brown, finely fringed olive-green; greater and median coverts dark brown, edged and tipped pale olive-green; lesser coverts olive-green. Underwingcoverts and axillaries pale yellow with dusky bases. Bill large and stout (A), heavier than bill of S. donaldsoni, horn-brown with base of lower mandible whitish; eyes brown; legs dark brown, reddish brown or blackish. ADULT X: similar to ad. Y, ground colour of upperparts slightly duller and browner; underparts slightly duller and paler, breast with more prominent gorget of narrow dark streaks; bill less robust than in Y. SIZE (5 YY, 1X): wing, Y 87–90 (882), X 85; tail, Y 61–67 (628), X 58; bill, Y 18–185 (182), X 18; tarsus, Y 185–20 (190), X 195. IMMATURE: juv. streaked olive-buff and dark brown above, rump greenish olive-yellow; superciliary stripe buffy brown; underparts olive-yellow, chin to breast and flanks narrowly streaked blackish brown. TAXONOMIC NOTE: generally treated formerly as a subspecies of S. donaldsoni; but sexes of donaldsoni are very different whilst those of buchanani are similar, the X being Y-plumaged. S. buchanani is also larger (longer-winged and stouter-billed) than S. donaldsoni.

Serinus buchanani

?

Field Characters. Length 14–15 cm. Large, with heavy pale pink bill. Plainer and lighter green above than Northern Grosbeak Canary S. donaldsoni, rump less contrastingly yellow, no distinct facial pattern, underparts greener,

488

FRINGILLIDAE

3- or 4-s intervals, ‘sreeeet . . . wriseet . . . sew . . . sreet . . . wreet . . . ’ (Zimmerman et al. 1996, q.v. for further variations). Calls, ‘seep’, ‘pee-chu’ and loud ‘tweea’. General Habits. Inhabits park-like country with scattered thorn trees and scrub on arid plains (S Ethiopia), open acacia bush (Somalia), and open bush, acacia steppe and semi-desert scrub (N Kenya). Solitary or in pairs, once 3 together (Ethiopia) and sometimes in small flocks (Somalia). Usually rather silent; in breeding season Y sings from prominent tree-top perch. Food. Not known.

Breeding Habits. Solitary nester. NEST: several found but none described. EGGS: 2 (3 clutches, Kenya). LAYING DATES: 2 YY had probably bred in July, YY singing Mar, Nov (Benson 1946); Kenya, Mar (1), June (2), July (1), Sept (1). Nothing further known.

Key References Benson, C.W. (1947)., Clement, P. et al. (1993), Farnsworth, S.J. et al. (2000), Zimmerman, D.A. et al. (1996).

Plate 29

Serinus buchanani Hartert. Southern Grosbeak-Canary. Serin de Buchanan.

(Opp. p. 458)

Serinus buchanani Hartert, 1919. Bull. Br. Orn. Club 39, p. 50; Maktan (Voi), British East Africa. Forms a superspecies with S. donaldsoni and S. sulphuratus. Range and Status. Endemic resident and wanderer, S Kenya and NE Tanzania. Kenya, uncommon resident in Olorgesaillie area and southeast to Tsavo East Nat. Park; northeast to Mua Hills. Records on Tana R. in Meru and Garissa may refer to this species or to S. donaldsoni (Benson 1947). Tanzania, uncommon, as mapped: c. 34 records in c. 19 localities (N.E. Baker, pers. comm.); west to Mangola in L. Eyasi depression. Description. ADULT Y: upperparts dull olive-green, top of head narrowly streaked dusky brown, mantle, back and scapulars mottled dark brown, rump and uppertail-coverts brighter, more yellowish, plain apart from a few dusky shaft streaks. Tail feathers dark brown, outer webs narrowly edged olive-green. Superciliary stripe and short stripe below eye pale yellow, narrow and poorly marked. Lores, ear-coverts and cheeks to malar region dusky olive, tinged yellowish green. Underparts olive-yellow, washed dull greyish green on throat and breast, sides of breast and flanks with a few narrow dark brown streaks. Remiges dark brown, primaries and secondaries narrowly edged olive-green, tertials more broadly edged dull olive-green and tipped paler olive-green; primary coverts and alula dark brown, finely fringed olive-green; greater and median coverts dark brown, edged and tipped pale olive-green; lesser coverts olive-green. Underwingcoverts and axillaries pale yellow with dusky bases. Bill large and stout (A), heavier than bill of S. donaldsoni, horn-brown with base of lower mandible whitish; eyes brown; legs dark brown, reddish brown or blackish. ADULT X: similar to ad. Y, ground colour of upperparts slightly duller and browner; underparts slightly duller and paler, breast with more prominent gorget of narrow dark streaks; bill less robust than in Y. SIZE (5 YY, 1X): wing, Y 87–90 (882), X 85; tail, Y 61–67 (628), X 58; bill, Y 18–185 (182), X 18; tarsus, Y 185–20 (190), X 195. IMMATURE: juv. streaked olive-buff and dark brown above, rump greenish olive-yellow; superciliary stripe buffy brown; underparts olive-yellow, chin to breast and flanks narrowly streaked blackish brown. TAXONOMIC NOTE: generally treated formerly as a subspecies of S. donaldsoni; but sexes of donaldsoni are very different whilst those of buchanani are similar, the X being Y-plumaged. S. buchanani is also larger (longer-winged and stouter-billed) than S. donaldsoni.

Serinus buchanani

?

Field Characters. Length 14–15 cm. Large, with heavy pale pink bill. Plainer and lighter green above than Northern Grosbeak Canary S. donaldsoni, rump less contrastingly yellow, no distinct facial pattern, underparts greener,

Serinus sulphuratus especially breast, belly yellow, not white. Larger and heavier-billed than Brimstone Canary S. sulphuratus, with brighter rump, indistinct face pattern, no malar stripe, streaks on breast and flanks. Voice. Tape-recorded (B, McVIC, PEAR). Calls, a whistled ‘tyew-oo-lee’ or ‘tyew-ee-yew’, and long-drawn ‘seeeek’. Song ‘a series of low, almost guttural ticking or chipping notes interspersed with much higher-pitched and longer squealing notes of penetrating quality: ‘chrk chrk chrk chrrrrr seeeeeee, tk-tk chrk chrk chrk chrk tsik tsur squeeeeeeeeeeee’ (Zimmerman et al. 1996). General Habits. Inhabits dry thorn scrub and acacia savanna; often in Commiphora thickets. At Mangola, Tanzania, in scattered low thorn bush in denuded

489

grassland. Solitary or in pairs. Forages on ground; also feeds weaver-like on Acacia fruits in situ. Sings from treetops, after rains have promoted green growth. Food. Said to eat grass and weed seeds (Clement et al. 1993). Breeding Habits. Solitary nester. NEST: one was a flattened saucer made of rootlets, twiglets, spider web and vegetable down, built low down in tree. EGGS: pale blue with purple-black dots and a few short lines. SIZE: c. 195  150. Nothing further known. Key Reference Zimmerman, D.A. et al. (1996).

Serinus sulphuratus (Linnaeus). Brimstone Canary; Bully Canary. Serin soufre´.

Plate 29

Loxia sulphurata Linnaeus, 1766. Syst. Nat., ed. 12, 1, p. 305; Cape of Good Hope.

(Opp. p. 458)

Forms a superspecies with S. buchanani and S. donaldsoni. Range and Status. Endemic resident, E African and Rift Valley mountains north to NE Uganda, Angola to N Mozambique, and mainly within 200 km of coast from S Mozambique to SW Cape Province. E Zaı¨re, Lendu Plateau to Kivu (common, at least locally, at 760–2130 m) and Itombwe (very common, 990–2770 m); Marungu highlands (Kasiki). Uganda, up to 2300 m in Impenetrable (Bwindi) Forest and 2400 m in Virunga Volcanoes; W shores of L. Victoria; in belt from L. Albert to W L. Kyoga and Entebbe; Mt Morongole and Mt Elgon. Common in W Rwanda and W Burundi. Kenya, widespread and locally common, seldom over 2400 m, from Mt Elgon, Cheranganis, Maralal, Mt Kenya and Nyambenis to L. Victoria, Mara Game Res., Narok, Nairobi and Machakos Districts (Zimmerman et al. 1996); occasional in Taita Hills. Tanzania, in N, south to Kibondo and Kasulu and from Mara and Serengeti to Loliondo; also on Mt Kilimanjaro. S Tanzania, on Ufipa Plateau and from Mbeya to Iringa, Njombe, Mahenge, Livingstone and Matengo Highlands (common and widespread in Matengo); old records from ˆ i, sparse but wideDar es Salaam and Mikindani. Malaw spread, frequent above 900 m; at all altitudes up to 2400 m on Nyika Plateau but not above 1500 m on Mt Mulanje. S Zaı¨re and Zambia, widespread and locally common, as mapped; absent from Luangwa Valley. Angola, frequent from N Huı´la, Huambo, Cuanza Sul and whole of central Plateau to Bengo, Cuanza Norte, Malanje and Bie´; E Moxico. Mozambique, probably far more widespread north of 20
S than shown; south of 20
S, coast south to Inhaca I. (where frequent) and uncommon in most regions south of 24
S (where probably >50,000 birds: Parker 1999). Botswana, no records (Penry 1994) although old records from extreme NE shown by Hall and Moreau (1970). Zimbabwe and

Serinus sulphuratus

? ?

South Africa, as mapped; in Free State, only in Zastron area in SE, where local (Nuttall 2000). Description. S. s. sharpei Neumann (includes ‘frommi’, ‘shelleyi’ and ‘loveridgei’): Kenya, Uganda and E Zaı¨re to Angola, Zambia, Zimbabwe and central Mozambique. ADULT Y: frontal band bright yellow; rest of upperparts bright olive-green with faint narrow dusky streaks, slightly yellower on uppertail-coverts. Tail feathers blackish brown, outer webs edged bright olive-green. Broad superciliary stripe bright yellow; upper cheek, ear-coverts and sides of neck bright olive-green; lores darker olive-green;

Serinus sulphuratus especially breast, belly yellow, not white. Larger and heavier-billed than Brimstone Canary S. sulphuratus, with brighter rump, indistinct face pattern, no malar stripe, streaks on breast and flanks. Voice. Tape-recorded (B, McVIC, PEAR). Calls, a whistled ‘tyew-oo-lee’ or ‘tyew-ee-yew’, and long-drawn ‘seeeek’. Song ‘a series of low, almost guttural ticking or chipping notes interspersed with much higher-pitched and longer squealing notes of penetrating quality: ‘chrk chrk chrk chrrrrr seeeeeee, tk-tk chrk chrk chrk chrk tsik tsur squeeeeeeeeeeee’ (Zimmerman et al. 1996). General Habits. Inhabits dry thorn scrub and acacia savanna; often in Commiphora thickets. At Mangola, Tanzania, in scattered low thorn bush in denuded

489

grassland. Solitary or in pairs. Forages on ground; also feeds weaver-like on Acacia fruits in situ. Sings from treetops, after rains have promoted green growth. Food. Said to eat grass and weed seeds (Clement et al. 1993). Breeding Habits. Solitary nester. NEST: one was a flattened saucer made of rootlets, twiglets, spider web and vegetable down, built low down in tree. EGGS: pale blue with purple-black dots and a few short lines. SIZE: c. 195  150. Nothing further known. Key Reference Zimmerman, D.A. et al. (1996).

Serinus sulphuratus (Linnaeus). Brimstone Canary; Bully Canary. Serin soufre´.

Plate 29

Loxia sulphurata Linnaeus, 1766. Syst. Nat., ed. 12, 1, p. 305; Cape of Good Hope.

(Opp. p. 458)

Forms a superspecies with S. buchanani and S. donaldsoni. Range and Status. Endemic resident, E African and Rift Valley mountains north to NE Uganda, Angola to N Mozambique, and mainly within 200 km of coast from S Mozambique to SW Cape Province. E Zaı¨re, Lendu Plateau to Kivu (common, at least locally, at 760–2130 m) and Itombwe (very common, 990–2770 m); Marungu highlands (Kasiki). Uganda, up to 2300 m in Impenetrable (Bwindi) Forest and 2400 m in Virunga Volcanoes; W shores of L. Victoria; in belt from L. Albert to W L. Kyoga and Entebbe; Mt Morongole and Mt Elgon. Common in W Rwanda and W Burundi. Kenya, widespread and locally common, seldom over 2400 m, from Mt Elgon, Cheranganis, Maralal, Mt Kenya and Nyambenis to L. Victoria, Mara Game Res., Narok, Nairobi and Machakos Districts (Zimmerman et al. 1996); occasional in Taita Hills. Tanzania, in N, south to Kibondo and Kasulu and from Mara and Serengeti to Loliondo; also on Mt Kilimanjaro. S Tanzania, on Ufipa Plateau and from Mbeya to Iringa, Njombe, Mahenge, Livingstone and Matengo Highlands (common and widespread in Matengo); old records from ˆ i, sparse but wideDar es Salaam and Mikindani. Malaw spread, frequent above 900 m; at all altitudes up to 2400 m on Nyika Plateau but not above 1500 m on Mt Mulanje. S Zaı¨re and Zambia, widespread and locally common, as mapped; absent from Luangwa Valley. Angola, frequent from N Huı´la, Huambo, Cuanza Sul and whole of central Plateau to Bengo, Cuanza Norte, Malanje and Bie´; E Moxico. Mozambique, probably far more widespread north of 20
S than shown; south of 20
S, coast south to Inhaca I. (where frequent) and uncommon in most regions south of 24
S (where probably >50,000 birds: Parker 1999). Botswana, no records (Penry 1994) although old records from extreme NE shown by Hall and Moreau (1970). Zimbabwe and

Serinus sulphuratus

? ?

South Africa, as mapped; in Free State, only in Zastron area in SE, where local (Nuttall 2000). Description. S. s. sharpei Neumann (includes ‘frommi’, ‘shelleyi’ and ‘loveridgei’): Kenya, Uganda and E Zaı¨re to Angola, Zambia, Zimbabwe and central Mozambique. ADULT Y: frontal band bright yellow; rest of upperparts bright olive-green with faint narrow dusky streaks, slightly yellower on uppertail-coverts. Tail feathers blackish brown, outer webs edged bright olive-green. Broad superciliary stripe bright yellow; upper cheek, ear-coverts and sides of neck bright olive-green; lores darker olive-green;

490

FRINGILLIDAE

streaks on breast (X); face pattern distinctive: yellow supercilium, olive lores and ear-coverts, olive malar stripe between yellow cheeks and throat. Larger and heavierbilled than any yellow canary outside E Africa; S birds have varying amount of green on underparts; most like Y Yellow Canary S. flaviventris, which has similar face pattern but much smaller bill, and is bright yellow below.

yellow patch on rear of lower cheek bordered by broad olivegreen malar stripe. Underparts bright yellow, breast and flanks variably washed olive-green, especially on sides; centre of breast often golden yellow. Remiges blackish, primaries narrowly edged yellowish green, secondaries edged olive-green, tertials broadly edged and tipped bright olive-green; primary coverts and alula blackish, finely edged olive-green; greater and median coverts blackish, edged olive-green and broadly tipped yellowish; lesser coverts bright olive-green. Underwing-coverts greyish white, tipped yellow, strongly yellow near carpal edge; axillaries yellow. Bill large and stout, deep at the base (A), edge of maxilla plain or notched (B), dark brown or horn-brown with paler lower mandible; eyes dark brown; legs dark brown. ADULT X: similar to ad. Y, but upperparts slightly duller olive-green; supercilium, cheek patch, chin and throat slightly paler yellow; breast and flanks more strongly washed with green, and yellow of lower underparts less intense. SIZE (10 YY, 10 XX): wing, Y 73–81 (765), X 75–80 (767); tail, Y 54–60 (576), X 50–58 (533); bill, Y 145–155 (150), X 145–155 (152); tarsus, Y 165–19 (178), X ˆ i, Hanmer 2002), wing, Y (n ¼ 84) 70– 17–185 (179). Also (Malaw 78 (744), X (n ¼ 80) 68–78 (733); tail, Y (n ¼ 3) 50–58 (531), X (n ¼ 28) 47–58 (521). WEIGHT: Uganda (n ¼ 22, unsexed) 182– 255 (202); Kenya (n ¼ 35, unsexed) 186–265 (226); Zimbabwe, Y (n ¼ 1) 166, X (n ¼ 3) 174, 175, 218, unsexed (n ¼ 50) 179– ˆ i, Y (n ¼ 98) 154–227 (182), X (n ¼ 89) 150– 239 (211); Malaw 232 (182), imm. (unsexed, n ¼ 52) 152–201 (178) (Hanmer 2002). NESTLING: naked except for long tufts of down on head and back; skin pink-brown. S. s. wilsoni (Roberts) (includes ‘languens’): S Mozambique (south of Save R.) and South Africa (Transvaal south to Free State, KwaZulu-Natal and adjacent E Cape Province). Slightly darker green above. Larger, with heavier bill: wing, Y (n ¼ 6) 80–84 (822), bill, Y (n ¼ 6) 16–17 (164). WEIGHT: (n ¼ 10, unsexed) 22– 302 (258). S. s. sulphuratus (Linnaeus): South Africa (SW and S Cape Prov., intergrading with wilsoni). Much darker olive-green than wilsoni on upperparts and side of head; darker and greener on breast and flanks. Wing, Y (n ¼ 8) 80–85 (824). WEIGHT: (n ¼ 32, unsexed) 25–33 (287).

Field Characters. Length 135–16 cm. Large, with heavy yellow-brown bill. Variable; N birds smaller and plainer than the 2 grosbeak-canaries S. donaldsoni and S. buchanani, which they meet in Kenya and Tanzania; olive-green above, lightly streaked, green rump not contrasting with back; bright yellow below, unstreaked (Y) or with a few

Voice. Tape-recorded (72, 86, 88, 99, 104, B, C, F, LEM). Song rather slower and lower-pitched than other canaries, a mixture of warbles, churrs, liquid twitters and high silvery goldfinch-like notes, usually including characteristic long dry rolling trills; sometimes in 3–5 s phrases, ‘shya-shya-plew-plew-shee-shee-shee-jaja-dididrrrrr’, with short pauses in between, sometimes longer and more continuous. Varies regionally and individually; each bird typically begins his song in the same way but gives it a different ending. Y has quiet subsong of trills and warbles; X also sings, usually when on nest, her song louder and sweeter than Y’s (Skead 1960). Calls, a hard ‘drit’, also ‘‘unmusical, scratchy semi-trills, ‘swirriwirrit’, ‘prr zip’ and ‘chirr-rirr-rip’’’ (Skead 1960); the ‘swirriwirrit’ also used in alarm. X on nest gives deep, rounded whistle, ‘poy’. General Habits. Inhabits moist, open, bushy country with scattered trees, broad-leaved woodlands, savannas and farmland, from sea-level up to nearly 2800 m; small grassy patches alongside streams and marshes, and wooded kloofs, clearings in woods with second-growth shrubs and tufty grass (Katanga); commonest in ecotones between woodland and grassland, e.g. on Inhaca I., Mozambique, occurs amongst halophile vegetation between forest and mangrove, and in cultivated fields (de Boer and Bento 1999); montane grassland with scattered bushes; open gardens, stands of conifers, eucalypts and wattles, edges of cultivation and weedy arable land; in mountains, bracken-briar and scattered bushes (Zimbabwe); favours bush and rank vegetation along streams and around farmland (South Africa). Solitary, in pairs and sometimes flocks of c. 4–12; uncommonly up to 30 birds congregate at rich food source, but not as gregarious as other serins. Mixes with other serins. Forages mainly on ground, hopping and making short, brisk runs, and feeds also in bushes and trees; mandibulates large seeds to crack them open. Rather quiet and unobtrusive, but in breeding season perches conspicuously on fences, bushtops and roofs. Less active than most congeners. Flight fast, only slightly undulating. Resident, but some indication of seasonal movements in Zimbabwe and South Africa (Harrison et al. 1997). Food. Seeds, small fruits and flowers; buds, leaves and shoots; eats more fruits than do congeners. Seeds of garden Aspilia, zinnias, sunflowers and labiates (Kenya); seeds mainly of Olea (43 out of 63 feeding observations: Milewski 1978a), also of Diospyros, Pelargonium, Pinus, Cassytha, Cotoneaster, Psoralea, Raphanus, Acacia, Othonna, Euryops, Chrysanthemoides, Stellaria, Eriocephalus and Chenopodium (SW Cape Prov.); seeds of Digitaria horizontalis, Raphanus rapanistrum, Scutia myrtina, Emex australis, Euphorbia ingens, Lycium, Ligustrum lucidum, Ficus burttdayvi, Ehretia and

Serinus flaviventris

491

Scutia, and in captivity eats seeds of numerous named grasses and herbs, fruits of Rhus dentata, Harpephyllum caffrum, Vaccinium exul, Grewia hexamita, G. villosa etc., leaves of Senecio and buds of Sisymbrium capense (South Africa, Brickell and Konigkramer 1997). Aspilia and sunflowers kernels fed to young (Kenya). Breeding Habits. Solitary nester; monogamous; territorial. Y sings from prominent treetop perch when X attends nest. NEST: deep cup made of twiglets, plant stems, rootlets and some dry grass, often mixed with a little old, heavy cobweb and fibres, warmly and solidly lined with woolly plant down, often yellowish or greyish (C). Can incorporate string, wool and cows’ hair. Cobweb is mainly around rim and where nest is supported on fork; sometimes outside of nest made largely of rootlets, or fine dry grass; some nests have a few small green or dead leaves inside, or felted composite flowers, asclepiad down, and soft shoots of Gnaphalium and Helichrysum (Kenya). Int. diam. c. 50, int. depth c. 40. Placed in fork near top of shrub or wellbranched sapling at 1–6 m (mainly 2–3 m) above ground, e.g. Rhus tree; sometimes in tall conifer and spiny-leaved Hakea tree, well hidden or almost open to view, often in bunch of bananas. Built by X alone. ˆ i, mainly 2–3, av. of 46 EGGS: 3–4 (E Africa), 2–4 (Malaw clutches 27; South Africa, av. of 30 clutches 28). 3 eggs were once laid over 4 day period. Immaculate pure white; or white faintly tinged greenish or bluish, usually with scattered small (sometimes large) speckles and twirls of brown or black at large end. SIZE: (Zaı¨re, Katanga, n ¼ 25) 162–206  126–140 (184  134); (Zimbabwe and South Africa, n ¼ 52) 174–215  131–153 (193  140). LAYING DATES: E Africa, Region A, Oct, Region B, Apr– Jan (21 records evenly distributed), Region C, Jan, Mar– Oct (30 records, mainly Mar and Aug–Sept), Region D, Jan, Apr–June, Oct; Tanzania (Iringa), Mar; Zaı¨re, ˆ i, Katanga, Sept–Oct, once May (Vincent 1949); Malaw Dec–Feb, Apr–Oct, mainly May–Aug; Angola, (breeding condition Aug, fledgling Sept); Zimbabwe, July–Mar, mainly Aug–Oct; South Africa, W Cape July–Nov, E Cape May–Dec, KwaZulu-Natal Oct–Jan. INCUBATION: by X alone. Disturbed bird once gave ‘broken-wing’ distraction display (H.N. Chittenden in

Ginn et al. 1989). Period: 14 days (Kenya) and (southern Africa, n ¼ 6) 125–17, av. 145 days. DEVELOPMENT AND CARE OF YOUNG: quills on head, back and wings start to open on day 6; young well-feathered and fill the nest on day 10. Young brooded by X. She greets incoming Y with quivering wings and open bill and calls ‘suii suiii’; Y feeds her with small portions, by regurgitation; brooding X raises herself off young as she receives each portion from Y, and passes it to a chick. During long absence of Y from nest, X herself regurgitates a little food for the young (van Someren 1956). Later, Y and X feed brood separately. Transfer of food is always careful and gentle. At first X eats chicks’ faecal sacs; later, they are removed; later still, young excrete over side of nest. Young bird preens and eats its dry feather sheaths. X shelters brood from strong sun. Nestling period: 15 days (Kenya), 15 and 21 days (southern Africa). Young out of nest cared for by both parents. BREEDING SUCCESS/SURVIVAL: est. survival rate of birds >6 months old, 52% per year; 2 XX, ringed as ads, recaptured >7 years later (Hanmer 2002). Key References Brickell, N. and Konigkramer, T. (1997), Clancey, P.A. (1972), Clement, P. et al. (1993), Ginn, P.J. et al. (1989), Hanmer, D.B. (2002), Harrison, J.A. et al. (1997), Skead, C.J. (1960), Maclean, G.L. (1993), van Someren, G.V.L. (1956), Vincent, A.W. (1949).

Serinus flaviventris (Swainson). Yellow Canary. Serin de Sainte-He´le `ne.

Plate 29

Crithagra flaviventris Swainson, 1828. Zool. J., 3, p. 348; based on ‘Yellow-bellied gr[osbeak]’ of Latham, 1783, Gen. Synopsis Birds, 2, 1, p. 138, Cape of Good Hope.

(Opp. p. 458)

Range and Status. Endemic resident and wanderer, southern Africa. Introduced into Ascension I. and St Helena I. Angola, uncommon or rare, from Cuando Cubango to S Huı´la and coastal plain of Namibe (Dean 2000). Zambia, specimen at Sinjembele in SW Barotse Prov. and possibly 1 in Mwinilunga (Dowsett and Dowsett-Lemaire 1979). Zimbabwe, old record from Matetsi. Namibia, Botswana and South Africa as mapped (Harrison et al. 1997). Namibia, rare to uncommon in Cunene and Cubango R. valleys and

along margins of Namib Desert, frequent southeastwards towards Botswana and N Cape Prov. borders. Botswana, scarce in N and E but frequent to very common where shown south of 21
S – the common canary of the Kalahari savannas. South Africa, common and widespread in semiarid country, almost throughout, east to about 26
E; abundant in some coastal regions; in Transvaal common in SW Highveld and Kalahari thornveld areas, east to about Ogies, Bethal and Standerton; frequent on Pietersburg Plateau; records from Wakkerstroom and Vivo; throughout

Serinus flaviventris

491

Scutia, and in captivity eats seeds of numerous named grasses and herbs, fruits of Rhus dentata, Harpephyllum caffrum, Vaccinium exul, Grewia hexamita, G. villosa etc., leaves of Senecio and buds of Sisymbrium capense (South Africa, Brickell and Konigkramer 1997). Aspilia and sunflowers kernels fed to young (Kenya). Breeding Habits. Solitary nester; monogamous; territorial. Y sings from prominent treetop perch when X attends nest. NEST: deep cup made of twiglets, plant stems, rootlets and some dry grass, often mixed with a little old, heavy cobweb and fibres, warmly and solidly lined with woolly plant down, often yellowish or greyish (C). Can incorporate string, wool and cows’ hair. Cobweb is mainly around rim and where nest is supported on fork; sometimes outside of nest made largely of rootlets, or fine dry grass; some nests have a few small green or dead leaves inside, or felted composite flowers, asclepiad down, and soft shoots of Gnaphalium and Helichrysum (Kenya). Int. diam. c. 50, int. depth c. 40. Placed in fork near top of shrub or wellbranched sapling at 1–6 m (mainly 2–3 m) above ground, e.g. Rhus tree; sometimes in tall conifer and spiny-leaved Hakea tree, well hidden or almost open to view, often in bunch of bananas. Built by X alone. ˆ i, mainly 2–3, av. of 46 EGGS: 3–4 (E Africa), 2–4 (Malaw clutches 27; South Africa, av. of 30 clutches 28). 3 eggs were once laid over 4 day period. Immaculate pure white; or white faintly tinged greenish or bluish, usually with scattered small (sometimes large) speckles and twirls of brown or black at large end. SIZE: (Zaı¨re, Katanga, n ¼ 25) 162–206  126–140 (184  134); (Zimbabwe and South Africa, n ¼ 52) 174–215  131–153 (193  140). LAYING DATES: E Africa, Region A, Oct, Region B, Apr– Jan (21 records evenly distributed), Region C, Jan, Mar– Oct (30 records, mainly Mar and Aug–Sept), Region D, Jan, Apr–June, Oct; Tanzania (Iringa), Mar; Zaı¨re, ˆ i, Katanga, Sept–Oct, once May (Vincent 1949); Malaw Dec–Feb, Apr–Oct, mainly May–Aug; Angola, (breeding condition Aug, fledgling Sept); Zimbabwe, July–Mar, mainly Aug–Oct; South Africa, W Cape July–Nov, E Cape May–Dec, KwaZulu-Natal Oct–Jan. INCUBATION: by X alone. Disturbed bird once gave ‘broken-wing’ distraction display (H.N. Chittenden in

Ginn et al. 1989). Period: 14 days (Kenya) and (southern Africa, n ¼ 6) 125–17, av. 145 days. DEVELOPMENT AND CARE OF YOUNG: quills on head, back and wings start to open on day 6; young well-feathered and fill the nest on day 10. Young brooded by X. She greets incoming Y with quivering wings and open bill and calls ‘suii suiii’; Y feeds her with small portions, by regurgitation; brooding X raises herself off young as she receives each portion from Y, and passes it to a chick. During long absence of Y from nest, X herself regurgitates a little food for the young (van Someren 1956). Later, Y and X feed brood separately. Transfer of food is always careful and gentle. At first X eats chicks’ faecal sacs; later, they are removed; later still, young excrete over side of nest. Young bird preens and eats its dry feather sheaths. X shelters brood from strong sun. Nestling period: 15 days (Kenya), 15 and 21 days (southern Africa). Young out of nest cared for by both parents. BREEDING SUCCESS/SURVIVAL: est. survival rate of birds >6 months old, 52% per year; 2 XX, ringed as ads, recaptured >7 years later (Hanmer 2002). Key References Brickell, N. and Konigkramer, T. (1997), Clancey, P.A. (1972), Clement, P. et al. (1993), Ginn, P.J. et al. (1989), Hanmer, D.B. (2002), Harrison, J.A. et al. (1997), Skead, C.J. (1960), Maclean, G.L. (1993), van Someren, G.V.L. (1956), Vincent, A.W. (1949).

Serinus flaviventris (Swainson). Yellow Canary. Serin de Sainte-He´le `ne.

Plate 29

Crithagra flaviventris Swainson, 1828. Zool. J., 3, p. 348; based on ‘Yellow-bellied gr[osbeak]’ of Latham, 1783, Gen. Synopsis Birds, 2, 1, p. 138, Cape of Good Hope.

(Opp. p. 458)

Range and Status. Endemic resident and wanderer, southern Africa. Introduced into Ascension I. and St Helena I. Angola, uncommon or rare, from Cuando Cubango to S Huı´la and coastal plain of Namibe (Dean 2000). Zambia, specimen at Sinjembele in SW Barotse Prov. and possibly 1 in Mwinilunga (Dowsett and Dowsett-Lemaire 1979). Zimbabwe, old record from Matetsi. Namibia, Botswana and South Africa as mapped (Harrison et al. 1997). Namibia, rare to uncommon in Cunene and Cubango R. valleys and

along margins of Namib Desert, frequent southeastwards towards Botswana and N Cape Prov. borders. Botswana, scarce in N and E but frequent to very common where shown south of 21
S – the common canary of the Kalahari savannas. South Africa, common and widespread in semiarid country, almost throughout, east to about 26
E; abundant in some coastal regions; in Transvaal common in SW Highveld and Kalahari thornveld areas, east to about Ogies, Bethal and Standerton; frequent on Pietersburg Plateau; records from Wakkerstroom and Vivo; throughout

492

FRINGILLIDAE

Serinus flaviventris

Lesotho; in E Cape Prov. east to about Grahamstown, Stutterheim and border with Griqualand East; in KwaZulu-Natal occurs only marginally along Lesotho border; once at Cathedral Peak. Densities of 17 birds in 40 ha in S Karoo, Cape Prov. (J.M. Winterbottom in Ginn et al. 1989), and 74 pairs along c. 63 km of Modder R. (av. 11.4 pairs per km) (Kopij 2000). In Mar–Aug up to 26 non-breeding birds winter on Bird I. (Lambert’s Bay, W Cape Prov.) in probably 40,000 birds (Parker 1999); Transvaal, common in hilly areas of Escarpment and Bushveld regions and in Witwatersrand, Suikerbosrand and wooded parts of central highveld, but scarce elsewhere (Tarboton et al. 1987); Free State, locally common in Sasolburg area in N and in mountains in E; KwaZulu-Natal and E Cape Prov., frequent and widespread; SW Cape Prov., has undergone large westerly expansion in last 35 years but still uncommon: a decade ago no nearer to SW coast (Cape St Martin to Cape Agulhas) than 20–40 km (Hockey et al. 1989) but now seen there not infrequently (Harrison et al. 1997). Resident, with some evidence locally for seasonal and altitudinal movements and nomadism (Harrison et al. 1997). Description. S. g. canicapilla (Dubus): Guinea to Central African Republic. ADULT Y: forehead to nape streaked buffish white and dark brown; hindneck, sides of neck, mantle and scapulars to uppertail-coverts plain greyish brown or earth brown; tail dark

Serinus gularis

? ?

?

?

??

brown. Broad supercilium buffy white, extending back to side of nape. Lores, ear-coverts and cheeks plain dark brown. Chin and throat pale greyish buff, chin with small dusky spots or short streaks; breast and flanks greyish brown, grading to pale greyish buff on belly and undertail-coverts. Upperwing feathers dark brown, flight feathers finely edged pale buff, tertials edged buffy brown and tipped buff, greater coverts broadly edged buffy brown and tipped paler brown, median coverts tipped pale brown; wingbars inconspicuous, posterior one most prominent. Underwingcoverts and axillaries pale greyish brown. Bill horn brown, lower mandible whitish; eyes light brown; legs horn brown to flesh. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 77–85 (800), X 71–82 (776); tail, Y 50–55 (528), X 49–54 (520); bill, Y 13–145 (137), X 125–14 (133); tarsus, Y 14–155 (146), X 14–15 (146). IMMATURE: juv. paler above than ad. with distinct diffuse streaking, and spotted below across breast and on flanks; tertials, greater and median coverts with more prominent pale buff tips. NESTLING (S. g. gularis): hatches with skin pinkish red, pale grey down on crown, superciliaries, occiput, humerals, back, rump, secondaries, flanks and crurals; minute yellow filaments on tail and primary tracts; bill yellow, gape edges red, mouth and tongue red, legs pale pink (Skead 1960). S. g. montanorum Bannerman: highlands of E Nigeria and W Cameroon; at 1200–2000 m. Like canicapilla but slightly darker brown above, darker and browner below. S. g. elgonensis (Ogilvie-Grant): Sudan to W Kenya. Like canicapilla but slightly paler greyish brown above. S. g. benguellensis (Reichenow): Angola to W Zambia. Darker, richer brown above than canicapilla, pale crown streaks browner; darker and browner below, chin streaks darker.

Serinus gularis S. g. mendosus Clancey: NE Botswana, Zimbabwe and adjacent highland Mozambique. Upperparts paler and greyer than in canicapilla and faintly mottled; lacks streaks on chin; white chin and throat contrast with pale grey-brown breast and flanks; belly greyish white. Larger billed, with longer tail and much longer tarsus: SIZE (1 Y, 4 XX): wing, Y 79, X 77–79 (788), tail, Y 62, X 59–61 (600), bill, Y 15, X 145–15 (146); tarsus, Y 17, X 17–175 (171). WEIGHT: Y (n ¼ 3) 143–168 (152), X (n ¼ 5) 105–195 (159), unsexed (n ¼ 172) 114–21 (160). S. g. gularis (Smith): N and W Transvaal and adjacent Botswana to W Free State and N Cape Prov. (including Griqualand West). Like mendosus, but rather darker and browner (less grey) on breast and flanks, slightly browner above. Slightly paler above than canicapilla with whiter throat, few or no chin streaks, and distinctly mottled upperparts. WEIGHT: (Transvaal races, unsexed, n ¼ 50) 17–25 (201). S. g. endemion (Clancey): SE Transvaal, Swaziland and S Mozambique (Lebombo Range), through E Free State, Lesotho and Natal to Cape Prov. (east of Great Fish R.). Like gularis but slightly darker and greyer, with dark-spotted chin. SIZE (6 YY): wing, 75–80 (777), tail 57–64 (608), bill 15–165 (157), tarsus 16–175 (169). S. g. humilis (Bonaparte): SW and S Cape Province (east to Great Fish R.). Darker grey-brown above than endemion, more strongly mottled, pale head streaks buffy brown (not whitish); chin and throat grey-buff, the former well-mottled. Bill still larger: Y (n ¼ 3) 16–17 (165), unsexed (n ¼ 7) 16–165 (163). WEIGHT: SW Cape, unsexed (n ¼ 112) 167–248 (203) (Ward 2002b).

Field Characters. Length 13–16 cm. A wide-ranging seedeater, plain-looking except for long white superciliary stripe; upperparts lightly streaked, pale streaks on top of head, solid brown face patch; white throat contrasts with plain brown breast, chin with dark mottling in some races. Differs from all similar seedeaters except Brown-rumped S. tristriatus (which it does not overlap) by unstreaked underparts; from Streaky S. striolatus by unbroken face patch without separate malar stripe; from Black-eared S. mennelli by brown face patch not sharply defined with white below; very like Reichard’s Seedeater S. reichardi, but latter is streaked below. Voice. Tape-recorded (88, 99, 104, B, C, F, WALK). One of the finest singers among the canaries. Song varies regionally: bird in Nigeria (104) gave powerful song lasting over 30 s, a remarkable variety of sweet liquid trills, buzzes and rattles, at different speeds and pitches, reminiscent of domestic canary S. canaria; in southern Africa songs medium-paced, with many clear sweet whistles and some hard rattles, usually shorter (3–4 s); same phrase may be repeated, and may also be continued into longer (up to 17 s) warbling. Calls include brief ‘tssrr’ or ‘tsr-tsip’, hard ‘jip’, longer ‘tsee-sweep’ and a rolled, nasal ‘zoo-weeyoweeyip’. For further renditions see Skead (1960) and Zimmerman et al. (1996). General Habits. Inhabits well-wooded savanna, forest clearings, millet fields (Ghana, Togo); degraded guineansavanna and farmland, relatively undisturbed dry woodland with Anogeissus schimperi trees, near sandy streams with well-timbered banks and open, grazed glades between scattered clumps of shrubs (N Nigeria, Fry 1965), orchardbush savanna and small farm patches and scattered large shade trees (S Nigeria, NE Zaı¨re); light woodland savanna,

rough, open grassland dotted with clumps of bushes and copses of deciduous trees dominated by Brachystegia randii, Berlinia globiflora and Muhatja, also open hillsides with scattered trees and bushes, mountains, edges of Acacia and Terminalia woodland savanna, secondary growth, cultivation and gardens (Zimbabwe, Vincent 1949, Irwin 1981); visits garden bird tables; deciduous woodland, savanna, edges of forest and plantations, wooded valleys and kloofs, and stands of Protea caffra in hill country (Transvaal); scrub on mountain slopes, open bush, edges of evergreen forest, and thornveld (KwaZulu-Natal). Occurs usually singly, in pairs or family parties; sometimes in flocks of about 8; larger flocks (20–50, and once 95 birds) can gather at flowering plants, e.g. aloes. Occasionally feeds with other serins and seedeaters, but less gregarious than most congeners; joins mixed-species foraging flocks in Zimbabwe Brachystegia woods. Feeds on ground but mainly in seeding heads of herbs, grasses and cereals, e.g. Sorghum and bulrush millet Pennisetum americanum (A), and in bushes, aloes and trees. Bird on ground jumps up at head of grass or herb and pecks seeds off, or bends the plant and holds it down with one foot whilst taking seeds from its head (W.R.J. Dean in Ginn et al. 1989). Rolls a large seed in bill, mandibulating it so as to remove seedcoat and swallow kernel. Enters mealie (maize) store and scrapes maize grains with bill. Bird perched in mulberry Morus bush plucks fruit from above its head, transfers it to a foot, holds it between foot and twig perch, repositions it using bill if it slips, and nibbles at it, eating it in small portions (Milstein 1983). Probes base of flowers to extract nectar; nibbles stamens, anthers and petals; tears open unopened Aloe flowers, eats anthers but ignores stamens. Plucks whole Tecomaria flower, splits calyx and bottom of corolla tube with bill, sucks nectar out then drops the flower; or tears corolla tube of flower in situ down its whole length and then sucks the nectar (Skead 1960). Flies out to take winged termites and hoverflies; once 8 birds hawked termites together. Quiet and unobtrusive; even when Y calls from canopy or treetop in breeding season it attracts little attention (W.R.J. Dean in Ginn et al. 1989). Gait on ground a short hop; flight strongly undulating, leisurely; does not fly high. Often raises crown feathers when perched, not only when alarmed. Flies to take cover inside bush when disturbed.

507

508

FRINGILLIDAE

Food. Seeds, small fruits, nectar, buds, flower parts and insects. Very fond of aloe stamens (South Africa). Seeds only, in 20 stomachs (Nigeria), fruits of Anogeissus schimperi (Nigeria); many seeds, 12 termites and 4 small caterpillars (5 stomachs, Zaı¨re). Caterpillars sometimes fed to nestlings. In southern Africa, seeds of Protea roupelliae, Celtis africana, Acacia karoo, Pennisetum glaucum, Carica papaya, Hyparrhenia hirta, Nicotiana tabacum, pines, Casuarina, Opuntia, sunflower and Sorghum; feeds at Maytenus, Aspalanthus and Chenopodium (together 64% of 54 feeding occasions), Aloe, Nymania, Chirona, Lampranthus, Melianthus, Salsola, Eriocephalus and Olea plants (Milewski 1978a); stamens of Aloe, peach and apricot flowers; nectar of Kniphofia, Aloe marlothii, A. candelabrum, A. arborescens, A. ferox, A. humilis, A. variegata, Tecomaria capensis and Hibiscus; fruits of Morus, Euphorbia ingens and wild and garden Lantana and figs; buds and petals of Prunus persica, P. armeniaca, Salvia africana, Erythrina, Amaranthus hybridus, Taretes erecta, T. patula, Rhus pyroides, Morus mesozygia, Lycium afrum, L. austrinum, Lantana rugosa and Ficus (Brickell and Konigkramer 1997). Breeding Habits. Solitary nester; presumably monogamous; perhaps mildly colonial – 2 or probably several pairs may nest close together; little evidence of territorial animosity (Skead 1960). Y sings from perch often high up, e.g. top of tall Casuarina tree, also from bush or telephone pole or within woodland canopy, and sometimes sings in circling flight. Y sings in ‘butterfly’ flight with slow, regular wingbeats, but ‘butterflying’ less common than in most congeners. Often a bird leaves a party of 3–4, flies up and ‘rows’ wings for up to 200 m, singing all the time. 2 pairs were once on adjacent bushes; both YY sang and from time to time one or both postured at its X by pointing bill skyward, swaying head slowly and sidling up to X (Skead 1960), with body held horizontally and slightly raised and fanned tail (W.R.J. Dean in Ginn et al. 1989). Once Y sang at perch, another flew in and perched beside it; singer immediately fluffed its feathers out and slowly swayed from side to side whilst uttering throaty chirrups (Skead 1960). 2 flying, singing birds often flutter down, spiralling, bills almost touching. NEST: a small, neat cup made in 3 parts (base, lining, inner lining) with soft, dry petioles bound with cobweb, around soft, fibrous, dry grass inside, smoothly and compactly lined with vegetable down, often bright reddish (Zimbabwe); firmly attached to supporting twigs by cobweb (B). Also made of grass, dead leaves, small dead twiglets up to 50 mm long and scraps of bark and paper, all firmly bound with coarse cobweb inside and out, lined with very thin slivers of bark, with innermost lining c. 4 mm thick of wool and vegetable down (e.g. of Protea). Occasionally nests are bulky. One contained guineafowl feathers; another lined with kapok stolen from nest of Cape Penduline Tit Anthoscopus minutus. SIZE: ext. diam. 64–105, int. diam. 51–75, ext. depth up to 51, int. depth 26– 39. Sited 15–12 (usually 2–3) m up in shrub or quite large tree (e.g. Parinarium mobola), on twiggy, multiple forks in middle of shrub near top, or slung between horizontal forks, or in vertical fork, or near end of a lower branch of tall tree; often on cluster of pine cones or seed pods; once behind peeling bark, once on old nest of Rock Martin

Hirundo fuligula; usually well screened by foliage. Nest built by X only, Y accompanying her and singing. EGGS: 2–4, usually 2–3, av. of 33 clutches (Zimbabwe) 23. Laid (and hatch) on successive days. Shell smooth, sometimes rough. White or very pale greenish blue, speckled with chocolate brown, mauve-brown and pale violet, mainly in a zone around broad end; sometimes with a few blackish hairstreaks. SIZE: (n ¼ 52) 170–212  126–150 (185  138). LAYING DATES: Nigeria, (breeding condition, July; territorial YY singing, June–Aug); Angola, (juv., Jan); Zimbabwe, Sept–Mar, mainly Sept–Jan (98 out of 111 records); Transvaal, Oct–Jan; KwaZulu-Natal, Sept–Feb. INCUBATION: by X only, starting with 3rd egg. X fed on nest by Y, X lifting head to receive food piecemeal but making no sound; Y feeds her by regurgitation, up to 14 times. Having been fed, X keeps food and continues to incubate or brood, or follows Y away, or she often feeds the brood, also piecemeal. Period: 125–15 days, av. 133. DEVELOPMENT AND CARE OF YOUNG: see NESTLING above; on day 2 skin darker; day 4, bill pale horn with yellow cutting edges, primaries, secondaries and their coverts emerging; day 5, quills on back, breast, flanks and belly, eyes opening; day 6, quills on head and tail; day 7, quills on chin and cloaca, feathers on secondary coverts, back, humerals, breast, flanks and belly; day 8, feathers on crown and tail; day 9, eyes fully open (Skead 1960). Young brooded by X for c. 5 days. Later, each parent feeds chicks, directly, with mixture of seeds and some caterpillars and grit. Parents swallow chicks’ faecal sacs for first 8 days; thereafter, chicks deposit faeces on nest rim. Nestling period: c. 17 days. Out of nest, chicks cared for by Y and X who give each young 6–10 billfulls of food per feed. Hungry young calls intermittently but chicks do not yammer perpetually; they solicit from incoming parent with wings quivering. BREEDING SUCCESS/SURVIVAL: no information. Key References Brickell, N. and Konigkramer, T. (1997), Ginn, P.J. et al. (1989), Harrison, J.A. et al. (1997), Irwin, M.P.S. (1977, 1981), Maclean, G.L. (1993), Milstein, P. le S. (1983), Skead, C.J. (1960), Vincent, A.W. (1949), Ward, V.L. (2002b).

Serinus reichardi

509

Serinus reichardi (Reichenow). Reichard’s Seedeater; Stripe-breasted Seedeater. Serin de Plate 30 Reichard. (Opp. p. 459) Poliospiza reichardi Reichenow, 1882. J. Orn., 30, p. 209; Kakoma, Tabora, Tanganyika. Forms a superspecies with S. gularis. Range and Status. Endemic resident, NE Ethiopia to NE Namibia. Ethiopia, locally common in W Highlands, and in SE Highlands where range closely approaches or even overlaps with that of similar-looking Brown-rumped Seedeater S. tristriatus; records in and just south of Nechisar Nat. Park (Safford et al. 1993). Sudan, uncommon, near borders in S and SE (Yambio, Yei, Didinga, Boma; Nov–May). Kenya, uncommon and local in highlands of W and centre at 1600– 1800 m, sometimes up to 2000 m, from N and S slopes of Mt Elgon (Timau, Mangiki), Kongelai Escarpment and Muhoroni (near Kisumu) east to Mt Ololokwe, from Mt Nyiru south to Limuru, and within that compass occurring on Elgeyu, Tambach and Laikipia Escarpments and Tugen Hills and at Kacheliba, Kabarnet, near Marigat, Eldama Ravine and L. Nakuru. Burundi, pair once at Kiofi (Giofi), Kumoso (Gaugris et al. 1981). Tanzania, c. 60 records, locally common at 500–1500 m in Iringa Highlands between Malangali and Iringa (e.g. Ngwazi, 1830 m, Moyer et al. 1990), elsewhere sparse from Kibondo, central Tabora Prov., NW Rukwa Prov. and Ufipa Plateau to Mbeya, Songea, Matengo, Masasi and east of Rufiji R. near Monga; flock of >20 at Tatanda (Ufipa); frequent to common in Matengo Highlands (Sassi and Zimmer 1941); probably widespread at low densities in much of W and SE Tanzania (N.E. Baker, pers. comm.). Zaı¨re, uncommon, north to 10
S in SW and S of upper Katanga, record from near Moba on L. Tanganyika. Zambia, occurs in Brachystegia and Baikiaea woodland, range overlapping broadly that of Black-eared Seedeater S. mennelli, but S. reichardi less common than it; known from 41% of Zambia squares, as mapped (R.J. Dowsett, pers. comm.); absent from Luangwa and Middle Zambezi Valleys and from west of Zambezi in Balovale. Zimbabwe, record from Chambonda ˆ i, uncommon, at 900– Vlei, Victoria Falls, Mar 1995. Malaw 2000 m, from Nyika Plateau and Misuku Hills to Lilongwe (Dzalanyama Mts), Dedza and Mangochi (Namwera), south to about 16
S (Hall and Moreau 1970). Mozambique, Furancungo, Tete Prov.; record from W Nampula Province. Description. S. r. reichardi (Reichenow): south of L. Victoria. ADULT Y: forehead to hindneck streaked buffy white and dark brown; mantle, back and scapulars rich earth brown, diffusely streaked darker brown; rump and uppertail-coverts plain earth brown; tail dark brown. Broad superciliary stripe whitish, extending back to side of nape; lores, cheeks and ear-coverts dark brown. Chin and throat white with faint buff wash; breast and flanks grey-buff, suffused darker grey-brown on sides and flanks, with band of dark brown streaks across upper breast and more diffuse streaks on lower breast and flanks; belly and undertailcoverts buffish white. Upperwing feathers dark brown, remiges narrowly edged buffy brown when fresh, tertials tipped grey-buff, greater coverts edged buffy brown and tipped grey-buff, forming a pale narrow wing-bar, median coverts tipped grey-buff, forming a less conspicuous wing-bar. Underwing-coverts and axillaries pale grey-buff. Bill slate; eyes brown; legs black. Sexes alike. SIZE (10 YY, 10 XX): wing, Y 76–81 (790), X 76–83 (784); tail, Y 48–56

Serinus reichardi

(537), X 49–54 (509); bill, Y 12–135 (128), X 115–135 (128); tarsus, Y 15–16 (154), X 15–155 (153). IMMATURE: juv. paler, more tawny above than ad., with streaking more pronounced; breast and flanks with short tawny-brown streaks; tertials edged and tipped tawny-buff; well-demarcated tawny-buff tips to greater and median coverts, forming distinct wing-bars. S. g. striatipectus (Sharpe): Ethiopia to S Sudan and Kenya. Sharp streaking on underparts extends to chin, throat, lower breast and flanks. Slightly tawnier above than nominate race; washed browner, less greyish, on breast and flanks. WEIGHT: (Ethiopia, n ¼ 2, unsexed) 143, 150 (NW Kenya, n ¼ 16, unsexed) 19–22 (205). TAXONOMIC NOTE: the form striatipectus has been attributed to S. gularis although it is sympatric with S. g. elgonensis on Mt Elgon and Kongolai Escarpment (Kenya) and does not respond to playback songs of Southern African S. gularis (Zimmerman et al. 1996). The taxonomic picture is obscured by considerable variation of these serins as well as by their scarcity.

Field Characters. Length 125–13 cm. A brown seedeater with long white supercilium, streaked above and below but less boldly than Streaky Seedeater S. striolatus; face patch solid brown, without separate malar stripe. Distinguished from Streaky-headed and Brown-rumped Seedeaters S. gularis and S. tristriatus by streaked underparts, from Black-eared Seedeater S. mennelli by brown, not black, face patch, not clearly demarcated below with white.

510

FRINGILLIDAE

Voice. Tape-recorded (102, B, LEM, McVIC). Song a mixture of sweet and buzzy trills interspersed with unmusical twittering and much repetition of single notes and phrases, e.g. ‘djee-djee-djee-djee-tchueee-tchueeetchueee-tititititit-tsitsi-tsuee-tsuee-tsuitit-tsuiti’, often mingled with extensive imitations of other bird species (Zimmerman et al. 1996, q.v. for further variations). Calls include nasal ‘zee-zwee’ and ‘zu-weeo’; trilled ‘chirry’ or ‘chirru’ and sibilant ‘tissy’. General Habits. In Kenya striatipectus inhabits shrubby escarpments with scattered trees; further south, nominate reichardi is restricted to little-disturbed Brachystegia and Baikiaea woodlands; in Matengo Highlands, S Tanzania, in Faurea-Protea-Uapaka-Brachystegia-dominated woods; once in a mealie garden (Mozambique, Vincent 1936). Habits of both races are poorly known. In Kenya, shy and inconspicuous except when Y sings from a treetop or in courtship flight; territories may be large, since Y sings over a considerable area (Zimmerman et al. 1996). Occurs in pairs or flocks of 3–5 or 10–20; forages on ground; freely uses trees, and flock once seen flycatching from treetops. ‘A common upland species of gardens, cultivated land or

Brachystegia woodland, congregating into large flocks and doing damage to both corn and soft fruit at times’ (Mackworth-Praed and Grant 1960) may refer to S. gularis rather than S. reichardi. Food. No information. Breeding Habits. Solitary nester, or breeds in colonies of 4–5 pairs (Lynes 1934); territorial (see above). Y courts using ‘butterflying’ display flights, singing all the while. NEST and EGGS: ‘Nest of rootlets and stems lined with soft material usually at some height in a tree . . . Eggs 2–4, faint bluish white with a few dots and streaks of purplish black; c. 20  15’ (Mackworth-Praed and Grant 1960) may refer to S. gularis rather than S. reichardi. No subsequent descriptions known. LAYING DATES: E Africa, Region A, July, Region C, Jan– Mar; Zaı¨re, Lubumbashi, (gonads enlarged until Aug); Zambia, (gonads active Feb, Mar, Aug); Zaı¨re, (breeding starts in Jan). Key References Zimmerman, D.A. et al. (1998), Lynes, H. (1934, 1938), Vincent, J. (1936).

Plate 30

Serinus tristriatus Ru ¨ ppel. Brown-rumped Seedeater. Serin `a trois raies.

(Opp. p. 459)

Serinus tristriatus Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 97; Taranta Mts passes, 8000 ft, Abyssinia. Range and Status. Endemic resident, Ethiopian and adjacent highlands. Ethiopia and Eritrea, common at 1700–2450 m, uncommon at 1060–1700 m, abundant above 2150 m (Zinner, 2001) and recorded up to 3300 m (Clement et al. 1993); 93 caught in an Addis Ababa garden in 2 months (Tyler 1979). Somalia, locally very common, between 1200 and 1970 m. Description. S. t. tristriatus Ru ¨ ppell: Eritrea and Ethiopia. ADULT Y: forehead to hindneck earth brown with narrow darker brown streaks; rest of upperparts greyish earth brown, mantle with a few faint darker shaft streaks. Tail dark brown. Narrow supercilium buffy white. Lores, cheeks and ear-coverts dark earth brown, merging with paler side of neck. Chin and upper throat greyish white, with a few blackish brown streaks at sides, merging into grey-brown lower throat, breast, flanks, belly and undertailcoverts. Remiges dull dark brown, primaries with narrow buff edges when fresh, tips of tertials slightly paler and greyer; rest of upperwing feathers dull brown. Underwing-coverts and axillaries greyish brown. Bill dusky; eyes light brown; legs light brown. Sexes alike. SIZE (8 YY, 5 XX): wing, Y 67–69 (685), X 65–69 (678); tail, Y 53–57 (558), X 53–59 (548); bill, Y 125–15 (138), X 13–14 (136); tarsus, Y 165–185 (175), X 17–18 (174). WEIGHT: (n ¼ 27, unsexed) 124–176 (158). IMMATURE: juv. more streaked above than ad.; underparts greybuff, with whiter chin and throat, and short dark brown streaks from chin to flanks and sides of belly. S. t. pallidior Phillips: N Somalia. Rather paler and greyer above and below than nominate tristriatus, streaks on mantle more pronounced.

Field Characters. Length 13 cm. A small plain brown seedeater of Ethiopian highlands; distinguished from

Serinus tristriatus

sympatric Streaky and Reichard’s Seedeaters S. striolatus and S. reichardi and Ankober Serin S. ankoberensis by short white supercilium and lack of any streaking, from Streaky by solid brown face without separate dark malar stripe.

510

FRINGILLIDAE

Voice. Tape-recorded (102, B, LEM, McVIC). Song a mixture of sweet and buzzy trills interspersed with unmusical twittering and much repetition of single notes and phrases, e.g. ‘djee-djee-djee-djee-tchueee-tchueeetchueee-tititititit-tsitsi-tsuee-tsuee-tsuitit-tsuiti’, often mingled with extensive imitations of other bird species (Zimmerman et al. 1996, q.v. for further variations). Calls include nasal ‘zee-zwee’ and ‘zu-weeo’; trilled ‘chirry’ or ‘chirru’ and sibilant ‘tissy’. General Habits. In Kenya striatipectus inhabits shrubby escarpments with scattered trees; further south, nominate reichardi is restricted to little-disturbed Brachystegia and Baikiaea woodlands; in Matengo Highlands, S Tanzania, in Faurea-Protea-Uapaka-Brachystegia-dominated woods; once in a mealie garden (Mozambique, Vincent 1936). Habits of both races are poorly known. In Kenya, shy and inconspicuous except when Y sings from a treetop or in courtship flight; territories may be large, since Y sings over a considerable area (Zimmerman et al. 1996). Occurs in pairs or flocks of 3–5 or 10–20; forages on ground; freely uses trees, and flock once seen flycatching from treetops. ‘A common upland species of gardens, cultivated land or

Brachystegia woodland, congregating into large flocks and doing damage to both corn and soft fruit at times’ (Mackworth-Praed and Grant 1960) may refer to S. gularis rather than S. reichardi. Food. No information. Breeding Habits. Solitary nester, or breeds in colonies of 4–5 pairs (Lynes 1934); territorial (see above). Y courts using ‘butterflying’ display flights, singing all the while. NEST and EGGS: ‘Nest of rootlets and stems lined with soft material usually at some height in a tree . . . Eggs 2–4, faint bluish white with a few dots and streaks of purplish black; c. 20  15’ (Mackworth-Praed and Grant 1960) may refer to S. gularis rather than S. reichardi. No subsequent descriptions known. LAYING DATES: E Africa, Region A, July, Region C, Jan– Mar; Zaı¨re, Lubumbashi, (gonads enlarged until Aug); Zambia, (gonads active Feb, Mar, Aug); Zaı¨re, (breeding starts in Jan). Key References Zimmerman, D.A. et al. (1998), Lynes, H. (1934, 1938), Vincent, J. (1936).

Plate 30

Serinus tristriatus Ru ¨ ppel. Brown-rumped Seedeater. Serin `a trois raies.

(Opp. p. 459)

Serinus tristriatus Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 97; Taranta Mts passes, 8000 ft, Abyssinia. Range and Status. Endemic resident, Ethiopian and adjacent highlands. Ethiopia and Eritrea, common at 1700–2450 m, uncommon at 1060–1700 m, abundant above 2150 m (Zinner, 2001) and recorded up to 3300 m (Clement et al. 1993); 93 caught in an Addis Ababa garden in 2 months (Tyler 1979). Somalia, locally very common, between 1200 and 1970 m. Description. S. t. tristriatus Ru ¨ ppell: Eritrea and Ethiopia. ADULT Y: forehead to hindneck earth brown with narrow darker brown streaks; rest of upperparts greyish earth brown, mantle with a few faint darker shaft streaks. Tail dark brown. Narrow supercilium buffy white. Lores, cheeks and ear-coverts dark earth brown, merging with paler side of neck. Chin and upper throat greyish white, with a few blackish brown streaks at sides, merging into grey-brown lower throat, breast, flanks, belly and undertailcoverts. Remiges dull dark brown, primaries with narrow buff edges when fresh, tips of tertials slightly paler and greyer; rest of upperwing feathers dull brown. Underwing-coverts and axillaries greyish brown. Bill dusky; eyes light brown; legs light brown. Sexes alike. SIZE (8 YY, 5 XX): wing, Y 67–69 (685), X 65–69 (678); tail, Y 53–57 (558), X 53–59 (548); bill, Y 125–15 (138), X 13–14 (136); tarsus, Y 165–185 (175), X 17–18 (174). WEIGHT: (n ¼ 27, unsexed) 124–176 (158). IMMATURE: juv. more streaked above than ad.; underparts greybuff, with whiter chin and throat, and short dark brown streaks from chin to flanks and sides of belly. S. t. pallidior Phillips: N Somalia. Rather paler and greyer above and below than nominate tristriatus, streaks on mantle more pronounced.

Field Characters. Length 13 cm. A small plain brown seedeater of Ethiopian highlands; distinguished from

Serinus tristriatus

sympatric Streaky and Reichard’s Seedeaters S. striolatus and S. reichardi and Ankober Serin S. ankoberensis by short white supercilium and lack of any streaking, from Streaky by solid brown face without separate dark malar stripe.

Serinus flavivertex Voice. Tape-recorded (B, SINC). Various sibilant calls, ‘pss-chip’, ‘pss-chu’, ‘wee-chu’, ‘chip’, ‘zit’, ‘pss-chuwee’. Song described as high-pitched but weak 3–4 note ‘sip-siptwis-twis’ or ‘sissis-sissis’ (Clement et al. 1993); this sounds like repetition of call notes rather than a full song. General Habits. Inhabits woodland, bushes, forest clearings and edges, juniper parkland, juniper scrub and undergrowth, in mountainous, in rocky country; also in large gardens as in Addis Ababa. Occurs singly or in pairs in breeding season and in pairs or small flocks out of it, when loose flocks sometimes wander widely. Perches low in trees and on ground. Food. ‘All kinds of seeds and seedlings’ (Cheesman and Sclater 1935–1936), and so has a reputation for damaging plants in gardens; mainly seeds of herbs, small plants and shrubs (Clement et al. 1993).

511

Breeding Habits. Solitary nester; monogamous; territorial. Y and X ruffle body plumage in Head-and-Tail-up display, Y accompanies X during nest-building, Y presents nest material during courtship, and helps to collect material but does not help build the nest (Hemmer 1976, van den Elzen 1985). NEST: a very neat cup made of fine, dry grass stems mixed with some spider-web, lined with fibres and animal hairs or wool; placed on horizontal fork in leafy tree or bush, commonly a juniper, at ‘any height’ above ground. EGGS: 3. Greenish white or pale greenish blue, with brown, violet, blackish or reddish spots, chiefly at large end. SIZE: c. 19  14. LAYING DATES: Ethiopia, Apr–Oct; Eritrea, June. Nothing further known. Key References Clement, P. et al. (1993), Hemmer, W. (1976), Mackworth-Praed, C.W. and Grant, C.H.B. (1960).

Subgenus Serinus Koch

Finches with sexes alike or nearly so; plumage mainly yellow-olive, streaky, some species with grey hindcrowns, napes and necks, others black-headed; rumps plain yellow. Lowland and montane. YY display by swaying with partly opened wings and transport nest material, XX build nest, juvs have red gapes. Nest rims accumulate a thick layer of nestling faeces (i.e. serinus, canaria, canicollis, nigriceps: Nicolai 1960, van den Elzen 1985; biology of other species insufficiently known). 9 species, 7 occurring in Africa; 3 endemic: nigriceps (montane, Ethiopia), flavivertex (montane, eastern Africa, Angola) and canicollis (lowland, South Africa). Grey-necked syriacus (Syria, Israel, vagrant in Africa) and canaria (Canary Is; not in Africa) would appear to be closely related with canicollis. S. corsicana (not in Africa) is an allospecies of S. citrinella. S. serinus (N Africa, W Europe) is heavily striped, with sexes not alike and with Ochrospiza cheek pattern. The remaining 2 are citrinella (SW European mountains) and the aberrantly coloured pusillus (SW Asia), both vagrant in Africa. S. canicollis, flavivertex, serinus, canaria and citrinella were treated by Hall and Moreau (1970) as composing a superspecies. S. flavivertex and canicollis have been treated as conspecific by most authorities. S. syriacus has been considered conspecific with serinus (see Sibley and Monroe 1990), but we discern no close alliance. Some representatives of this subgenus are close to Alario spp. in reproductive characters and others to Carduelis spp. in plumage and bill morphology.

Serinus flavivertex (Blanford). Yellow-crowned Canary. Serin a ` calotte jaune.

Plate 30

Crithagra flavivertex Blanford, 1869. Ann. Mag. Nat. Hist., ser. 4, 4, p. 330; Tigre´, Abyssinia, type from Adigrat, Tigre´ Prov., fide W.L. Sclater, 1930, Syst. Av. Aethiop., p. 817.

(Opp. p. 459)

Forms a superspecies with S. canicollis. Range and Status. Endemic resident, highlands in Eritrea, ˆ i. Ethiopia, E Africa, W Angola and west of L. Malaw Eritrea, uncommon to locally common at 1675–2400 m (Zinner 2001). Ethiopia, rarely below 1650 m, locally common above that level, from Adigrat near Eritrean border to Yabelo in S, west to Injibara, Gojam Prov., where occurs plentifully at 2440 m. Sudan, uncommon to common from 1800 m to >2400 m in Imatong Mts. Uganda, Kigezi

district. Kenya, common to abundant at 2200–4000 m, occurring down to 1600 m in rainy season, and up to 4300 m; widespread, from Mt Elgon and Cheranganis to Mau, Aberdares and Mt Kenya; ubiquitous at Mau Narok; in forest on Mt Loma, west of Lodwar. Zaı¨re, widespread and locally very common, e.g. highlands southwest of L. Kivu; in Itombwe, common at 2070–2770 m; not in Marungu Mts. Occurs in extreme W Rwanda and Burundi,

Serinus flavivertex Voice. Tape-recorded (B, SINC). Various sibilant calls, ‘pss-chip’, ‘pss-chu’, ‘wee-chu’, ‘chip’, ‘zit’, ‘pss-chuwee’. Song described as high-pitched but weak 3–4 note ‘sip-siptwis-twis’ or ‘sissis-sissis’ (Clement et al. 1993); this sounds like repetition of call notes rather than a full song. General Habits. Inhabits woodland, bushes, forest clearings and edges, juniper parkland, juniper scrub and undergrowth, in mountainous, in rocky country; also in large gardens as in Addis Ababa. Occurs singly or in pairs in breeding season and in pairs or small flocks out of it, when loose flocks sometimes wander widely. Perches low in trees and on ground. Food. ‘All kinds of seeds and seedlings’ (Cheesman and Sclater 1935–1936), and so has a reputation for damaging plants in gardens; mainly seeds of herbs, small plants and shrubs (Clement et al. 1993).

511

Breeding Habits. Solitary nester; monogamous; territorial. Y and X ruffle body plumage in Head-and-Tail-up display, Y accompanies X during nest-building, Y presents nest material during courtship, and helps to collect material but does not help build the nest (Hemmer 1976, van den Elzen 1985). NEST: a very neat cup made of fine, dry grass stems mixed with some spider-web, lined with fibres and animal hairs or wool; placed on horizontal fork in leafy tree or bush, commonly a juniper, at ‘any height’ above ground. EGGS: 3. Greenish white or pale greenish blue, with brown, violet, blackish or reddish spots, chiefly at large end. SIZE: c. 19  14. LAYING DATES: Ethiopia, Apr–Oct; Eritrea, June. Nothing further known. Key References Clement, P. et al. (1993), Hemmer, W. (1976), Mackworth-Praed, C.W. and Grant, C.H.B. (1960).

Subgenus Serinus Koch

Finches with sexes alike or nearly so; plumage mainly yellow-olive, streaky, some species with grey hindcrowns, napes and necks, others black-headed; rumps plain yellow. Lowland and montane. YY display by swaying with partly opened wings and transport nest material, XX build nest, juvs have red gapes. Nest rims accumulate a thick layer of nestling faeces (i.e. serinus, canaria, canicollis, nigriceps: Nicolai 1960, van den Elzen 1985; biology of other species insufficiently known). 9 species, 7 occurring in Africa; 3 endemic: nigriceps (montane, Ethiopia), flavivertex (montane, eastern Africa, Angola) and canicollis (lowland, South Africa). Grey-necked syriacus (Syria, Israel, vagrant in Africa) and canaria (Canary Is; not in Africa) would appear to be closely related with canicollis. S. corsicana (not in Africa) is an allospecies of S. citrinella. S. serinus (N Africa, W Europe) is heavily striped, with sexes not alike and with Ochrospiza cheek pattern. The remaining 2 are citrinella (SW European mountains) and the aberrantly coloured pusillus (SW Asia), both vagrant in Africa. S. canicollis, flavivertex, serinus, canaria and citrinella were treated by Hall and Moreau (1970) as composing a superspecies. S. flavivertex and canicollis have been treated as conspecific by most authorities. S. syriacus has been considered conspecific with serinus (see Sibley and Monroe 1990), but we discern no close alliance. Some representatives of this subgenus are close to Alario spp. in reproductive characters and others to Carduelis spp. in plumage and bill morphology.

Serinus flavivertex (Blanford). Yellow-crowned Canary. Serin a ` calotte jaune.

Plate 30

Crithagra flavivertex Blanford, 1869. Ann. Mag. Nat. Hist., ser. 4, 4, p. 330; Tigre´, Abyssinia, type from Adigrat, Tigre´ Prov., fide W.L. Sclater, 1930, Syst. Av. Aethiop., p. 817.

(Opp. p. 459)

Forms a superspecies with S. canicollis. Range and Status. Endemic resident, highlands in Eritrea, ˆ i. Ethiopia, E Africa, W Angola and west of L. Malaw Eritrea, uncommon to locally common at 1675–2400 m (Zinner 2001). Ethiopia, rarely below 1650 m, locally common above that level, from Adigrat near Eritrean border to Yabelo in S, west to Injibara, Gojam Prov., where occurs plentifully at 2440 m. Sudan, uncommon to common from 1800 m to >2400 m in Imatong Mts. Uganda, Kigezi

district. Kenya, common to abundant at 2200–4000 m, occurring down to 1600 m in rainy season, and up to 4300 m; widespread, from Mt Elgon and Cheranganis to Mau, Aberdares and Mt Kenya; ubiquitous at Mau Narok; in forest on Mt Loma, west of Lodwar. Zaı¨re, widespread and locally very common, e.g. highlands southwest of L. Kivu; in Itombwe, common at 2070–2770 m; not in Marungu Mts. Occurs in extreme W Rwanda and Burundi,

Serinus flavivertex Voice. Tape-recorded (B, SINC). Various sibilant calls, ‘pss-chip’, ‘pss-chu’, ‘wee-chu’, ‘chip’, ‘zit’, ‘pss-chuwee’. Song described as high-pitched but weak 3–4 note ‘sip-siptwis-twis’ or ‘sissis-sissis’ (Clement et al. 1993); this sounds like repetition of call notes rather than a full song. General Habits. Inhabits woodland, bushes, forest clearings and edges, juniper parkland, juniper scrub and undergrowth, in mountainous, in rocky country; also in large gardens as in Addis Ababa. Occurs singly or in pairs in breeding season and in pairs or small flocks out of it, when loose flocks sometimes wander widely. Perches low in trees and on ground. Food. ‘All kinds of seeds and seedlings’ (Cheesman and Sclater 1935–1936), and so has a reputation for damaging plants in gardens; mainly seeds of herbs, small plants and shrubs (Clement et al. 1993).

511

Breeding Habits. Solitary nester; monogamous; territorial. Y and X ruffle body plumage in Head-and-Tail-up display, Y accompanies X during nest-building, Y presents nest material during courtship, and helps to collect material but does not help build the nest (Hemmer 1976, van den Elzen 1985). NEST: a very neat cup made of fine, dry grass stems mixed with some spider-web, lined with fibres and animal hairs or wool; placed on horizontal fork in leafy tree or bush, commonly a juniper, at ‘any height’ above ground. EGGS: 3. Greenish white or pale greenish blue, with brown, violet, blackish or reddish spots, chiefly at large end. SIZE: c. 19  14. LAYING DATES: Ethiopia, Apr–Oct; Eritrea, June. Nothing further known. Key References Clement, P. et al. (1993), Hemmer, W. (1976), Mackworth-Praed, C.W. and Grant, C.H.B. (1960).

Subgenus Serinus Koch

Finches with sexes alike or nearly so; plumage mainly yellow-olive, streaky, some species with grey hindcrowns, napes and necks, others black-headed; rumps plain yellow. Lowland and montane. YY display by swaying with partly opened wings and transport nest material, XX build nest, juvs have red gapes. Nest rims accumulate a thick layer of nestling faeces (i.e. serinus, canaria, canicollis, nigriceps: Nicolai 1960, van den Elzen 1985; biology of other species insufficiently known). 9 species, 7 occurring in Africa; 3 endemic: nigriceps (montane, Ethiopia), flavivertex (montane, eastern Africa, Angola) and canicollis (lowland, South Africa). Grey-necked syriacus (Syria, Israel, vagrant in Africa) and canaria (Canary Is; not in Africa) would appear to be closely related with canicollis. S. corsicana (not in Africa) is an allospecies of S. citrinella. S. serinus (N Africa, W Europe) is heavily striped, with sexes not alike and with Ochrospiza cheek pattern. The remaining 2 are citrinella (SW European mountains) and the aberrantly coloured pusillus (SW Asia), both vagrant in Africa. S. canicollis, flavivertex, serinus, canaria and citrinella were treated by Hall and Moreau (1970) as composing a superspecies. S. flavivertex and canicollis have been treated as conspecific by most authorities. S. syriacus has been considered conspecific with serinus (see Sibley and Monroe 1990), but we discern no close alliance. Some representatives of this subgenus are close to Alario spp. in reproductive characters and others to Carduelis spp. in plumage and bill morphology.

Serinus flavivertex (Blanford). Yellow-crowned Canary. Serin a ` calotte jaune.

Plate 30

Crithagra flavivertex Blanford, 1869. Ann. Mag. Nat. Hist., ser. 4, 4, p. 330; Tigre´, Abyssinia, type from Adigrat, Tigre´ Prov., fide W.L. Sclater, 1930, Syst. Av. Aethiop., p. 817.

(Opp. p. 459)

Forms a superspecies with S. canicollis. Range and Status. Endemic resident, highlands in Eritrea, ˆ i. Ethiopia, E Africa, W Angola and west of L. Malaw Eritrea, uncommon to locally common at 1675–2400 m (Zinner 2001). Ethiopia, rarely below 1650 m, locally common above that level, from Adigrat near Eritrean border to Yabelo in S, west to Injibara, Gojam Prov., where occurs plentifully at 2440 m. Sudan, uncommon to common from 1800 m to >2400 m in Imatong Mts. Uganda, Kigezi

district. Kenya, common to abundant at 2200–4000 m, occurring down to 1600 m in rainy season, and up to 4300 m; widespread, from Mt Elgon and Cheranganis to Mau, Aberdares and Mt Kenya; ubiquitous at Mau Narok; in forest on Mt Loma, west of Lodwar. Zaı¨re, widespread and locally very common, e.g. highlands southwest of L. Kivu; in Itombwe, common at 2070–2770 m; not in Marungu Mts. Occurs in extreme W Rwanda and Burundi,

512

FRINGILLIDAE

Serinus flavivertex

? ?

e.g. Bururi Forest. Tanzania, high ground in Crater and Mbulu highlands, Mts Hanang, Meru and Kilimanjaro; on Kilimanjaro, at 2200–>2500 m on W slopes and 1500– >2900 m on S slopes; in S, widespread in highlands from near Ikomba to Mbeya, Njombe, Iringa and Udzungwas. ˆ i, on Nyika up to Zambia, only on Nyika Plateau. Malaw 2440 m; frequent in Viphya Mts east of Mzimba, at 1830 m; sparse in Mafinga Mts and Misuku Hills. Angola, locally common from Serra da Chela and Tundavala (Huı´la), Huambo and W Bie´ to S Cuanza Sul. Description. S. f. flavivertex (Blanford): Eritrea to N Tanzania. ADULT Y: forehead and forecrown golden yellow, merging into olive-green hindcrown; nape, hindneck, sides of neck, mantle, scapulars and back olive-green with dusky brown streaks; rump and uppertail-coverts greenish yellow. Tail feathers blackish, T1– T5 edged bright greenish yellow; undersides dark grey-brown. Broad greenish yellow superciliary stripe extends back to side of nape; below this a dusky green stripe through lores and behind eye; ear-coverts dark olive-green. Cheeks to chin, throat, breast, flanks and upper belly greenish yellow, greener on sides of breast and flanks, with diffuse dusky streaks; lower belly, vent and thighs white; undertail-coverts pale yellow. Remiges blackish, middle and distal part of primaries (proximal to emarginations) narrowly edged yellow, distal part of secondaries and tertials more broadly edged bright greenish yellow; primary coverts and alula blackish; greater and median coverts blackish, finely edged and broadly tipped bright yellow; lesser coverts olive-green. On closed wing, a prominent yellow panel, separated by black secondary bases from 2 broad bright yellow wing-bars. Bill slate above, brown below; eyes brown; legs black. ADULT X: forehead and forecrown greenish yellow with dusky streaks; hindcrown to back and scapulars dull brownish olive-green, streaked dusky brown; rump and uppertail-coverts yellowish green. Tail as ad. Y, but feather edging more greenish. Supercilium greenish yellow. Cheeks to chin, throat, breast and upper belly dull yellowish green, sides of breast and flanks duller with narrow dusky streaks; belly more extensively white than in Y; undertail-coverts

yellowish white. Wings as ad. Y, but yellow edges and tips of tertials, greater coverts and median coverts narrower and more greenish. SIZE (10 YY, 9 XX): wing, Y 78–81 (805), X 74–78 (757); tail, Y 50–55 (531), X 49–52 (502); bill, Y 12–13 (123), X 11–13 (120); tarsus, Y 15–16 (156), X 15–16 (153). WEIGHT: unsexed, Kenya (n ¼ 29) 13–16 (140). IMMATURE: juv. has upperparts broadly streaked dark brown and pale tawny-buff; sides of head and underparts pale yellowish buff with sharp narrow dark brown streaks. Tail feathers dark brown, edged yellow. Wing feathers browner than in ad., outer secondaries and tertials edged pale buff, tips of greater and median coverts brownish buff. S. f. sassi Neumann: SW Uganda, E Zaı¨re, S Tanzania, N ˆ i and NE Zambia. Differs from nominate flavivertex in Malaw having black on tail feathers confined to central streak, outer border broadly greenish yellow, inner webs tinged greenish. WEIGHT: (S Tanzania) Y (n ¼ 3) 155–16 (156), 2 XX 125, 175, unsexed (n ¼ 9) 13–17 (149). S. f. huillensis Sousa: Angola. Y has crown, cheeks and underparts brighter purer yellow than in nominate race; both sexes more yellowish green above, with streaking almost absent; shows prominent yellow tail panel, but wing-bars obscured by broad pale yellow edges to greater and median coverts.

Field Characters. Length 115–125 cm. A highland canary; longish tail gives slim shape. Y marked by golden yellow forecrown and black and yellow wings; yellow supercilium curls round behind olive cheek patch, underparts and rump yellow, upperparts yellow-green with light streaking; wing black with 2 broad yellow wing-bars and yellow panel on folded flight feathers. X duller and more streaked, less yellow on crown, belly white; immature buffy brown, streaked above and below, wing an echo of ads’, with buff bars and panel. Y of pale Angolan race huillensis almost unstreaked above. Separated altitudinally from Yellowfronted and White-bellied Canaries S. mozambicus and S. dorsostriatus, told from both by black and yellow wings, from Yellow-fronted by longer tail, lack of black lores and moustache. Voice. Tape-recorded (B, C, KAE P, NOR, PEA). Song a continuous bustling, tinkling jumble of sweet notes and trills, many with quality of European Goldfinch Carduelis carduelis, lasting 6–10 s, often up to 15 s; several YY may sing in chorus (Zimmerman et al. 1996). Calls include a liquid trill, often repeated in series, also a rising ‘sweet pea’ or ‘peet’. General Habits. In E Africa inhabits clearings in and edges of highland forest, moorlands, also pastures, cultivation and gardens; Acacia mollissima trees bordering L. Delia, Eritrea, where also occurs in eucalypts, junipers and upland scrub; clearings in juniper woods at 1675– 1830 m (Yabelo, S Ethiopia); Podocarpus trees in high forest (Imatong Mts, Sudan); savanna and borders of forest (Itombwe, Zaı¨re); montane grassland with scattered bushes ˆ i). (Malaw In pairs, parties of 4–6 after breeding, and commonly forms flocks of hundreds, at least at higher elevations. Forages mainly on ground, also (Imatong Mts, Sudan) on Podocarpus trees, herbs and in Sorghum crops (Nikolaus 1979). Flight less undulating than in most congeners.

Serinus canicollis Resident; wanders; a partial vertical migrant in Kenya ˆ i, where absent from Nyika Plateau (see above) and Malaw in Jan and from Viphya Mts in Mar–July. Food. Seeds up to size of rice kernels. Breeding Habits. Solitary nester; territorial. Y sings from very top of conifer and launches itself into slow ‘butterfly’ courtship flight. NEST: open cup made of rootlets, lined with dry petals, placed 1–25 m above ground in health, Protea bush or (commonly) in Grevillea tree (Chapin 1954). ˆ i); at L. Kivu, Zaı¨re, ‘broods EGGS: 4 (9 clutches, Malaw of only 2 are frequent’ (Chapin 1954). White to pale green,

513

sparingly spotted with dark brown and grey. SIZE: (n ¼ ?) 177–182  130–133. LAYING DATES: Ethiopia, June; Sudan, Nov; E Africa: Region A, Oct–Nov, Region D, Dec–Feb, May–Sept, with pronounced peak in May–June (14 out of 23 records), but at Mau Narok breeds in all months (nesting or feeding young) with peaks in Apr–May and Oct–Nov, at start of rains and start of dry season (Brown and Britton 1980); Uganda, L. Bunyoni, (breeding condition Apr–June); Zaı¨re, L. Kivu, (breeds from Aug to Mar), Itombwe, (breeding condition ˆ i, Oct–Nov; Angola, (oviduct egg Apr, juvs Oct); Malaw Sept, breeding condition Oct). Nothing further known.

Serinus canicollis (Swainson). Cape Canary. Serin du Cap.

Plate 30

Crithagra canicollis Swainson, 1838 (1837?). Anim. Menag., p. 317; Africa, restricted to Cape of Good Hope by Vincent, 1952, Check List Birds South Africa, p. 115.

(Opp. p. 459)

Forms a superspecies with S. flavivertex. Range and Status. Endemic resident, vertical migrant and wanderer, SE South Africa and highlands of Zimbabwe/ Mozambique border. Introduced to Re´union I., where scarce (R. Safford, pers. comm.). Zimbabwe, at 1350–2300 m in Inyanga Highlands, Mt. Inyangani, south to Melsetter in Chimanimani Mts; common at Stapleford; known to occur down to 900 m in E Inyanga (Aug) and visits Mt Selinda and Chipinga Uplands (June). Mozambique, present in mountains along Zimbabwe border, north to headwaters of R. Pungwe. South Africa as mapped; uncommon in Kamieskroon area, W Cape Prov., common to very common in Cape Peninsula and greater Cape Town (Oatley 2000). In Free State, locally very common resident in E mts; isolated populations west to Bloemfontein, Franklin Game Res., Naval Hill and Kimberley (Nuttall 2000). In Transvaal, common in E Highveld and Escarpment region including Soutpansberg and Blouberg; west across Highveld to Suikersbosrand and Magliesberg hills where local; also local in Waterberg; elsewhere in Transvaal a rare vagrant (Tarboton et al. 1987). Common in W Swaziland, and Lesotho. Range and abundance thought to have increased in Karoo, with planting of exotic trees such as pines and poplars. Description. S. c. thompsonae Roberts: Transvaal and Natal to E Griqualand. ADULT Y: forehead golden yellow, streaked with green, grading on hindcrown to olive-green streaked with yellow; nape to upper mantle, rear of ear-coverts and sides of neck pale grey, forming a broad shawl; lower mantle, back and scapulars deep olive-green, feathers with fine dusky shaft streaks, tips tinged grey when fresh; rump and uppertail-coverts greenish yellow. Tail feathers dark brown, tinged yellow, outer webs edged bright yellow; undersides pale yellow. Supercilium yellow but not forming contrasting stripe; band through lores and mark behind eye dusky; front of ear-coverts dull olive-green. Cheeks, chin, throat and upper breast yellow with golden or brownish tinge; sides of breast and flanks yellow, washed olive-green; lower breast and upper belly lighter, cleaner yellow; lower belly, vent and thighs greyish white; undertail-coverts yellow. Remiges blackish,

Serinus canicollis

primaries narrowly edged bright yellow proximal to emarginations, secondaries edged olive-green and tertials more broadly so; primary coverts and alula blackish, fringed olive-green; greater coverts blackish with olive-green edges, broader and yellower distally and around tips, forming distinct wing-bar; median and lesser coverts yellowish olive-green. Underwingcoverts and axillaries greyish white. Bill pointed (A), dark horn above, pale horn to creamy white below; eyes brown; legs dark brown or dark pinkish brown. ADULT X: forehead to forecrown greenish yellow with narrow dusky streaks; hindcrown to upper mantle, sides of neck and rear of ear-coverts greyish, tinged

Serinus canicollis Resident; wanders; a partial vertical migrant in Kenya ˆ i, where absent from Nyika Plateau (see above) and Malaw in Jan and from Viphya Mts in Mar–July. Food. Seeds up to size of rice kernels. Breeding Habits. Solitary nester; territorial. Y sings from very top of conifer and launches itself into slow ‘butterfly’ courtship flight. NEST: open cup made of rootlets, lined with dry petals, placed 1–25 m above ground in health, Protea bush or (commonly) in Grevillea tree (Chapin 1954). ˆ i); at L. Kivu, Zaı¨re, ‘broods EGGS: 4 (9 clutches, Malaw of only 2 are frequent’ (Chapin 1954). White to pale green,

513

sparingly spotted with dark brown and grey. SIZE: (n ¼ ?) 177–182  130–133. LAYING DATES: Ethiopia, June; Sudan, Nov; E Africa: Region A, Oct–Nov, Region D, Dec–Feb, May–Sept, with pronounced peak in May–June (14 out of 23 records), but at Mau Narok breeds in all months (nesting or feeding young) with peaks in Apr–May and Oct–Nov, at start of rains and start of dry season (Brown and Britton 1980); Uganda, L. Bunyoni, (breeding condition Apr–June); Zaı¨re, L. Kivu, (breeds from Aug to Mar), Itombwe, (breeding condition ˆ i, Oct–Nov; Angola, (oviduct egg Apr, juvs Oct); Malaw Sept, breeding condition Oct). Nothing further known.

Serinus canicollis (Swainson). Cape Canary. Serin du Cap.

Plate 30

Crithagra canicollis Swainson, 1838 (1837?). Anim. Menag., p. 317; Africa, restricted to Cape of Good Hope by Vincent, 1952, Check List Birds South Africa, p. 115.

(Opp. p. 459)

Forms a superspecies with S. flavivertex. Range and Status. Endemic resident, vertical migrant and wanderer, SE South Africa and highlands of Zimbabwe/ Mozambique border. Introduced to Re´union I., where scarce (R. Safford, pers. comm.). Zimbabwe, at 1350–2300 m in Inyanga Highlands, Mt. Inyangani, south to Melsetter in Chimanimani Mts; common at Stapleford; known to occur down to 900 m in E Inyanga (Aug) and visits Mt Selinda and Chipinga Uplands (June). Mozambique, present in mountains along Zimbabwe border, north to headwaters of R. Pungwe. South Africa as mapped; uncommon in Kamieskroon area, W Cape Prov., common to very common in Cape Peninsula and greater Cape Town (Oatley 2000). In Free State, locally very common resident in E mts; isolated populations west to Bloemfontein, Franklin Game Res., Naval Hill and Kimberley (Nuttall 2000). In Transvaal, common in E Highveld and Escarpment region including Soutpansberg and Blouberg; west across Highveld to Suikersbosrand and Magliesberg hills where local; also local in Waterberg; elsewhere in Transvaal a rare vagrant (Tarboton et al. 1987). Common in W Swaziland, and Lesotho. Range and abundance thought to have increased in Karoo, with planting of exotic trees such as pines and poplars. Description. S. c. thompsonae Roberts: Transvaal and Natal to E Griqualand. ADULT Y: forehead golden yellow, streaked with green, grading on hindcrown to olive-green streaked with yellow; nape to upper mantle, rear of ear-coverts and sides of neck pale grey, forming a broad shawl; lower mantle, back and scapulars deep olive-green, feathers with fine dusky shaft streaks, tips tinged grey when fresh; rump and uppertail-coverts greenish yellow. Tail feathers dark brown, tinged yellow, outer webs edged bright yellow; undersides pale yellow. Supercilium yellow but not forming contrasting stripe; band through lores and mark behind eye dusky; front of ear-coverts dull olive-green. Cheeks, chin, throat and upper breast yellow with golden or brownish tinge; sides of breast and flanks yellow, washed olive-green; lower breast and upper belly lighter, cleaner yellow; lower belly, vent and thighs greyish white; undertail-coverts yellow. Remiges blackish,

Serinus canicollis

primaries narrowly edged bright yellow proximal to emarginations, secondaries edged olive-green and tertials more broadly so; primary coverts and alula blackish, fringed olive-green; greater coverts blackish with olive-green edges, broader and yellower distally and around tips, forming distinct wing-bar; median and lesser coverts yellowish olive-green. Underwingcoverts and axillaries greyish white. Bill pointed (A), dark horn above, pale horn to creamy white below; eyes brown; legs dark brown or dark pinkish brown. ADULT X: forehead to forecrown greenish yellow with narrow dusky streaks; hindcrown to upper mantle, sides of neck and rear of ear-coverts greyish, tinged

514

FRINGILLIDAE

olive-green; lower mantle to back and scapulars dull olive-green with narrow dusky shaft streaks, more prominent than in Y; rump and uppertail-coverts yellowish olive-green. Tail as in ad. Y but feather edges more greenish yellow. Dusky stripe through lores to behind eye; front of ear-coverts and cheeks dull olivegreen, finely streaked dusky. Chin, throat and upper breast dull pale olive-green, tinged grey; sides of breast and flanks greyish olive-green with a few dusky shaft streaks; lower breast and upper belly pale, dull yellow, grading to greyish white on lower belly and vent; undertail-coverts pale yellow. Wings as ad. Y, but median and lesser coverts duller and greener, outer borders of tertials and greater coverts duller, greener and narrower. SIZE (10 YY, 10 XX): wing, Y 75–83 (790), X 72–78 (758); tail, Y 52–60 ( 5 6 0 ) , X 50–55 (523); bill, Y 115–125 (122), X 11–13 (120); tarsus, Y 15–165 (155), X 15–16 (154). WEIGHT (this race and nominate canicollis): (n ¼ 83, unsexed) 13–195 (155). IMMATURE: juv. broadly streaked above with dark brown and pale buffish or tawny-brown; sides of head and underparts pale buff, with dark brown streaks, narrow on belly and undertailcoverts; breast, belly and flanks tinged yellow. Tail feathers dark brown, tinged and edged yellow. Wing feathers browner than in ad., outer secondaries and tertials edged pale buff, edges and tips of greater coverts and broad tips of median and lesser coverts pale buffish brown. NESTLING (S. c. canicollis): at about 1 week skin pink, greyer on feather tracts; palate bright carmine; edge of mouth and gape pale yellow; a narrow black line at edge of palate between the carmine and pale yellow (B) (photo in Steyn 1996). S. c. canicollis (Swainson): W and SW Cape Prov. to E Cape and W Free State. Y slightly duller (less golden) olive-green above than thompsonae. S. c. griseitergum Clancey: Zimbabwe, Mozambique. Y has forehead, crown and face greener than in thompsonae, shawl deeper, more bluish, grey, and back darker green with more pronounced streaking; underparts tinged greener. X colder on upperparts, less brownish than in nominate race, with blacker streaking; forehead greener; chin and throat greyer.

buzzes; scrambled character apparent from sonagram in Maclean (1993). Some notes have tinny quality but do not quite have the timbre of European Goldfinch Carduelis carduelis. Twittering calls may run into song without interruption. Song continuous for up to 20 s without pause; during height of breeding season full song bout of individual may last for 15 min; in colony, when one bird starts to sing it sets off others, and this may develop into chorus lasting up to 30 min (Skead 1960). When singing from perch, bird’s head is raised, body sways gently from side to side, and throat ‘‘seems ready to burst with song’’ (Skead 1960). X does not sing. Calls thin and sweet, ‘peet’, descending ‘tswee-yew’ and rising ‘way-yee’ or more nasal ‘dzoo-wee’, also short tittering trills, which may be given in flight; alarm a plaintive ‘sweet’ with anxious overtone. In courtship Y makes mewing ‘meh, meh, meh’ when approaching X; fighting birds give harsh, rasping ‘swiz, swiz . . . ’; chicks in nest give plaintive ‘cheeet’, and fledglings a variety of hunger calls, ‘cheetoo’, ‘chirree-reet’ and ‘chee-reet’ (Skead 1960).

Field Characters. Length 11–14 cm. A slim canary with longish tail, readily identified by grey shawl from nape to mantle and sides of neck (continuing across breast in X), and by plain face without any distinct markings; crown and throat brownish gold, becoming yellow on underparts; folded wing mainly green with indistinct pale bar. X like Y but duller and more streaked; juv. buffy brown, streaked above and below, tinged yellow on underparts, with broad pale buff bar on wing; can look confusingly like Forest Canary S. scotops, but seldom strays far from family group with grey-naped ad. (Oatley 2000). Yellow-fronted Canary S. mozambicus has short tail, some grey on nape but black and yellow striped face.

General Habits. In Zimbabwe inhabits montane grassland, bracken-briar and hillsides with Philippia, Protea, Leucosidea and other scrub; frequent in plantations of Pinus patula, attracted especially to recently cut-over sections with pine slash (Irwin 1981). In South Africa, edges of open grassland with thin and scattered woods, copses, pine plantations, fynbos, gardens, parks, playing fields, road verges, fallow fields, arable land, orchards and vineyards; grassveld in Drakensberg Mts; occasionally in treeless areas, coastal dunes and on rocky and sandy beaches. Occurs singly, in pairs and family parties; even in breeding season in flocks commonly of up to c. 12 birds and often up to 50–70; after breeding, can congregate in hundreds, up to 500, in e.g. slashed Pinus patula plantation or at a ripe sunflower crop. Forages on ground where gait a hop, and amongst scrub, proteas, weedy plants and short, bushy growth on mountain slopes; takes seeds from fruiting heads by perching on stems next to them. Often forages in company of other seed-eating birds. Confiding. Flight undulating, but less so than in some smaller congeners. Roosts in large flocks, in trees; at a site near Grahamstown, E Cape Prov., small flock arrives to roost every Apr and builds up to c. 50 birds in May and June, sometimes with influx of 200 (Skead 1960). Flock flies to water at any time of day and birds drink communally. Evidently a vertical migrant in Zimbabwe (see above). A distinct increase in reporting rates in spring in SW and SE Cape (Harrison et al. 1997), and species appears to move out of arid N parts of its range during summer (Hockey et al. 1989); bird in S Cape Prov. recovered 80 km from place where ringed; bird of nominate race once found in range of thompsonae in E Transvaal; wanders locally in Karoo, north to Victoria West where perhaps only a nonbreeding visitor; flock stays in an area for weeks, then suddenly moves away.

Voice. Tape-recorded (20, 33, 58, 72, 88, 99, 104, B, C, F, LEM, PAY). Song, given from prominent perch or in flight, pleasing and sustained, a bustling jumble of brief sweet and sharp notes with a few short churrs and low

Food. Seeds of Pinus patula, Bidens pilosa, Alternanthera pungens, Psidium guajava, Eriobotrya japonica, Ursinia, Senecio, Osteospermum, Venidium, Cephalaria, sunflower, grasses and cereals; buds and petals of Salvia splendens,

Serinus canicollis S. chamelaeagnea and Buddleia (Skead 1960, Brickell and Konigkramer 1997). A few insects, e.g. grasshoppers, given by Y to incubating mate. In SW Cape Prov., 36% of 530 feeds were at Inula, 31% at Senecio, 11% at Metalasia, 9% at Stoebe, 7% at Chenopodium and the rest at Athanasia, heathers, Gnidia, Amaranthus, Sonchus, Stellaria, Elytropappus and Eriocephalus (Milewski 1978a); particularly fond of seeds (achenes) of Compositae, and feeds on waysides where Arctotheca, Arctotis, Senecio and Ursinia abound (Oatley 2000). Breeding Habits. Solitary or semi-colonial nester; monogamous; territorial. In colony of 10 nests, some only 3–5 m apart; 3 old and 11 current nests along 300 m transect in 2 ha patch of Leucosidea sericea trees on Vaal R. (Wolff and Jacobsen 1980); colony of 24 nests in a stand of exotic trees (R. Martin in Ginn et al. 1989). An oak, pine or other tree next to a farmhouse is used for nesting year after year; one such site used for 16 years, another for 30 years (Skead 1960). Y has ‘butterfly’ courtship flight with slow, deliberate wingbeats, bird sometimes in sustained song; this courtship flight coincides with beginning of nest-building. Singing strongly, one bird chases another in flight; rivals can also fight by grappling in the air, falling to ground interlocked, accompanied by harsh screeches (Skead 1960). Courting Y approaches a perched X by edging along perch towards her, swaying his body as he does so, singing or silently or uttering a mewing call, the approach ending in a sudden dash which may put X to flight, when Y follows; sidling Y once seen carrying piece of fluff in bill as he approached X. Once, Y alighted next to nest-building X carrying material in her bill, sang at her and hopped over her to perch on other side; she stretched body and swayed slowly from side to side. Birds in a colony often gape and posture at each other in mild aggression. NEST: thick-walled cup made of fine rootlets, long Helichrysum tendrils and weed stems, with cobweb and a few tiny twigs intermixed (B, C), also often lichens, moss, pine needles, leaves and bits of rag and string, compactly lined with soft plant down, hair, thin Helichrysum tendrils, sheep’s wool, cow hair and usually a few small feathers, with rootlets around rim. Rim rootlets are characteristic of this species; and appear to act as suitable substrate for eventual accumulation of thick coat of nestlings’ faeces. Ext. diam. 77–89, int. diam. 46–56, ext. depth 46–51, int. depth 26–33. Species seldom makes use of indigenous vegetation for nesting but places nest mainly in exotics: to 18 m up in clumps of tall pines or oaks, and smaller trees such as peach and citrus; rarely in native Leucosidea sericea; on a fork or horizontal branch, 18–6 m above ground, uncommonly down to 10 m (in vineyards) and up to 18 m. X searching for new nest site industriously moves along tree branches, squats down in a fork and rotates body on it several times, then moves to another fork. Nest built by X alone, taking up to 2 weeks or once 22 days; she starts by laying down a horizontal ring of long tendrils, building first the base then the sides upon it. Y often accompanies her back and forth on material-gathering trips, and sings or gives tittering call whilst she builds. EGGS: 2–5, usually 3–4, av. (n ¼ 74) 32. Laid on successive days. White, or palest greenish or bluish, with

scattered chocolate-brown and small black speckles, mainly at broad end; sometimes sparingly speckled or blotched or scrolled with reddish brown or greyish purple. SIZE: (n ¼ 60) 158–189  116–136 (172  128).

LAYING DATES: Zimbabwe, Sept–Dec and Feb, mainly Oct–Dec, particularly Nov; South Africa, SW Cape Prov., Aug–Dec (mainly Sept–Oct); E Cape Prov., July–Dec; KwaZulu-Natal and Transvaal, Aug–Dec; Free State, Dec–Mar. INCUBATION: eggs generally all hatch on same day, over a 12-h period; some clutches hatch over 2 days. Incubation begins with 3rd egg laid, even when complete clutch will be of 4; by X alone, sitting tightly, fed on nest from outset by Y; X greets Y’s arrival with repetitive tittering call; Y stands on edge of nest and gives food piecemeal, taking

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c. 30 s; X is fed by regurgitation but rarely Y brings whole insect in bill. Period: 12–16 (usually 12–14) days; 2 neighbouring clutches of 4 eggs were laid concurrently, one hatched in 13 and 14 days, the other in 15 and 16 days (Skead 1960). DEVELOPMENT AND CARE OF YOUNG: eggshells carried away; hatchlings brooded by day by X for 2 days and thereafter less and less; Y feeds them from outset (sometimes having to push his bill under brooding X to do so); from day 3 X starts to feed them too. Young fed by regurgitation, with seed and grit. X sleeps on nest at night for first 8–10 days. Both parents remove gelatinous faecal sacs up to 8th day; thereafter droppings deposited on rim of nest are ignored by parents. In rain or under hot sun, X stands on nest rim and spreads wings to shade young, or she may sit on them even when herself heat-stressed. Young crane for food with necks stretched up and bills wide open; they show fear from day 12 by crouching in bottom of nest. They emerge from nest 2 days before

flying, perch on adjacent twigs and utter begging calls for the first time. Nestling period: 155–185 days. On first flights young flutter from nest and are weak, awkward and vulnerable to predators; they do not keep close together and do not stay near nest for long. Fed by both parents; fledgling begs persistently and waves wings with loose, slow, flapping, rotary motion; young may leap onto incoming parent’s back with impatience (Skead 1960). BREEDING SUCCESS/SURVIVAL: of 78 eggs in one colony, 7 (9%) infertile, 75% hatched and 38 chicks fledged (49% success). Fledglings taken by Fiscal Shrike Lanius collaris (Skead 1960). Not certainly parasitized by any cuckoo. Key References Clement, P. et al. (1993), Ginn, P.J. et al. (1989), Harrison, J.A. et al. (1997), Oatley, T. (2000), Skead, C.J. (1948, 1960), Maclean, G.L. (1993), Milewski, A.V. (1978a, b), Vincent, A.W. (1949).

Plate 31

Serinus serinus (Linnaeus). European Serin. Serin cini.

(Opp. p. 474)

Fringilla Serinus Linnaeus, 1766. Syst. Nat., ed. 12, 1, p. 320; S Europe. Forms a superspecies with S. canaria (Canary Is). Range and Status. Europe east to 32
E, north to S England, S Denmark and Estonia; Mediterranean islands, Turkey, Armenia; Canary Is, NW Africa, N Libya, Lebanon and N Israel. Partial migrant, vacating N and central Europe in winter, moving south into S European and N African breeding range and into N Egypt and Sinai. Resident in and winter visitor to N Morocco, south to Sidi Ifni on coast, High Atlas Mts, and about Taourirt in NE, N Algeria south to Aı¨n Sefra, Laghouat, Messaad and Biskra, N Tunisia south to Beni Abbe`s, Gafsa and Gabe`s, coastal plain of NW Libya east to Tawarga, NE Libya southwest of Al Bayda, and a few probably breed in Egypt in Nile Delta (singing YY in May–June in Cairo, El Khatatba, Shatanuf and Aby Rauwash). Up to 12 pairs bred at Rafah, N Sinai, Apr 1996. Common in NW Africa, in Morocco from coast up to 2700 m (Massif du Toubkal), locally abundant up to 2000 m; in Algeria widespread in open cedar woods up to 2150 m. Sparse winter visit to Cyrenaica and Fezzan (Libya: Cyrene, Barce, Tobruk, Al Adem, Murzuq, Sarir); singing YY at Leptis Magna, Wadi el Kouf, Cyrene, Misratah and Sabratah, Apr 1998 (Massa 1999). Regular in Egypt on N coast and in Nile Delta and Valley, once south to Aswan. Flourishes near human habitation and European range has expanded considerably during 20th century; probably still on the increase. Colonized Tripolitania, Libya, in 1930s after acacias were introduced there. 5–8 million pairs in Spain and Portugal. In Morocco, av. 125 pairs per km2 of forest in breeding season, 18 pairs per km2 in maquis and 36 per km2 of semi-arid Thuja scrub; flocks of 200 and 250 occur in Mar (Ke´nitra, Bargha). Near Gibraltar, winter

Serinus serinus

?

densities of c. 8 birds per km2 in maquis, 2 per km2 in matorral and 63 per km2 in pasture. Description. ADULT Y (breeding): forehead, forecrown, sides of crown and sides of hindneck bright yellow; crown, nape, middle of hindneck and upper mantle olive with fine black streaks; lower mantle and back olive with short, broad, blackish streaks; rump

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c. 30 s; X is fed by regurgitation but rarely Y brings whole insect in bill. Period: 12–16 (usually 12–14) days; 2 neighbouring clutches of 4 eggs were laid concurrently, one hatched in 13 and 14 days, the other in 15 and 16 days (Skead 1960). DEVELOPMENT AND CARE OF YOUNG: eggshells carried away; hatchlings brooded by day by X for 2 days and thereafter less and less; Y feeds them from outset (sometimes having to push his bill under brooding X to do so); from day 3 X starts to feed them too. Young fed by regurgitation, with seed and grit. X sleeps on nest at night for first 8–10 days. Both parents remove gelatinous faecal sacs up to 8th day; thereafter droppings deposited on rim of nest are ignored by parents. In rain or under hot sun, X stands on nest rim and spreads wings to shade young, or she may sit on them even when herself heat-stressed. Young crane for food with necks stretched up and bills wide open; they show fear from day 12 by crouching in bottom of nest. They emerge from nest 2 days before

flying, perch on adjacent twigs and utter begging calls for the first time. Nestling period: 155–185 days. On first flights young flutter from nest and are weak, awkward and vulnerable to predators; they do not keep close together and do not stay near nest for long. Fed by both parents; fledgling begs persistently and waves wings with loose, slow, flapping, rotary motion; young may leap onto incoming parent’s back with impatience (Skead 1960). BREEDING SUCCESS/SURVIVAL: of 78 eggs in one colony, 7 (9%) infertile, 75% hatched and 38 chicks fledged (49% success). Fledglings taken by Fiscal Shrike Lanius collaris (Skead 1960). Not certainly parasitized by any cuckoo. Key References Clement, P. et al. (1993), Ginn, P.J. et al. (1989), Harrison, J.A. et al. (1997), Oatley, T. (2000), Skead, C.J. (1948, 1960), Maclean, G.L. (1993), Milewski, A.V. (1978a, b), Vincent, A.W. (1949).

Plate 31

Serinus serinus (Linnaeus). European Serin. Serin cini.

(Opp. p. 474)

Fringilla Serinus Linnaeus, 1766. Syst. Nat., ed. 12, 1, p. 320; S Europe. Forms a superspecies with S. canaria (Canary Is). Range and Status. Europe east to 32
E, north to S England, S Denmark and Estonia; Mediterranean islands, Turkey, Armenia; Canary Is, NW Africa, N Libya, Lebanon and N Israel. Partial migrant, vacating N and central Europe in winter, moving south into S European and N African breeding range and into N Egypt and Sinai. Resident in and winter visitor to N Morocco, south to Sidi Ifni on coast, High Atlas Mts, and about Taourirt in NE, N Algeria south to Aı¨n Sefra, Laghouat, Messaad and Biskra, N Tunisia south to Beni Abbe`s, Gafsa and Gabe`s, coastal plain of NW Libya east to Tawarga, NE Libya southwest of Al Bayda, and a few probably breed in Egypt in Nile Delta (singing YY in May–June in Cairo, El Khatatba, Shatanuf and Aby Rauwash). Up to 12 pairs bred at Rafah, N Sinai, Apr 1996. Common in NW Africa, in Morocco from coast up to 2700 m (Massif du Toubkal), locally abundant up to 2000 m; in Algeria widespread in open cedar woods up to 2150 m. Sparse winter visit to Cyrenaica and Fezzan (Libya: Cyrene, Barce, Tobruk, Al Adem, Murzuq, Sarir); singing YY at Leptis Magna, Wadi el Kouf, Cyrene, Misratah and Sabratah, Apr 1998 (Massa 1999). Regular in Egypt on N coast and in Nile Delta and Valley, once south to Aswan. Flourishes near human habitation and European range has expanded considerably during 20th century; probably still on the increase. Colonized Tripolitania, Libya, in 1930s after acacias were introduced there. 5–8 million pairs in Spain and Portugal. In Morocco, av. 125 pairs per km2 of forest in breeding season, 18 pairs per km2 in maquis and 36 per km2 of semi-arid Thuja scrub; flocks of 200 and 250 occur in Mar (Ke´nitra, Bargha). Near Gibraltar, winter

Serinus serinus

?

densities of c. 8 birds per km2 in maquis, 2 per km2 in matorral and 63 per km2 in pasture. Description. ADULT Y (breeding): forehead, forecrown, sides of crown and sides of hindneck bright yellow; crown, nape, middle of hindneck and upper mantle olive with fine black streaks; lower mantle and back olive with short, broad, blackish streaks; rump

Serinus serinus bright yellow, uppertail coverts dark grey-brown, with oliveyellow edges; tail feathers blackish with narrow olive-yellow edges. Lores, line through eye, moustachial stripe and ear-coverts dark olive-green; cheeks and line immediately under eye yellow. Chin, throat and breast yellow; belly white; sides of breast and flanks with blackish streaks forming 2–3 long lines; undertailcoverts white. All wing feathers blackish, median coverts tipped pale buff, greater coverts edged olive-green and tipped pale buff, tertials broadly edged olive, secondaries and primaries narrowly edged yellowish; underwing coverts and axillaries pale grey; underside of tail dark grey. Bill dark grey or dark horn-brown; eyes black-brown; legs flesh-brown. ADULT Y (non-breeding): similar, but yellows less bright. ADULT X: like Y, but in place of bright yellow, forehead, sides of head and neck, throat and breast are dull yellowish, streaked with blackish brown. SIZE (Portugal, Spain, Algeria, Italy): wing, Y (n ¼ 22) 68–76 (716), X (n ¼ 6) 66– 69 (675); tail, Y (n ¼ 45) 46–55 (506), X (n ¼ 19) 46–53 (481); bill to skull, Y (n ¼ 24) 98–106 (103), X (n ¼ 7) 99–113 (105); bill to nostril, Y (n ¼ 24) 55–63 (58), X (n ¼ 7) 55–61 (57); tarsus, Y (n ¼ 38) 128–144 (136), X (n ¼ 13) 126–143 (134). WEIGHT: (Tunisia, May) unsexed (n ¼ 15) 105–130 (113). IMMATURE: juv. like ad. X but duller, buffier, without olive, and with streaky rump. Whole upperparts dark grey-brown with broad, pale buff or yellowish buff streaks; sides of neck yellowish buff; tail feathers with broader buff margins than in ad.; underparts very pale buff, whitish towards vent, with thin, dark grey streaks; greater upperwing-coverts and tertials with broader, warmer buff margins than in ad. NESTLING: upperparts and upperwing downy; down pale grey, scant but quite long.

Field Characters. Length 115 cm. Small, streaked greenish finch with short stubby bill and bright yellow rump. Y has bright yellow head and breast with olive crown and face patch; X duller, browner above, head and breast streakier, with much less yellow. Told from Eurasian Siskin Carduelis spinus by lack of yellow in wings and tail. Darker, streakier and shorter-tailed than Syrian Serin S. syriacus. Voice. Tape-recorded (62–73, 76–89, 105, 110–113, B, C). Song rapid, high-pitched jingle of silvery, tinkling notes on one pitch, often continuing for 15 s; sometimes a series of shorter phrases (3–5 s); may contain some buzzy notes, but more musical than jingling song of Corn Bunting Miliaria calandra. Short (05–1 s) drier metallic trill given in flight; also (alarm?) gives nasal, buzzy ‘dzooya’ or upslurred ‘dza-dweee’. For further calls and sonagrams see Cramp and Perrins (1994). General Habits. Inhabits warm, dry lowlands with plenty of scattered trees, and sunny, wooded foothills and mountain slopes, ascending to subalpine zone in Morocco; commonest in suburban gardens with deciduous, coniferous and ornamental trees and large shrubs with open spaces between, and in farmland with tree avenues, tall hedges, copses and plentiful scattered trees; stands of Pinus radiata, Abies alba, Thuja and Juniperus oxycedrus, parkland, orchards, plantations, citrus and olive groves, vineyards, farmyards, kitchen gardens, mosaics of conifers and deciduous trees around open fields, wooded pasture, city parks; in south of range, common in gorges and similar places with plenty green, woody growth, and frequent amongst palms and around villages in semi-desert towards

margins of Sahara. In breeding season Y favours solitary, tall, columnar trees with pointed, spindly, open top, such as cyprus or silver birch, and sings also from wires and rooftops. Usually in pairs; in late summer in family parties; in winter in flocks of 5–15 and occasionally (Israel) hundreds or even thousands. Spends much time on ground, foraging often with other finches, moving with short, rapid, shuffling hops; also forages in shrubs and small trees, where agile, clinging to thinnest, outer branches and bending down to peck energetically at vegetation, pulling catkins and buds to bits. Extracts ripe seeds from Compositae flowering heads, after pulling off petals and bracts. Feeds among tall herbs; in NW Africa commonly feeds on tops of mugwort Artemisia vulgaris (Compositae). Occasionally stands on weak herb stem, holding it to the ground, to peck at seeding head. Bold and confiding. Foraging flocks mobile and restless. Flight buoyant, fast, deeply undulating, rising with rapid wingbeats then falling with closed wings; undulations less regular than in many larger finches, and has skipping or dancing quality. Sings all year. Outside breeding season roosts communally in woods, birds dispersing several km by day to feed. In Morocco resident, breeding population supplemented in winter by visitors from Europe; an altitudinal migrant in Haut Atlas. In autumn, N European birds move to SW and SE, travelling on a broad front; Spain is a major wintering area; mainly Spanish birds migrate by day across Strait of Gibraltar from 2nd week Sept to 1st week Oct; in one year the 6th commonest visible migrant species there, with 15,000 seen; at Settat, NW Morocco, in foothills 70 km inland from Casablanca, main ‘autumn’ passage in Nov and early Dec and spring return from late Feb to early May. At Gibraltar, spring passage from late Mar to mid or late Apr. 1 bird ringed in Spain recovered in Morocco; 1 ringed in Tunisia recovered in Sicily. Winter visitor to Egypt from mid Sept to mid Mar, a few to late Apr. Food. Artemisia vulgaris (Morocco). In Europe, seeds, buds and young flowers of wide variety of plants including spruce, larch, birch, elm, alder, mulberry, nettle, hemp, dock, knotgrass, buckwheat, chickweed, poppy, rose, crucifers, mugwort and other composites, grasses and small-seeded cereals (Cramp and Perrins 1994). Breeding Habits. Monogamous; nests solitarily or in loose groups. Territorial; territory, often of c. 1 ha, used for courtship, nesting and feeding; av. of c. 80 m between territory centres (Germany). Territory boundaries not well defined; territory defended by bird making song-flights, which sometimes go well into territory of adjacent pair. Singing Y has Advertising posture, with upright stance, slightly drooping wings, half-cocked tail, head stretched up and turning from side to side, ruffled throat feathers and trembling body. After a few s of singing next to his mate he flies steeply up to make Song-flight and, with spread tail, ruffled plumage and exaggerated, slow beats of outstretched wings, follows an erratic course 10–20 m above territory, rolling somewhat about his long axis, making a wide arc then descending, slowly at first, followed by a

517

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‘parachute drop’, to alight on same perch next to X, where he continues to sing (Cramp and Perrins 1994). He may then adopt Advertising posture again, quiver wings, bow and strut in front of X, leading either to aerial chasing or to copulation. 1–2 broods. NEST: small and compact, made mainly of fine twigs and stalks, with some strips of bark, roots, grass and moss; neatly and thickly lined with hair, small feathers, plant down and rootlets (Cramp and Perrins 1994). Ext. diam. 85–100, int. diam. 45–70, ext. depth 45–50, int. depth 30– 40. Built commonly in citrus, spruce, lime, yew, pine, larch, elm, elder, apple and pear trees, generally about two thirds out from trunk; av. height (131 nests, central Europe) 29 m, max. 6 m. X prospects for nest-site, accompanied constantly by Y. Nest built mainly or entirely by X, taking 4–11 days; Y sometimes brings material; X and Y gather material almost entirely from ground. EGGS: 3–5 (Europe), mainly 3–4, av. (185 clutches, E Spain) 37. Sub-elliptical, smooth, slightly glossy; bluish or greenish white, sparsely spotted and streaked with rustred and purple-brown, mainly at broad end. SIZE: (n ¼ 337) 144–178  110–133 (161  120). Weight c. 121 g.

LAYING DATES: NW Africa, from mid Feb; Morocco Mar–May (earliest, nestlings Berkane late Feb, latest a clutch on 26th May). INCUBATION: by X, sitting tightly and for lengthy periods; starts with penultimate or last egg; X courtship-fed by Y late in incubation period. Period (n ¼ 9) 127 days. DEVELOPMENT AND CARE OF YOUNG: young brooded only by X. Y makes feeding visits to nest, regurgitating food to X who delivers it to young. Later, young fed and cared for by Y and X equally. X eats faecal sacs in 1st week; from 7– 9 days young defaecate on rim of nest, which becomes heavily encrusted. Nestling period (n ¼ 14) 13–18 days, av. 152 days; brood leaves nest over 1–2 days, dispersing into nest tree. Young independent c. 9 days after leaving nest. BREEDING SUCCESS/SURVIVAL: in E Spain, of 290 eggs, 67% hatched and 33% produced flying young; losses due to predation (by Magpie Pica pica, other corvids, cats), farming activities, infertility, nest damage, poor weather and starvation of young (Cramp and Perrins 1994).

Key References Clement, P. et al. (1993), Cramp, S. and Perrins, C.M. (1994), Isenmann, P. and Moali, A. (2000).

Plate 31

Serinus syriacus Bonaparte. Syrian Serin; Tristram’s Serin. Serin syriaque.

(Opp. p. 474)

Serinus syriacus Bonaparte, 1850. Consp. Avium 1, p. 523; ‘‘ex As. occ. Bischerra’’. Endemic to mountains in S Syria, Lebanon, N Israel (Mt Hermon, at 900–1750 m) and Jordan (Dana Nat. Res.: Khoury 2001); migrant, wintering from Syria to S Israel, Sinai (e.g. 8–10, Ras El Naqb at southern tip, Jan 1997), Jordan and Iraq; in Sinai a scarce winter visitor and passage migrant, but hundreds in some years. Vagrant, Egypt: 1 Helwan, Dec 1919, 1 Bahig, Nov 1969, pair (the Y singing) Tuna el Gebel, Feb 1979 (Short and Horne 1981), 2 east of Cairo, Sept 1994 (Bull. Afr. Bird Club 1995, 2, 123). Adult Y: bright yellow forehead, lores, around eyes, chin, throat and upper breast; rest of head pale grey; mantle pale grey, dark-streaked; rump and sides of rather long tail yellow; wings black but scapulars, lesser, median and greater wingcoverts yellow, tertials broadly and secondaries and primaries narrowly edged yellow; underparts unstreaked, belly whitish; bill and eyes blackish, legs brown. Adult X similar but yellows less intense and crown, mantle, lesser wingcoverts and flanks streaky. Ad. in flight looks yellowish, with yellow rump and upperwing-coverts in strong contrast with blackish tail, primaries and secondary bases. Paler and yellower than European Serin S. serinus, more uniform, less streaky; tail and wings with some yellow, less contrasting yellow rump. Juv. more distinctive: wings and tail variegated, blackish, and buffy where ad. is yellow; rest of plumage rather plain light buffy brown, with slight streaks on mantle and breast; lower mandible horn-coloured. Length 12–14 cm; wing, Y 75–80 (778), X 71–77 (746); tail Y 52–57 (552), X 52–56 (534); weight, Y 10–14 (114), X 11–12 (113). Breeds in mountain cedar and juniper woods but in winter occurs in low-lying semi-desert acacia scrub and Artemisia steppe with scattered trees and tall shrubs (Khoury 2000), wadis, orchards and gardens. Eats small seeds, taken from ground and in trees and bushes, moves freely between treetops and ground. In winter forms flocks of up to 600 birds. Bounding flight. Flight call ‘tsirr’ or ‘tirrrh’, drier and less ringing than that of European Serin.

Plate 31

Serinus citrinella (Pallas). Citril Finch. Venturon montagnard.

(Opp. p. 474)

Fringilla citrinella Pallas, 1764. In Vroeg, Cat. raisonne´, Oiseaux, adumbrat., p. 3; ‘‘Holland’’. Confined to SW European mountains: S Germany, W Austria, NE France, N Italy, N and central Spain (S. c. citrinella). Vertical and short-distance migrant; rare vagrant north to Poland, west to Britain and south to Morocco and Algeria. 1, Ceuta, extreme N Morocco, Mar 1991 and another there Apr 1991; 2 El Kala, Algeria, in 19th century. African vagrants are presumably this species but might possibly have been the newly separted S. corsicana, sedentary on Corsica and Sardinia (Isenmann and Moali 2000). Adult Y has greenish yellow face (forehead, around eyes, chin and throat), bluish grey crown, hindneck and ear-coverts, olive mantle, yellowish rump, blackish tail, greenish yellow underparts with narrow grey band across upper breast and

518

FRINGILLIDAE

‘parachute drop’, to alight on same perch next to X, where he continues to sing (Cramp and Perrins 1994). He may then adopt Advertising posture again, quiver wings, bow and strut in front of X, leading either to aerial chasing or to copulation. 1–2 broods. NEST: small and compact, made mainly of fine twigs and stalks, with some strips of bark, roots, grass and moss; neatly and thickly lined with hair, small feathers, plant down and rootlets (Cramp and Perrins 1994). Ext. diam. 85–100, int. diam. 45–70, ext. depth 45–50, int. depth 30– 40. Built commonly in citrus, spruce, lime, yew, pine, larch, elm, elder, apple and pear trees, generally about two thirds out from trunk; av. height (131 nests, central Europe) 29 m, max. 6 m. X prospects for nest-site, accompanied constantly by Y. Nest built mainly or entirely by X, taking 4–11 days; Y sometimes brings material; X and Y gather material almost entirely from ground. EGGS: 3–5 (Europe), mainly 3–4, av. (185 clutches, E Spain) 37. Sub-elliptical, smooth, slightly glossy; bluish or greenish white, sparsely spotted and streaked with rustred and purple-brown, mainly at broad end. SIZE: (n ¼ 337) 144–178  110–133 (161  120). Weight c. 121 g.

LAYING DATES: NW Africa, from mid Feb; Morocco Mar–May (earliest, nestlings Berkane late Feb, latest a clutch on 26th May). INCUBATION: by X, sitting tightly and for lengthy periods; starts with penultimate or last egg; X courtship-fed by Y late in incubation period. Period (n ¼ 9) 127 days. DEVELOPMENT AND CARE OF YOUNG: young brooded only by X. Y makes feeding visits to nest, regurgitating food to X who delivers it to young. Later, young fed and cared for by Y and X equally. X eats faecal sacs in 1st week; from 7– 9 days young defaecate on rim of nest, which becomes heavily encrusted. Nestling period (n ¼ 14) 13–18 days, av. 152 days; brood leaves nest over 1–2 days, dispersing into nest tree. Young independent c. 9 days after leaving nest. BREEDING SUCCESS/SURVIVAL: in E Spain, of 290 eggs, 67% hatched and 33% produced flying young; losses due to predation (by Magpie Pica pica, other corvids, cats), farming activities, infertility, nest damage, poor weather and starvation of young (Cramp and Perrins 1994).

Key References Clement, P. et al. (1993), Cramp, S. and Perrins, C.M. (1994), Isenmann, P. and Moali, A. (2000).

Plate 31

Serinus syriacus Bonaparte. Syrian Serin; Tristram’s Serin. Serin syriaque.

(Opp. p. 474)

Serinus syriacus Bonaparte, 1850. Consp. Avium 1, p. 523; ‘‘ex As. occ. Bischerra’’. Endemic to mountains in S Syria, Lebanon, N Israel (Mt Hermon, at 900–1750 m) and Jordan (Dana Nat. Res.: Khoury 2001); migrant, wintering from Syria to S Israel, Sinai (e.g. 8–10, Ras El Naqb at southern tip, Jan 1997), Jordan and Iraq; in Sinai a scarce winter visitor and passage migrant, but hundreds in some years. Vagrant, Egypt: 1 Helwan, Dec 1919, 1 Bahig, Nov 1969, pair (the Y singing) Tuna el Gebel, Feb 1979 (Short and Horne 1981), 2 east of Cairo, Sept 1994 (Bull. Afr. Bird Club 1995, 2, 123). Adult Y: bright yellow forehead, lores, around eyes, chin, throat and upper breast; rest of head pale grey; mantle pale grey, dark-streaked; rump and sides of rather long tail yellow; wings black but scapulars, lesser, median and greater wingcoverts yellow, tertials broadly and secondaries and primaries narrowly edged yellow; underparts unstreaked, belly whitish; bill and eyes blackish, legs brown. Adult X similar but yellows less intense and crown, mantle, lesser wingcoverts and flanks streaky. Ad. in flight looks yellowish, with yellow rump and upperwing-coverts in strong contrast with blackish tail, primaries and secondary bases. Paler and yellower than European Serin S. serinus, more uniform, less streaky; tail and wings with some yellow, less contrasting yellow rump. Juv. more distinctive: wings and tail variegated, blackish, and buffy where ad. is yellow; rest of plumage rather plain light buffy brown, with slight streaks on mantle and breast; lower mandible horn-coloured. Length 12–14 cm; wing, Y 75–80 (778), X 71–77 (746); tail Y 52–57 (552), X 52–56 (534); weight, Y 10–14 (114), X 11–12 (113). Breeds in mountain cedar and juniper woods but in winter occurs in low-lying semi-desert acacia scrub and Artemisia steppe with scattered trees and tall shrubs (Khoury 2000), wadis, orchards and gardens. Eats small seeds, taken from ground and in trees and bushes, moves freely between treetops and ground. In winter forms flocks of up to 600 birds. Bounding flight. Flight call ‘tsirr’ or ‘tirrrh’, drier and less ringing than that of European Serin.

Plate 31

Serinus citrinella (Pallas). Citril Finch. Venturon montagnard.

(Opp. p. 474)

Fringilla citrinella Pallas, 1764. In Vroeg, Cat. raisonne´, Oiseaux, adumbrat., p. 3; ‘‘Holland’’. Confined to SW European mountains: S Germany, W Austria, NE France, N Italy, N and central Spain (S. c. citrinella). Vertical and short-distance migrant; rare vagrant north to Poland, west to Britain and south to Morocco and Algeria. 1, Ceuta, extreme N Morocco, Mar 1991 and another there Apr 1991; 2 El Kala, Algeria, in 19th century. African vagrants are presumably this species but might possibly have been the newly separted S. corsicana, sedentary on Corsica and Sardinia (Isenmann and Moali 2000). Adult Y has greenish yellow face (forehead, around eyes, chin and throat), bluish grey crown, hindneck and ear-coverts, olive mantle, yellowish rump, blackish tail, greenish yellow underparts with narrow grey band across upper breast and

518

FRINGILLIDAE

‘parachute drop’, to alight on same perch next to X, where he continues to sing (Cramp and Perrins 1994). He may then adopt Advertising posture again, quiver wings, bow and strut in front of X, leading either to aerial chasing or to copulation. 1–2 broods. NEST: small and compact, made mainly of fine twigs and stalks, with some strips of bark, roots, grass and moss; neatly and thickly lined with hair, small feathers, plant down and rootlets (Cramp and Perrins 1994). Ext. diam. 85–100, int. diam. 45–70, ext. depth 45–50, int. depth 30– 40. Built commonly in citrus, spruce, lime, yew, pine, larch, elm, elder, apple and pear trees, generally about two thirds out from trunk; av. height (131 nests, central Europe) 29 m, max. 6 m. X prospects for nest-site, accompanied constantly by Y. Nest built mainly or entirely by X, taking 4–11 days; Y sometimes brings material; X and Y gather material almost entirely from ground. EGGS: 3–5 (Europe), mainly 3–4, av. (185 clutches, E Spain) 37. Sub-elliptical, smooth, slightly glossy; bluish or greenish white, sparsely spotted and streaked with rustred and purple-brown, mainly at broad end. SIZE: (n ¼ 337) 144–178  110–133 (161  120). Weight c. 121 g.

LAYING DATES: NW Africa, from mid Feb; Morocco Mar–May (earliest, nestlings Berkane late Feb, latest a clutch on 26th May). INCUBATION: by X, sitting tightly and for lengthy periods; starts with penultimate or last egg; X courtship-fed by Y late in incubation period. Period (n ¼ 9) 127 days. DEVELOPMENT AND CARE OF YOUNG: young brooded only by X. Y makes feeding visits to nest, regurgitating food to X who delivers it to young. Later, young fed and cared for by Y and X equally. X eats faecal sacs in 1st week; from 7– 9 days young defaecate on rim of nest, which becomes heavily encrusted. Nestling period (n ¼ 14) 13–18 days, av. 152 days; brood leaves nest over 1–2 days, dispersing into nest tree. Young independent c. 9 days after leaving nest. BREEDING SUCCESS/SURVIVAL: in E Spain, of 290 eggs, 67% hatched and 33% produced flying young; losses due to predation (by Magpie Pica pica, other corvids, cats), farming activities, infertility, nest damage, poor weather and starvation of young (Cramp and Perrins 1994).

Key References Clement, P. et al. (1993), Cramp, S. and Perrins, C.M. (1994), Isenmann, P. and Moali, A. (2000).

Plate 31

Serinus syriacus Bonaparte. Syrian Serin; Tristram’s Serin. Serin syriaque.

(Opp. p. 474)

Serinus syriacus Bonaparte, 1850. Consp. Avium 1, p. 523; ‘‘ex As. occ. Bischerra’’. Endemic to mountains in S Syria, Lebanon, N Israel (Mt Hermon, at 900–1750 m) and Jordan (Dana Nat. Res.: Khoury 2001); migrant, wintering from Syria to S Israel, Sinai (e.g. 8–10, Ras El Naqb at southern tip, Jan 1997), Jordan and Iraq; in Sinai a scarce winter visitor and passage migrant, but hundreds in some years. Vagrant, Egypt: 1 Helwan, Dec 1919, 1 Bahig, Nov 1969, pair (the Y singing) Tuna el Gebel, Feb 1979 (Short and Horne 1981), 2 east of Cairo, Sept 1994 (Bull. Afr. Bird Club 1995, 2, 123). Adult Y: bright yellow forehead, lores, around eyes, chin, throat and upper breast; rest of head pale grey; mantle pale grey, dark-streaked; rump and sides of rather long tail yellow; wings black but scapulars, lesser, median and greater wingcoverts yellow, tertials broadly and secondaries and primaries narrowly edged yellow; underparts unstreaked, belly whitish; bill and eyes blackish, legs brown. Adult X similar but yellows less intense and crown, mantle, lesser wingcoverts and flanks streaky. Ad. in flight looks yellowish, with yellow rump and upperwing-coverts in strong contrast with blackish tail, primaries and secondary bases. Paler and yellower than European Serin S. serinus, more uniform, less streaky; tail and wings with some yellow, less contrasting yellow rump. Juv. more distinctive: wings and tail variegated, blackish, and buffy where ad. is yellow; rest of plumage rather plain light buffy brown, with slight streaks on mantle and breast; lower mandible horn-coloured. Length 12–14 cm; wing, Y 75–80 (778), X 71–77 (746); tail Y 52–57 (552), X 52–56 (534); weight, Y 10–14 (114), X 11–12 (113). Breeds in mountain cedar and juniper woods but in winter occurs in low-lying semi-desert acacia scrub and Artemisia steppe with scattered trees and tall shrubs (Khoury 2000), wadis, orchards and gardens. Eats small seeds, taken from ground and in trees and bushes, moves freely between treetops and ground. In winter forms flocks of up to 600 birds. Bounding flight. Flight call ‘tsirr’ or ‘tirrrh’, drier and less ringing than that of European Serin.

Plate 31

Serinus citrinella (Pallas). Citril Finch. Venturon montagnard.

(Opp. p. 474)

Fringilla citrinella Pallas, 1764. In Vroeg, Cat. raisonne´, Oiseaux, adumbrat., p. 3; ‘‘Holland’’. Confined to SW European mountains: S Germany, W Austria, NE France, N Italy, N and central Spain (S. c. citrinella). Vertical and short-distance migrant; rare vagrant north to Poland, west to Britain and south to Morocco and Algeria. 1, Ceuta, extreme N Morocco, Mar 1991 and another there Apr 1991; 2 El Kala, Algeria, in 19th century. African vagrants are presumably this species but might possibly have been the newly separted S. corsicana, sedentary on Corsica and Sardinia (Isenmann and Moali 2000). Adult Y has greenish yellow face (forehead, around eyes, chin and throat), bluish grey crown, hindneck and ear-coverts, olive mantle, yellowish rump, blackish tail, greenish yellow underparts with narrow grey band across upper breast and

Serinus nigriceps

519

grey sides of breast, and black wings boldly marked with yellow (lesser coverts; broad tips to secondary-coverts forming a short wing-bar; outer edges of tertials, secondaries and primaries); upper mandible blackish, lower brownish, eyes black, legs brown. Adult X similar but less yellowish, mantle streakier, and yellow wing bars much narrower. Juv. not safely distinguishable in field from juv. Eurasian Siskin Carduelis spinus, q.v. Length 115–12 cm; wing, Y 75–82 (782), X 75–79 (762); tail Y 51–58 (544), X 50–58 (538); weight 115–14, monthly averages from 122 to 129. Recalls Carduelis finches in form and behaviour, rather than serins Serinus. Inhabits spruce, larch and pine forests. Perches in trees and shrubs, also on wires and rocky ground. Gregarious in winter, often flocking with Common Redpolls C. flammea and Eurasian Siskins C. spinus. Forages mainly on ground, also in outer branches. Twittering song more musical than Eurasian Siskin; flight call a nasal, creaky, metallic ‘di’, ‘tweck’ or ‘chiht’, sometimes in rapid series.

Serinus nigriceps Ru ¨ ppell. Ethiopian Siskin; Black-headed Siskin. Serin a` te ˆte noire.

Plate 31

Serinus nigriceps Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 96, pl. 34, fig. 1; Simen Prov., 10,000 ft., Abyssinia.

(Opp. p. 474)

Range and Status. Endemic resident, Ethiopian highlands at 1800–4100 m (rarely below 1800 m), near eastern flank of W Highlands and in SE Highlands. Locally common to abundant, less numerous in farmland than formerly (Ash and Gullick 1989). Description. ADULT Y: head and neck brownish black, browner on ear-coverts, cheeks, chin and throat, extending to sooty brown bib on centre of upper breast. Rest of upperparts dull yellowish olive-green, sullied brownish, brighter and more yellow on lateral scapulars, rump and short uppertail-coverts. Tail black, feathers narrowly fringed buffish white or pale buffish yellow when fresh. Sides of breast and flanks dull olive-green, variably tinged with brown, merging with olive-yellow lower breast and upper belly and pale yellow or yellowish buff lower belly and undertailcoverts. Remiges blackish, primaries and secondaries finely edged buff and tipped greyish white, tertials broadly edged and tipped pale greenish grey; alula and primary coverts blackish, finely fringed buff; greater coverts blackish, edged and broadly tipped pale greenish yellow to form prominent wingbar; median coverts blackish, tipped pale greenish yellow; lesser coverts olive-yellow. Underwing-coverts and axillaries dull olive-green. Bill black or dark brown, shape varying somewhat (A); eyes brown; legs black or blackish brown. ADULT X: top of head sooty brown, crown usually invaded by dull olive-green streaking; hindneck to mantle, back and scapulars dull olive-green, tinged brownish and with diffuse dark brown streaking, greener on lateral scapulars; rump and short uppertail-coverts yellowish olive-green; long uppertailcoverts dull olive-green tipped pale grey. Lores to behind eye sooty brown; ear-coverts, cheeks to chin, throat and centre of upper breast dull greyish or brownish olive-green; sides of breast and flanks dull yellowish olive with diffuse dusky olive streaking; lower breast and upper belly pale olive-yellow, merging with buffish white lower belly and undertail-coverts. Tail and wings as ad. Y, but black feathers tinged brown, yellow tips to greater and median coverts paler and narrower, lesser coverts tinged brown. SIZE (10 YY, 4 XX): wing, Y 76–78 (769), X 73–77 (750); tail, Y 52–60 (542), X 50–54 (520); bill, Y 125–135 (130), X 12–135 (124); tarsus, Y 16–17 (167), X 155–17 (165). IMMATURE: juv. similar to ad. X, but more brownish above, entire head and upperparts with soft dark brown streaking; underparts dull yellowish or buffish white, heavily streaked; greater and median coverts tipped pale buff. TAXONOMIC NOTE: often placed in genus Carduelis.

Field Characters. Length 11–125 cm. Y with black head, bright yellow scapulars and broad pale wing-bars unmistakable (Oriole-Finch Linurgus olivaceus does not reach Ethiopia). X and juv. told from African Citril S. citrinelloides and Yellow-crowned Canary S. flavivertex by sooty brown

Serinus nigriceps

?

forehead, crown and lores without yellow supercilium; from African Citril by broader wing-bars on darker wing and lighter, more diffuse streaking; juv. from juv. Yellowcrowned Canary by olive-green rather than buffy-brown upperparts.

Voice. Tape-recorded (109, SM). Song a pleasant series of musical trills in frequently-repeated phrases, recalling Goldfinch C. carduelis but without the buzzing quality (Clement et al. 1993). Noisy chattering calls include twitters, chirps, buzzy ‘trrrr’ and hard ‘chit-chit-chit’.

Serinus nigriceps

519

grey sides of breast, and black wings boldly marked with yellow (lesser coverts; broad tips to secondary-coverts forming a short wing-bar; outer edges of tertials, secondaries and primaries); upper mandible blackish, lower brownish, eyes black, legs brown. Adult X similar but less yellowish, mantle streakier, and yellow wing bars much narrower. Juv. not safely distinguishable in field from juv. Eurasian Siskin Carduelis spinus, q.v. Length 115–12 cm; wing, Y 75–82 (782), X 75–79 (762); tail Y 51–58 (544), X 50–58 (538); weight 115–14, monthly averages from 122 to 129. Recalls Carduelis finches in form and behaviour, rather than serins Serinus. Inhabits spruce, larch and pine forests. Perches in trees and shrubs, also on wires and rocky ground. Gregarious in winter, often flocking with Common Redpolls C. flammea and Eurasian Siskins C. spinus. Forages mainly on ground, also in outer branches. Twittering song more musical than Eurasian Siskin; flight call a nasal, creaky, metallic ‘di’, ‘tweck’ or ‘chiht’, sometimes in rapid series.

Serinus nigriceps Ru ¨ ppell. Ethiopian Siskin; Black-headed Siskin. Serin a` te ˆte noire.

Plate 31

Serinus nigriceps Ru ¨ ppell, 1840. Neue Wirbelt., Vo ¨gel, p. 96, pl. 34, fig. 1; Simen Prov., 10,000 ft., Abyssinia.

(Opp. p. 474)

Range and Status. Endemic resident, Ethiopian highlands at 1800–4100 m (rarely below 1800 m), near eastern flank of W Highlands and in SE Highlands. Locally common to abundant, less numerous in farmland than formerly (Ash and Gullick 1989). Description. ADULT Y: head and neck brownish black, browner on ear-coverts, cheeks, chin and throat, extending to sooty brown bib on centre of upper breast. Rest of upperparts dull yellowish olive-green, sullied brownish, brighter and more yellow on lateral scapulars, rump and short uppertail-coverts. Tail black, feathers narrowly fringed buffish white or pale buffish yellow when fresh. Sides of breast and flanks dull olive-green, variably tinged with brown, merging with olive-yellow lower breast and upper belly and pale yellow or yellowish buff lower belly and undertailcoverts. Remiges blackish, primaries and secondaries finely edged buff and tipped greyish white, tertials broadly edged and tipped pale greenish grey; alula and primary coverts blackish, finely fringed buff; greater coverts blackish, edged and broadly tipped pale greenish yellow to form prominent wingbar; median coverts blackish, tipped pale greenish yellow; lesser coverts olive-yellow. Underwing-coverts and axillaries dull olive-green. Bill black or dark brown, shape varying somewhat (A); eyes brown; legs black or blackish brown. ADULT X: top of head sooty brown, crown usually invaded by dull olive-green streaking; hindneck to mantle, back and scapulars dull olive-green, tinged brownish and with diffuse dark brown streaking, greener on lateral scapulars; rump and short uppertail-coverts yellowish olive-green; long uppertailcoverts dull olive-green tipped pale grey. Lores to behind eye sooty brown; ear-coverts, cheeks to chin, throat and centre of upper breast dull greyish or brownish olive-green; sides of breast and flanks dull yellowish olive with diffuse dusky olive streaking; lower breast and upper belly pale olive-yellow, merging with buffish white lower belly and undertail-coverts. Tail and wings as ad. Y, but black feathers tinged brown, yellow tips to greater and median coverts paler and narrower, lesser coverts tinged brown. SIZE (10 YY, 4 XX): wing, Y 76–78 (769), X 73–77 (750); tail, Y 52–60 (542), X 50–54 (520); bill, Y 125–135 (130), X 12–135 (124); tarsus, Y 16–17 (167), X 155–17 (165). IMMATURE: juv. similar to ad. X, but more brownish above, entire head and upperparts with soft dark brown streaking; underparts dull yellowish or buffish white, heavily streaked; greater and median coverts tipped pale buff. TAXONOMIC NOTE: often placed in genus Carduelis.

Field Characters. Length 11–125 cm. Y with black head, bright yellow scapulars and broad pale wing-bars unmistakable (Oriole-Finch Linurgus olivaceus does not reach Ethiopia). X and juv. told from African Citril S. citrinelloides and Yellow-crowned Canary S. flavivertex by sooty brown

Serinus nigriceps

?

forehead, crown and lores without yellow supercilium; from African Citril by broader wing-bars on darker wing and lighter, more diffuse streaking; juv. from juv. Yellowcrowned Canary by olive-green rather than buffy-brown upperparts.

Voice. Tape-recorded (109, SM). Song a pleasant series of musical trills in frequently-repeated phrases, recalling Goldfinch C. carduelis but without the buzzing quality (Clement et al. 1993). Noisy chattering calls include twitters, chirps, buzzy ‘trrrr’ and hard ‘chit-chit-chit’.

520

FRINGILLIDAE

General Habits. Frequents moorland, grassland and open montane forest with giant lobelia, alchemillas (Alchemilla johnstoni, A. ellenbecki, A. abyssinica), giant heath and main food plants St John’s wort Hypericum and Hagenia; thickets of Euphorbia depauperata (Bale Mts); partly cultivated fields (Ankober, Addis Ababa: Francis and Shirihai 1999); sometimes around sheep folds and human habitation. In pairs when breeding but gregarious at other times, mainly in flocks of up to 10 birds; flocks sometimes of up to 100 birds or more. Rather flighty and restless. Perches boldly on twigs; slim; pattern-winged; long-winged like other high-altitude Ethiopian serins, these characters making it look more like a Carduelis finch than a typical lowland Serinus (B); when singing, posture and actions recall European Goldfinch C. carduelis or Common Linnet C. cannabina (Clement et al. 1993). Forages on ground, and by clinging to low bushes and tall herbs, taking seeds in situ. Food. Seeds, mainly of Hypericum lanceolatum (Guttiferae) and Hagenia abyssinica (Rosaceae). In captivity eats seeds of Panicum maximum, Capsella bursapastoris, Sonchus sp. and Stellaria media and buds of Oxalis pescaprae. Breeding Habits. Solitary nester; monogamous; territorial. NEST: a neat, well-made compact cup, composed of fine grasses and rootlets, lined with very fine grasses or grasslike matter, sometimes mixed with a little down; markedly deeper than nest of e.g. Cape Canary S. canicollis; placed low down in bush or on hanging branch.

EGGS: 2–3. Eggs bluish white, finely spotted with brown or with a few brown spots. SIZE: c. 18  13. LAYING DATES: May, June, Sept, Oct. INCUBATION: period 14 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: nestling period 17– 18 days (in captivity). BREEDING SUCCESS/SURVIVAL: no information.

Key References Brickell, N. and Konigkramer, T. (1997), Clement, P. et al. (1993).

Plate 31

Serinus pusillus (Pallas). Red-fronted Serin. Serin a ` front d’or.

(Opp. p. 474)

Passer pusillus Pallas, 1811. Zoogr. Rosso-Asiat., 2, p. 28; near Caucasus and Caspian Sea. S Turkey, Transcaucasia, Iran, mountains of central Asia east to Nepal, SW Tibet, Pamirs, Tien Shan, Tarbagatai and Sinkiang. Altitudinal and short-distance migrant; winter visitor to N Lebanon; vagrant Austria, Greece, Cyprus, Syria, Jordan and Egypt. A X at Bahig, Egypt, in Mar 1969 (several other records in Egyptian Sinai). A small, dark, stubby finch, sexes alike, ad. readily identified by black or blackish head and upper breast, scarlet forehead and warm ochre rump; rest of plumage strongly variegated with pale buff, rufescent mid-buffs and blackish browns; tertials, uppertail-coverts and tail feathers blackish with whitish buff edges; secondary coverts tipped buff, making a short, narrow but quite conspicuous wing-bar. Soft parts blackish. Imm. has dark brown forehead, plain brown or rufescent brown cheeks and ear-coverts, and slightly dark-streaked, buffy breast and flanks; base of lower mandible brownish. Length 12–13 cm; wing Y 72–77 (742), X 68–75 (722), tail Y 50–61 (557), X 49–57 (532); weight (Asia Minor) 8 YY 11–13 (119), 1 X 12. Keeps mainly above 2000 m but some descend to coasts in winter. Sometimes in leafy vegetation but perches mainly on dead branches, rocks and stones. Very agile in trees, feeding upside down at catkins. Forages, often with other finches, on open ground, eating mainly small seeds taken from surface, from low herbs and grasses by jumping up and fluttering; clings to stem to pick seeds from head of plant, bird’s weight sometimes pressing stem to the ground. Flies with rapid wingbeats and short, steep bounds, often trilling with a short, high-pitched, rippling twitter.

Genus Pseudochloroptila Wolters An endemic superspecies of 2 South African siskins or serin-like finches, YY with rufescent brown back, plain grey rump, unstreaked yellowish underparts, and bill finer (less stubby) than in Serinus. Y of one species quite large; Y of other species has black and white tips to remiges and rectrices. Courtship behaviour includes unique whirring flight by Y in front of X; nest placed in crevice or concealed on ground; nest constructed by both sexes; eggs pure white; incubation period (16 days) and fledgling period (16–18 days) each 2 days longer than in Serinus; nestling gapes uniformly red. Begging calls resemble those of Serinus scotops.

520

FRINGILLIDAE

General Habits. Frequents moorland, grassland and open montane forest with giant lobelia, alchemillas (Alchemilla johnstoni, A. ellenbecki, A. abyssinica), giant heath and main food plants St John’s wort Hypericum and Hagenia; thickets of Euphorbia depauperata (Bale Mts); partly cultivated fields (Ankober, Addis Ababa: Francis and Shirihai 1999); sometimes around sheep folds and human habitation. In pairs when breeding but gregarious at other times, mainly in flocks of up to 10 birds; flocks sometimes of up to 100 birds or more. Rather flighty and restless. Perches boldly on twigs; slim; pattern-winged; long-winged like other high-altitude Ethiopian serins, these characters making it look more like a Carduelis finch than a typical lowland Serinus (B); when singing, posture and actions recall European Goldfinch C. carduelis or Common Linnet C. cannabina (Clement et al. 1993). Forages on ground, and by clinging to low bushes and tall herbs, taking seeds in situ. Food. Seeds, mainly of Hypericum lanceolatum (Guttiferae) and Hagenia abyssinica (Rosaceae). In captivity eats seeds of Panicum maximum, Capsella bursapastoris, Sonchus sp. and Stellaria media and buds of Oxalis pescaprae. Breeding Habits. Solitary nester; monogamous; territorial. NEST: a neat, well-made compact cup, composed of fine grasses and rootlets, lined with very fine grasses or grasslike matter, sometimes mixed with a little down; markedly deeper than nest of e.g. Cape Canary S. canicollis; placed low down in bush or on hanging branch.

EGGS: 2–3. Eggs bluish white, finely spotted with brown or with a few brown spots. SIZE: c. 18  13. LAYING DATES: May, June, Sept, Oct. INCUBATION: period 14 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: nestling period 17– 18 days (in captivity). BREEDING SUCCESS/SURVIVAL: no information.

Key References Brickell, N. and Konigkramer, T. (1997), Clement, P. et al. (1993).

Plate 31

Serinus pusillus (Pallas). Red-fronted Serin. Serin a ` front d’or.

(Opp. p. 474)

Passer pusillus Pallas, 1811. Zoogr. Rosso-Asiat., 2, p. 28; near Caucasus and Caspian Sea. S Turkey, Transcaucasia, Iran, mountains of central Asia east to Nepal, SW Tibet, Pamirs, Tien Shan, Tarbagatai and Sinkiang. Altitudinal and short-distance migrant; winter visitor to N Lebanon; vagrant Austria, Greece, Cyprus, Syria, Jordan and Egypt. A X at Bahig, Egypt, in Mar 1969 (several other records in Egyptian Sinai). A small, dark, stubby finch, sexes alike, ad. readily identified by black or blackish head and upper breast, scarlet forehead and warm ochre rump; rest of plumage strongly variegated with pale buff, rufescent mid-buffs and blackish browns; tertials, uppertail-coverts and tail feathers blackish with whitish buff edges; secondary coverts tipped buff, making a short, narrow but quite conspicuous wing-bar. Soft parts blackish. Imm. has dark brown forehead, plain brown or rufescent brown cheeks and ear-coverts, and slightly dark-streaked, buffy breast and flanks; base of lower mandible brownish. Length 12–13 cm; wing Y 72–77 (742), X 68–75 (722), tail Y 50–61 (557), X 49–57 (532); weight (Asia Minor) 8 YY 11–13 (119), 1 X 12. Keeps mainly above 2000 m but some descend to coasts in winter. Sometimes in leafy vegetation but perches mainly on dead branches, rocks and stones. Very agile in trees, feeding upside down at catkins. Forages, often with other finches, on open ground, eating mainly small seeds taken from surface, from low herbs and grasses by jumping up and fluttering; clings to stem to pick seeds from head of plant, bird’s weight sometimes pressing stem to the ground. Flies with rapid wingbeats and short, steep bounds, often trilling with a short, high-pitched, rippling twitter.

Genus Pseudochloroptila Wolters An endemic superspecies of 2 South African siskins or serin-like finches, YY with rufescent brown back, plain grey rump, unstreaked yellowish underparts, and bill finer (less stubby) than in Serinus. Y of one species quite large; Y of other species has black and white tips to remiges and rectrices. Courtship behaviour includes unique whirring flight by Y in front of X; nest placed in crevice or concealed on ground; nest constructed by both sexes; eggs pure white; incubation period (16 days) and fledgling period (16–18 days) each 2 days longer than in Serinus; nestling gapes uniformly red. Begging calls resemble those of Serinus scotops.

520

FRINGILLIDAE

General Habits. Frequents moorland, grassland and open montane forest with giant lobelia, alchemillas (Alchemilla johnstoni, A. ellenbecki, A. abyssinica), giant heath and main food plants St John’s wort Hypericum and Hagenia; thickets of Euphorbia depauperata (Bale Mts); partly cultivated fields (Ankober, Addis Ababa: Francis and Shirihai 1999); sometimes around sheep folds and human habitation. In pairs when breeding but gregarious at other times, mainly in flocks of up to 10 birds; flocks sometimes of up to 100 birds or more. Rather flighty and restless. Perches boldly on twigs; slim; pattern-winged; long-winged like other high-altitude Ethiopian serins, these characters making it look more like a Carduelis finch than a typical lowland Serinus (B); when singing, posture and actions recall European Goldfinch C. carduelis or Common Linnet C. cannabina (Clement et al. 1993). Forages on ground, and by clinging to low bushes and tall herbs, taking seeds in situ. Food. Seeds, mainly of Hypericum lanceolatum (Guttiferae) and Hagenia abyssinica (Rosaceae). In captivity eats seeds of Panicum maximum, Capsella bursapastoris, Sonchus sp. and Stellaria media and buds of Oxalis pescaprae. Breeding Habits. Solitary nester; monogamous; territorial. NEST: a neat, well-made compact cup, composed of fine grasses and rootlets, lined with very fine grasses or grasslike matter, sometimes mixed with a little down; markedly deeper than nest of e.g. Cape Canary S. canicollis; placed low down in bush or on hanging branch.

EGGS: 2–3. Eggs bluish white, finely spotted with brown or with a few brown spots. SIZE: c. 18  13. LAYING DATES: May, June, Sept, Oct. INCUBATION: period 14 days (in captivity). DEVELOPMENT AND CARE OF YOUNG: nestling period 17– 18 days (in captivity). BREEDING SUCCESS/SURVIVAL: no information.

Key References Brickell, N. and Konigkramer, T. (1997), Clement, P. et al. (1993).

Plate 31

Serinus pusillus (Pallas). Red-fronted Serin. Serin a ` front d’or.

(Opp. p. 474)

Passer pusillus Pallas, 1811. Zoogr. Rosso-Asiat., 2, p. 28; near Caucasus and Caspian Sea. S Turkey, Transcaucasia, Iran, mountains of central Asia east to Nepal, SW Tibet, Pamirs, Tien Shan, Tarbagatai and Sinkiang. Altitudinal and short-distance migrant; winter visitor to N Lebanon; vagrant Austria, Greece, Cyprus, Syria, Jordan and Egypt. A X at Bahig, Egypt, in Mar 1969 (several other records in Egyptian Sinai). A small, dark, stubby finch, sexes alike, ad. readily identified by black or blackish head and upper breast, scarlet forehead and warm ochre rump; rest of plumage strongly variegated with pale buff, rufescent mid-buffs and blackish browns; tertials, uppertail-coverts and tail feathers blackish with whitish buff edges; secondary coverts tipped buff, making a short, narrow but quite conspicuous wing-bar. Soft parts blackish. Imm. has dark brown forehead, plain brown or rufescent brown cheeks and ear-coverts, and slightly dark-streaked, buffy breast and flanks; base of lower mandible brownish. Length 12–13 cm; wing Y 72–77 (742), X 68–75 (722), tail Y 50–61 (557), X 49–57 (532); weight (Asia Minor) 8 YY 11–13 (119), 1 X 12. Keeps mainly above 2000 m but some descend to coasts in winter. Sometimes in leafy vegetation but perches mainly on dead branches, rocks and stones. Very agile in trees, feeding upside down at catkins. Forages, often with other finches, on open ground, eating mainly small seeds taken from surface, from low herbs and grasses by jumping up and fluttering; clings to stem to pick seeds from head of plant, bird’s weight sometimes pressing stem to the ground. Flies with rapid wingbeats and short, steep bounds, often trilling with a short, high-pitched, rippling twitter.

Genus Pseudochloroptila Wolters An endemic superspecies of 2 South African siskins or serin-like finches, YY with rufescent brown back, plain grey rump, unstreaked yellowish underparts, and bill finer (less stubby) than in Serinus. Y of one species quite large; Y of other species has black and white tips to remiges and rectrices. Courtship behaviour includes unique whirring flight by Y in front of X; nest placed in crevice or concealed on ground; nest constructed by both sexes; eggs pure white; incubation period (16 days) and fledgling period (16–18 days) each 2 days longer than in Serinus; nestling gapes uniformly red. Begging calls resemble those of Serinus scotops.

Pseudochloroptila totta

521

Pseudochloroptila totta superspecies

1 P. totta 2 P. symonsi

1

2

Pseudochloroptila totta (Sparrman). Cape Siskin. Serin totta.

Plate 31

Loxia totta Sparrman, 1786. Mus. Carlsonianum, fasc. 1, pl. 18; Hottentot country, i.e. Cape Province.

(Opp. p. 474)

Forms a superspecies with P. symonsi. Range and Status. Endemic resident, SW Cape Prov., South Africa; widespread and frequent, sea level up to c. 1000 m, north to Kobe Mts and Nieuwoudtville, east to Addo Nat. Park and Grahamstown. Considered to be Nearthreatened (Collar et al. 1994), but not under any immediate threat (Harrison et al. 1997). Description. ADULT Y: top of head dull olive, streaked blackish brown; mantle to back and scapulars dark brown; rump and short uppertail-coverts dull olive-green; long uppertail-coverts greyish brown. Tail feathers greyish brown, tipped white, broadly so on T5–T6 and with short white extension along shaft of inner web of T6. Lores dull olive; cheeks and ear-coverts dull olive streaked yellowish. Chin to breast and flanks olive-yellow, tinged brown on sides of breast and flanks, chin and throat sometimes with a few blackish brown streaks; belly brownish white; thighs brown; undertail-coverts brown, fringed greyish white. Primaries and secondaries blackish brown, tipped white; tertials dark brown with fine white fringes at tips; upperwing-coverts and alula dark brown, greater coverts with lighter greyish white tips, forming a fine wing-bar. Underwing-coverts and axillaries buffy yellow. Bill sepia-horn above, whitish horn below with sepia tip; eyes sepia; legs sepia. ADULT X: top of head buffy brown streaked dark brown; rest of upperparts dark brown, rump and short uppertail-coverts more olive, long uppertail-coverts fringed buff. Tail as ad. Y. Lores and sides of head olive-brown, cheeks and ear-coverts streaked buff. Chin and throat pale buff with short blackish brown streaks; breast and flanks olive-yellow, tinged browner on sides and flanks; belly brownish white; undertail-coverts brown with pale fringes. Wings as ad. Y, but white tips to flight feathers narrower. SIZE (5 YY, 4 XX): wing, Y 69–70 (696), X 67–69

(680); tail, Y 44–48 (466), X 43–48 (453); bill, Y 11–13 (119), X 105–12 (114); tarsus, Y 155–165 (160), X 15–155 (153); further data in Claassen (2002). WEIGHT: Y (n ¼ 9) 13–14 (135), X (n ¼ 9) 12–14 (130) (Claassen 2002). IMMATURE: similar to ad. X (Maclean 1993). NESTLING: hatchling skin dark pink, closed eyes blackish, swollen gape pale yellow, palate carmine-pink; quite long down on at least crown and nape (photo in Steyn 1996).

Field Characters. Length 12–13 cm. A small brown and yellow canary, largely unstreaked, with shortish tail. Readily separable by range alone from Drakensberg Siskin S. symonsi, also by yellower underparts and diagnostic white tips to wing and tail feathers, conspicuous in flight. Yellow Canary S. flaviventris in same habitat has brighter and more extensive yellow on underparts, yellow-green upperparts and wings and long yellow supercilium. X duller, less yellow below, throat and breast streaked, white on wing-tips often lost through wear; juv. like X but duller still. Voice. Tape-recorded (99, F, GIB). Contact call an upslurred, nasal ‘chewy’ or double ‘chewy-peesoo’, also written ‘pee-chee’, giving rise to common name Peachy Canary; this is incorporated into song, together with sweet thin notes and liquid twitters, ‘chewy-chewy-chee-cheetsew-tsew-titsew-lirrrrrr-chewy’, quality sweet but light and thin, rather casual. Other calls high, sibilant, ‘tsip’, ‘tsi-chit’, ‘tsi-peew’.

Pseudochloroptila totta

521

Pseudochloroptila totta superspecies

1 P. totta 2 P. symonsi

1

2

Pseudochloroptila totta (Sparrman). Cape Siskin. Serin totta.

Plate 31

Loxia totta Sparrman, 1786. Mus. Carlsonianum, fasc. 1, pl. 18; Hottentot country, i.e. Cape Province.

(Opp. p. 474)

Forms a superspecies with P. symonsi. Range and Status. Endemic resident, SW Cape Prov., South Africa; widespread and frequent, sea level up to c. 1000 m, north to Kobe Mts and Nieuwoudtville, east to Addo Nat. Park and Grahamstown. Considered to be Nearthreatened (Collar et al. 1994), but not under any immediate threat (Harrison et al. 1997). Description. ADULT Y: top of head dull olive, streaked blackish brown; mantle to back and scapulars dark brown; rump and short uppertail-coverts dull olive-green; long uppertail-coverts greyish brown. Tail feathers greyish brown, tipped white, broadly so on T5–T6 and with short white extension along shaft of inner web of T6. Lores dull olive; cheeks and ear-coverts dull olive streaked yellowish. Chin to breast and flanks olive-yellow, tinged brown on sides of breast and flanks, chin and throat sometimes with a few blackish brown streaks; belly brownish white; thighs brown; undertail-coverts brown, fringed greyish white. Primaries and secondaries blackish brown, tipped white; tertials dark brown with fine white fringes at tips; upperwing-coverts and alula dark brown, greater coverts with lighter greyish white tips, forming a fine wing-bar. Underwing-coverts and axillaries buffy yellow. Bill sepia-horn above, whitish horn below with sepia tip; eyes sepia; legs sepia. ADULT X: top of head buffy brown streaked dark brown; rest of upperparts dark brown, rump and short uppertail-coverts more olive, long uppertail-coverts fringed buff. Tail as ad. Y. Lores and sides of head olive-brown, cheeks and ear-coverts streaked buff. Chin and throat pale buff with short blackish brown streaks; breast and flanks olive-yellow, tinged browner on sides and flanks; belly brownish white; undertail-coverts brown with pale fringes. Wings as ad. Y, but white tips to flight feathers narrower. SIZE (5 YY, 4 XX): wing, Y 69–70 (696), X 67–69

(680); tail, Y 44–48 (466), X 43–48 (453); bill, Y 11–13 (119), X 105–12 (114); tarsus, Y 155–165 (160), X 15–155 (153); further data in Claassen (2002). WEIGHT: Y (n ¼ 9) 13–14 (135), X (n ¼ 9) 12–14 (130) (Claassen 2002). IMMATURE: similar to ad. X (Maclean 1993). NESTLING: hatchling skin dark pink, closed eyes blackish, swollen gape pale yellow, palate carmine-pink; quite long down on at least crown and nape (photo in Steyn 1996).

Field Characters. Length 12–13 cm. A small brown and yellow canary, largely unstreaked, with shortish tail. Readily separable by range alone from Drakensberg Siskin S. symonsi, also by yellower underparts and diagnostic white tips to wing and tail feathers, conspicuous in flight. Yellow Canary S. flaviventris in same habitat has brighter and more extensive yellow on underparts, yellow-green upperparts and wings and long yellow supercilium. X duller, less yellow below, throat and breast streaked, white on wing-tips often lost through wear; juv. like X but duller still. Voice. Tape-recorded (99, F, GIB). Contact call an upslurred, nasal ‘chewy’ or double ‘chewy-peesoo’, also written ‘pee-chee’, giving rise to common name Peachy Canary; this is incorporated into song, together with sweet thin notes and liquid twitters, ‘chewy-chewy-chee-cheetsew-tsew-titsew-lirrrrrr-chewy’, quality sweet but light and thin, rather casual. Other calls high, sibilant, ‘tsip’, ‘tsi-chit’, ‘tsi-peew’.

524

FRINGILLIDAE

General Habits. Inhabits scrubby fynbos veld on rocky mountain slopes and tops, sheltered valleys in foothills of larger mountains, tall vegetation at edges of relict forest tracts, stands of tall Protea spp. along rivers and streams, and coastal forelands; large clearings in forest country near Knysna; occurrences in Little Karoo are of birds likely also to be in fynbos pockets; occasional in pine plantations and stands of rooikrans Acacia cyclops. A familiar bird in some gardens and parks in towns close to mountains, e.g. Ladysmith, 500 km south of Cyrenaica coast. Egypt, small numbers of chlorotica have bred since mid 1980s in Nile Delta and Suez Canal areas (El Khatotba, Birket el Hagg, Wadi el Natrun, Abu Rauwash, El Abbassa, Bilbeis, Port Said); frequent winter visitor (mainly chlorotica, a few aurantiiventris), mid Oct to early Apr, mainly along Mediterranean coast between Matruh and Alexandria, also inland, south to Faiyum, Wadi el Natrun, Nile valley south once to Isna; rare on Red Sea coast (Hurghada, Safaga). Mauritania, rare: 1 Nouakchott May– June 1980, and at Nouadhibou single bird and a pair, Oct 1984, 4 in Jan 1986, 5 in Mar 1986 and 2 (1 a singing Y) in May 1988. Breeding density in Morocco from 5 pairs per km2 in maquis and broad-leaved woods to 15 pairs per km2 in cedar scrub.

Carduelis chloris

Description. C. c. voousi Roselaar: Moroccan Atlas Mts, N Algeria. ADULT Y: crown and nape bright olive-green, tinged grey, merging with yellower forehead; hindneck and sides of neck to short uppertail-coverts bright olive-green, tinged more yellow on rump; long uppertail-coverts grey. Central 2 pairs of tail feathers (T1–T2) and distal part of others black, fringed grey; base of outer web of T2 yellow; entire basal and middle part of T3–T6 yellow apart from black shafts. Lores dusky olive; cheeks and earcoverts olive-green, the latter tinged grey. Chin to breast and flanks bright olive-green, yellower on chin, grading to light greenish yellow on belly, thighs and undertail-coverts. Flight feathers black, P6–P9 with yellow outer webs proximal to emarginations, P1–P5 edged yellowish green, secondaries with grey outer edges grading to greenish distally; tertials greenish grey; primary coverts blackish, fringed greyish green; alula blackish, outer web dull olive-green; greater coverts grey; median and lesser coverts bright olive-green. Underwing-coverts and axillaries pale yellow. Bill fairly stout, pale pinkish or greyish

Carduelis chloris

537

black nor brightly coloured, but in some spp. dark horn or slaty; slender and quite long with straight culmen, or (a few spp.) robust and short with curved culmen. Mostly slim, long-winged and quite long-tailed, at least relative to Serinus. Could be polyphyletic. About 36 species, including the redpolls and linnets ‘Acanthis’ and 2 species regarded by some (e.g. Clement et al. 1993) as Serinus: Holarctic (2), Nearctic (4), Palearctic (9), Neotropical (14), Arabian (2), Oriental (3) and African (2). The 2 ‘serins’ that we transfer to Carduelis are ankoberensis of Ethiopia and the little-known Oriental estherae of Indonesia and Philippines. C. estherae lives at 1400–3400 m and 1 subspecies, mindanensis, combines goldfinch wings with the body plumage of Ethiopia’s enigmatic ankoberensis, which lives at 2600–4250 m. It would be far-fetched to claim that estherae and ankoberensis are closely allied and perhaps each deserves a genus to itself; but they may have adapted to tropical mountaintops in parallel and be evolving convergently within or at the limits of Carduelis. 7 spp. in Africa: 2 endemic ( johannis, ankoberensis), 4 Palearctic, breeding in N Africa (spinus, chloris, carduelis, cannabina) and 1 a very rare vagrant ( flammea).

Carduelis chloris (Linnaeus). European Greenfinch. Verdier d’Europe.

Plate 32

Loxia chloris Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 174; Europe (Sweden).

(Opp. p. 475)

Range and Status. Europe east to 75
E in Kirgiziya and Tadzhikistan, north to Baltic and 70
N in Norway, south to Mediterranean Basin, Canaries, NW Africa and N Egypt. Accidental Faeroes, Madeira. Introduced to Azores, Uruguay, NE Argentina, SE Australia, New Zealand and Chatham Island. Resident and partial migrant. Dispersive resident and winter visitor. Morocco, frequent to common in Atlas Mts, south on Atlantic coast to about Cap Draa (voousi), and throughout northern lowlands to Atlantic and Mediterranean coasts (vanmarli); up to 1700 m in Moyen Atlas and 1100–1200 m (once 2000 m) in Haut Atlas. Algeria, widespread in N from coast south to northernmost oases (Djelfa, Messaad, Laghouat, Monts des Ksour, Aı¨n Sefra, once Be´ni Abbe`s); up to greatest altitude of cedars in Djurdjura and 1400 m in Aure`s); also a winter visitor from Europe, but proportion of residents to immigrants unknown. Tunisia, uncommon to frequent, widespread in lowlands and mountains, breeding south to Kairouan and Mahdia, known in breeding season south to Gabe`s, Matmata, Douz and Tozeur; not certainly known as a winter visitor from Europe – coastal movements occur at Djerba, but no passage evident at Cap Bon, where all birds ringed seem to be residents. Libya, scarce but regular winter visitor to Tripoli coast in Nov–Mar in small flocks (once flock of 80), and to Cyrenaica in Oct–Mar (flocks of up to 20); rare, Serir, >500 km south of Cyrenaica coast. Egypt, small numbers of chlorotica have bred since mid 1980s in Nile Delta and Suez Canal areas (El Khatotba, Birket el Hagg, Wadi el Natrun, Abu Rauwash, El Abbassa, Bilbeis, Port Said); frequent winter visitor (mainly chlorotica, a few aurantiiventris), mid Oct to early Apr, mainly along Mediterranean coast between Matruh and Alexandria, also inland, south to Faiyum, Wadi el Natrun, Nile valley south once to Isna; rare on Red Sea coast (Hurghada, Safaga). Mauritania, rare: 1 Nouakchott May– June 1980, and at Nouadhibou single bird and a pair, Oct 1984, 4 in Jan 1986, 5 in Mar 1986 and 2 (1 a singing Y) in May 1988. Breeding density in Morocco from 5 pairs per km2 in maquis and broad-leaved woods to 15 pairs per km2 in cedar scrub.

Carduelis chloris

Description. C. c. voousi Roselaar: Moroccan Atlas Mts, N Algeria. ADULT Y: crown and nape bright olive-green, tinged grey, merging with yellower forehead; hindneck and sides of neck to short uppertail-coverts bright olive-green, tinged more yellow on rump; long uppertail-coverts grey. Central 2 pairs of tail feathers (T1–T2) and distal part of others black, fringed grey; base of outer web of T2 yellow; entire basal and middle part of T3–T6 yellow apart from black shafts. Lores dusky olive; cheeks and earcoverts olive-green, the latter tinged grey. Chin to breast and flanks bright olive-green, yellower on chin, grading to light greenish yellow on belly, thighs and undertail-coverts. Flight feathers black, P6–P9 with yellow outer webs proximal to emarginations, P1–P5 edged yellowish green, secondaries with grey outer edges grading to greenish distally; tertials greenish grey; primary coverts blackish, fringed greyish green; alula blackish, outer web dull olive-green; greater coverts grey; median and lesser coverts bright olive-green. Underwing-coverts and axillaries pale yellow. Bill fairly stout, pale pinkish or greyish

540

FRINGILLIDAE

flesh; eyes dark brown; legs pale pinkish or purplish flesh. ADULT X: top of head to back and scapulars olive-green with diffuse grey-brown streaks; rump and short uppertail-coverts plain bright olive-green; long uppertail-coverts olive-grey. Central tail feathers (T1–T2) dark olive-grey, less blackish than in Y; pattern on T3–T6 sometimes as in Y, but yellow typically absent or forming basal wedge only on inner webs, and confined to border of base and middle of outer webs, not reaching shafts. Sides of head and neck dusky olive-green. Chin yellowish olive-green; throat, breast and flanks dusky olive-green, merging with dull greenish yellow belly, becoming whiter on thighs, vent and undertail-coverts. Flight feathers greyish black, yellow edges of outer primaries often narrower than in Y; tertials and greater coverts brownish grey, edged brownish olive; primary coverts and alula greyish black; median and lesser coverts dull olive-grey. Bare parts as ad. Y. SIZE (10 YY, 1 X): wing, Y 84–89 (860), X 83; tail, Y 51–57 (540), X 54; bill, Y 17–185 (179), X 18; tarsus, Y 17–19 (179), X 17. Also, wing, Y (n ¼ 9) 85–92 (888), X (n ¼ 6) 82–88 (850); tail, Y (n ¼ 9) 53–59 (533), X (n ¼ 6) 51–56 (535); bill, Y (n ¼ 9) 165–195 (179), X (n ¼ 5) 175–19 (180); tarsus, Y (n ¼ 9) 17–19 (180), X (n ¼ 6) 17–185 (178). IMMATURE: juv. has upperparts buffish olive-brown, mantle and scapulars with short dark grey streaks, rump paler and buffier; sides of head and neck olive-grey, tinged yellow, underparts pale yellow to whitish, tinged olive-brown on upper breast, sides of breast and flanks; tail as ad. of same sex, but feather tips narrower, more pointed. NESTLING: down greyish white, long and plentiful on crown, back and upperwing, short and scanty on thighs and vent; mouth bright red, more pinkish at borders, gape flanges yellowish white. C. c. vanmarli (Voous): N Moroccan lowlands; also NW Spain and Portugal. Y slightly darker and browner above than voousi, rump greener; darker and greener below, yellow on belly tinged greener and more restricted. Smaller, with smaller bill: wing, Y (n ¼ 15) 80–85 (828); bill, Y (n ¼ 13) 155–17 (162). C. c. aurantiiventris (Cabanis): N Tunisia and NW Libya; also central and E Spain to Italy and W Greece; winters throughout NW Africa; a few winter in Egypt. Like voousi, but slightly darker green above, rather more golden yellow below; bill smaller. SIZE (5 YY, Tunisia and Libya, Mar): wing, 82–86 (840), bill 155– 165 (161). WEIGHT: (Balearic Islands, breeding) Y (n ¼ 20) 24–30 (258), X (n ¼ 26) 21–31 (261); (France, breeding) Y (n ¼ 20) 240–300 (258), X (n ¼ 26) 210–310 (261). C. c. chlorotica (Bonaparte): Levant and N Egypt. Upperparts and throat to breast more yellowish olive-green than in previous races; tertials and greater coverts paler grey; yellow on tail and flight feathers slightly paler.

Field Characters. Length 15 cm. Plump olive-green finch with large head, black lores, stout pale bill and shortish tail; in flight, yellow patches on primaries and base of tail conspicuous, and yellow-green rump contrasts with olive back. X duller, olive-brown; juv. dull brown, lightly streaked, especially below; rump brown, but yellow still showing in wings and tail. Voice. Tape-recorded (62–73, 76–89, 105, 110–113, B, C). Song a series of rattling trills differing in length, speed and quality, some liquid, others drier, interspersed with long wheeze, ‘zhweeeee’, nasal ‘zhu-wee’, ‘zhuyo’ or ‘juweeo’, and pure sweet ‘pew’, often in series, ‘pew-pew-pew-pewpew’. Common call a dry twitter, similar to song trills; for complete catalogue see Cramp and Perrins (1994). General Habits. Inhabits cedar Thuja scrub, broad-leaved woodland, farmland, gardens, parks, orchards, olive

groves, citrus plantations, hedgerows, stubble and weedy fields. In palm groves at Errachidia (Morocco). In pairs when nesting, family parties afterwards, and in small flocks in non-breeding season (winter), usually of 3–8, seldom of >12 in Morocco but sometimes 20 and occasionally up to 80 further east in N Africa; mixes with feeding flocks of finches, buntings and sparrows, often Common Linnets C. cannabina, European Serins Serinus serinus, Yellowhammers Emberiza citrinella, Reed Buntings E. schoeniclus and Eurasian Tree Sparrows Passer montanus. Can form flocks of thousands (in Europe). Forages in trees, shrubs and sometimes in herb, grass and weed layer, but mainly on ground. Locomotion on ground a short, creeping hop when picking up seeds or a short series of fast hops with upright stance when interacting with other granivores. Rises from ground in strong and rapid flight, looking thickset and showing yellow in wings and spread tail, often twittering; flight over trees and open spaces undulating. Roosts gregariously (in Europe). Resident, dispersing locally in winter; also winter visitor in Sept–May; 2 birds ringed Belgium recovered in Morocco, 1915 km away; 1 ringed Balearic Is recovered Lakhdaria, Algeria. Migrates by day and night, in NW Europe on rather narrow front. Ringing studies in Europe show that northern populations are quite strongly migratory, many birds wintering south to N, NE and extreme S Spain; common winter visitor and passage migrant in Malta and Cyprus; aurantiiventris in Spain are partial migrants and probably move also into Morocco. About 20,000 birds cross Strait of Gibraltar in Oct–Nov (mainly late Oct and early Nov); spring passage there is less strong and more protracted, from early Feb to early May, mainly in 1st half Mar. Occasionally marked passage visible also on Moroccan coast, e.g. 200 moving south at Moulay-Bousselham (early Nov). Food. Not studied in Africa. In Europe seeds, often quite large and hard, mainly of cereals, composites, crucifers, Rosaceae and Polygonaceae; eats a few invertebrates in breeding season. Breeding Habits. Solitary nester, sometimes in loose colonies of 4–6 pairs (N Europe) or dozens (Israel), when nests at least 10 m apart; monogamous and polygynous (24% of nests in England, Y with 2 and sometimes 3–5 XX); not territorial, but pair defends immediate surrounds of nest. Y sings from tree-top or telephone post, in upright posture with wings and tail vibrating in time with ‘chi’ notes in song; he then often launches himself into characteristic song-flight up to c. 50 m long, weaving somewhat erratically and circling widely amongst or over tree-tops, with slow, deep wing-beats, body rolling from side to side. X sometimes sings, less vehemently than Y. In Head-forward aggressive display, bird crouches with head thrust forward at rival, bill often wide open, plumage sleeked. Courting Y hops and flies continually after X, and uses Head-forward display like aggressive one but with wrists and wing-tips raised to level of back and tail lowered and spread (A); Y also postures with head high, closed bill tilted up, wing-tips lowered, and tail slightly raised (B).

Carduelis carduelis

541

with fine grass, plant down, rootlets, feathers and hair. Ext. diam. 120, int. diam. 50, ext. height 75, int. depth 40. Sited in strong fork or against trunk of small tree (often a conifer), dense bush or in hedge or among creepers. Built by X, taking 8–12 days; Y often accompanies her on material-collecting forays. EGGS: 3–5 (av. of 50 clutches, 426). Sub-elliptical, smooth, slightly glossy. Greyish or bluish white, or sandy brown, sparsely spotted and blotched with reddish, purplish or blackish, mainly at broad end, sometimes forming a ring. SIZE: (aurantiiventris, n ¼ 113) 185–235  125–160 (205  148); WEIGHT: av. 2.1 (Cramp and Perrins 1994). LAYING DATES: NW Africa, Mar–May, mainly Apr–May. INCUBATION: by X, sitting often very tightly. Period: av. 129 days (Europe). DEVELOPMENT AND CARE OF YOUNG: young brooded for first 5–6 days by X; fed mainly by Y at first, then by Y and X more or less equally, by regurgitation. Nestling faeces removed by both parents, up to days 8 or 9. Nestling period: (n ¼ 160) 141–147 (144) days (Europe). BREEDING SUCCESS/SURVIVAL: in E Spain, 51% of 223 eggs in Mar–Apr resulted in flying young, 49% of 171 eggs in May, 26% of 79 eggs in June and 3% of 58 eggs in July; 10% of young died of starvation in Mar–Apr, rising to 47% in July; seed supply spoiled by the cutting of herb layer in orange grove where population nested (Cramp and Perrins 1994).

NEST:

strongly made open cup, thick-walled, made of moss, lichen and grass, on foundation of dry twigs; lined

Key References Clement, P. et al. (1993), Cramp, S. and Perrins, C.M. (1994), Isenmann, P. and Moali, A. (2000), The´venot, M. (1982).

Carduelis carduelis (Linnaeus). European Goldfinch. Chardonneret ´ele ´gant.

Plate 32

Fringilla Carduelis Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 180; Europe (Sweden).

(Opp. p. 475)

Range and Status. Europe, north to S Norway, S Sweden and S Finland, and W Asia east to Yenesei Valley, W Mongolia and Zaysan Basin; south to Canaries, NW Africa, NE Africa (Nile Valley and Delta), Levant, SW Iran and W Himalayan foothills; largely absent between Caspian Sea and L. Baikal. Resident and partial migrant; summer visitor in northernmost parts of range and in E-central Turkey; winter visitor to Sinai and Iraq. A popular cagebird since Roman times; prominent (illegally) in bird markets in Morocco from El Hedim to Mekne`s; introduced to Azores, Bermuda, locally in S America, S Australia and New Zealand; also to Santiago, Cape Verde Is, but extinct since 1960s. Resident and winter visitor. Morocco, abundant north of Haut Atlas Mts; common in Atlas Mts, up to 2600 m in spring and 3000 m in autumn; large flocks winter south to Marrakech and Errachidia (where also common in summer); south to Taliouine, Skoura, El Kelaa des Mgouna, Todra

Gorges, Tinerhir, Rgabi (Ouadi Draa) and Merzouga. Algeria, common and widespread from coast to northernmost oases (Biskra, Laghouat, Aı¨n Sefra, Be´char); up to 2100 m in Aure`s; disperses in winter, some birds to south of breeding range, in Ksour Mts and south to Be´ni Abbe`s, Hassi Touiel and El Gole`a. Tunisia, common and widespread in N, locally abundant, south to central plateau; absent from Kroumirie; scarce along Algerian border, south to Gafsa and on coast to Gabe`s; Zembra I; also an uncommon migrant at Cap Bon from Europe, late Sept to late Apr, mainly Oct–Mar. Libya, breeds in Tripoli area and south to Jebel Nafusa, where uncommon; in Cyrenaica breeds mainly in Benghazi area and Jebel Akhdar, north and east to Derna; records, mainly of small flocks, from Sirte coast, near Tobruk and in desert at Serir. Egypt, common almost throughout Nile Delta, on coast west to Matruh and in most parts of Nile Valley south to Aswan, also in Faiyum and Suez Canal area; breeds in Wadi el

Carduelis carduelis

541

with fine grass, plant down, rootlets, feathers and hair. Ext. diam. 120, int. diam. 50, ext. height 75, int. depth 40. Sited in strong fork or against trunk of small tree (often a conifer), dense bush or in hedge or among creepers. Built by X, taking 8–12 days; Y often accompanies her on material-collecting forays. EGGS: 3–5 (av. of 50 clutches, 426). Sub-elliptical, smooth, slightly glossy. Greyish or bluish white, or sandy brown, sparsely spotted and blotched with reddish, purplish or blackish, mainly at broad end, sometimes forming a ring. SIZE: (aurantiiventris, n ¼ 113) 185–235  125–160 (205  148); WEIGHT: av. 2.1 (Cramp and Perrins 1994). LAYING DATES: NW Africa, Mar–May, mainly Apr–May. INCUBATION: by X, sitting often very tightly. Period: av. 129 days (Europe). DEVELOPMENT AND CARE OF YOUNG: young brooded for first 5–6 days by X; fed mainly by Y at first, then by Y and X more or less equally, by regurgitation. Nestling faeces removed by both parents, up to days 8 or 9. Nestling period: (n ¼ 160) 141–147 (144) days (Europe). BREEDING SUCCESS/SURVIVAL: in E Spain, 51% of 223 eggs in Mar–Apr resulted in flying young, 49% of 171 eggs in May, 26% of 79 eggs in June and 3% of 58 eggs in July; 10% of young died of starvation in Mar–Apr, rising to 47% in July; seed supply spoiled by the cutting of herb layer in orange grove where population nested (Cramp and Perrins 1994).

NEST:

strongly made open cup, thick-walled, made of moss, lichen and grass, on foundation of dry twigs; lined

Key References Clement, P. et al. (1993), Cramp, S. and Perrins, C.M. (1994), Isenmann, P. and Moali, A. (2000), The´venot, M. (1982).

Carduelis carduelis (Linnaeus). European Goldfinch. Chardonneret ´ele ´gant.

Plate 32

Fringilla Carduelis Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 180; Europe (Sweden).

(Opp. p. 475)

Range and Status. Europe, north to S Norway, S Sweden and S Finland, and W Asia east to Yenesei Valley, W Mongolia and Zaysan Basin; south to Canaries, NW Africa, NE Africa (Nile Valley and Delta), Levant, SW Iran and W Himalayan foothills; largely absent between Caspian Sea and L. Baikal. Resident and partial migrant; summer visitor in northernmost parts of range and in E-central Turkey; winter visitor to Sinai and Iraq. A popular cagebird since Roman times; prominent (illegally) in bird markets in Morocco from El Hedim to Mekne`s; introduced to Azores, Bermuda, locally in S America, S Australia and New Zealand; also to Santiago, Cape Verde Is, but extinct since 1960s. Resident and winter visitor. Morocco, abundant north of Haut Atlas Mts; common in Atlas Mts, up to 2600 m in spring and 3000 m in autumn; large flocks winter south to Marrakech and Errachidia (where also common in summer); south to Taliouine, Skoura, El Kelaa des Mgouna, Todra

Gorges, Tinerhir, Rgabi (Ouadi Draa) and Merzouga. Algeria, common and widespread from coast to northernmost oases (Biskra, Laghouat, Aı¨n Sefra, Be´char); up to 2100 m in Aure`s; disperses in winter, some birds to south of breeding range, in Ksour Mts and south to Be´ni Abbe`s, Hassi Touiel and El Gole`a. Tunisia, common and widespread in N, locally abundant, south to central plateau; absent from Kroumirie; scarce along Algerian border, south to Gafsa and on coast to Gabe`s; Zembra I; also an uncommon migrant at Cap Bon from Europe, late Sept to late Apr, mainly Oct–Mar. Libya, breeds in Tripoli area and south to Jebel Nafusa, where uncommon; in Cyrenaica breeds mainly in Benghazi area and Jebel Akhdar, north and east to Derna; records, mainly of small flocks, from Sirte coast, near Tobruk and in desert at Serir. Egypt, common almost throughout Nile Delta, on coast west to Matruh and in most parts of Nile Valley south to Aswan, also in Faiyum and Suez Canal area; breeds in Wadi el

542

FRINGILLIDAE

Carduelis carduelis

Natrun, Bahariya, Farafra, Dakhla and Kharga in Western Desert; irregular visitor in Eastern Desert (Ain Sukhna, Hurghada, Safaga). Breeding density in Moroccan maquis scrub of 308 pairs per km2 (The´venot 1982). Very high densities of wintering birds in Gibraltar (650 birds per km2), where European Goldfinch is commonest bird in winter, and in N Morocco, where also one of the commonest. Description. C. c. parva Tschusi: Morocco to Libya; also S France, Iberia, Balearic Islands, Madeira, Azores and Canary Is. ADULT Y: forehead and forecrown to rear corner of eye, forecheeks, chin and upper throat dark red, forming bright face patch, crossed by black band through lores to below and immediately behind eye; mid-crown to nape black; hindneck, mantle, scapulars and back cinnamon-brown; rump and uppertailcoverts white, partly concealed by cinnamon-brown feather tips. Tail feathers black, T1–T2 (and sometimes T3) with white tips, inner web of (T3) T4 with small white patch, inner webs of T5–T6 with large white patch. Ear-coverts and hindcheeks buffy white, forming large vertical patch continuous with buffy white lower throat, and bordered behind by black band connecting with black nape. Sides of breast suffused rich cinnamon-brown, often connected by cinnamon-brown band across upper breast; flanks, patches at sides of belly and undertail-coverts tinged warm brown; rest of underparts white. Remiges black, inner primaries, secondaries and tertials broadly tipped white; middle and basal part of outer webs of P1–P8 and bases of secondaries golden yellow. Primary coverts and alula black; greater coverts golden yellow; median coverts black; lesser coverts black, fringed cinnamon-brown. Golden yellow on flight feathers and greater coverts forms broad band across closed wing. Underwing-coverts and axillaries greyish white, mottled blackish near carpal edge. Bill sharply pointed, pale horn or whitish, tipped dusky in nonbreeding season; eyes dark brown; legs pale flesh or light brown. ADULT X: similar to ad. Y, but red hardly extends to forecrown, rear corner of eye or upper throat; black on crown and nape sootier, whitish patch on ear-coverts and hindcheeks tinged browner; usually no mirror on inner web of T4; inner median

coverts typically greyer. SIZE (10 YY, 10 XX): wing, Y 73–78 (752), X 70–75 (726); tail, Y 43–48 (453), X 41–47 (447); bill, Y 15–165 (155), X 145–16 (147); tarsus, Y 14–155 (146), X 135–15 (143). WEIGHT: (Balearic Is) Y (n ¼ 25) 117–157 (137), X (n ¼ 21) 108–150 (127). IMMATURE: juv. has head and upperparts light greyish brown, head and neck speckled, and mantle, back and scapulars spotted or streaked dark grey-brown; nape, supercilium and below eye paler, more buffish; rump and uppertail-coverts buffy white, mottled grey; underparts pale grey-buff, whiter on belly; chin and throat speckled and breast and flanks more coarsely marked or streaked dark grey; tail feathers as ad. but more pointed, T4 without white patch on inner web; wings as ad. but median and lesser coverts tipped buff, and pale tips to remiges larger and buffier. NESTLING: down fairly long, dark grey, mainly on crown, upperwing, back and thighs; tongue and floor of mouth crimson, roof of mouth dark lilac, gape-flanges creamy white. C. c. niedecki Reichenow: Egypt, Levant, Cyprus, S Greece and Asia Minor to SW Iran. Less cinnamon than parva, paler and greyer. Larger: wing, Y (n ¼ 21) 77–84 (796). WEIGHT: (Cyprus) Y (n ¼ 59) 13–215 (151), X (n ¼ 35) 13–195 (145). C. c. carduelis (Linnaeus): Europe from central France and Denmark east to Urals; migrant to NW Africa. Like parva but larger: wing, Y (n ¼ 21) 78–84 (810). WEIGHT: (W Europe) Y (n ¼ 131) 13–20 (165), X (n ¼ 80) 131–19 (153). C. c. britannica (Hartert): Britain, Ireland and adjacent NW Europe; migrant to Morocco. Upperparts and belly patches slightly duller than nominate race, less cinnamon; slightly smaller, wing, Y (n ¼ 24) 75–80 (778).

Field Characters. Length 12–13 cm. At all ages shows unique flight pattern, black wings crossed by broad yellow band, white rump, and white-tipped black tail showing white side-panels when spread. Ad. unmistakable, with red face and black and white head; head of juv. plain brown. Tinkling call often heard before bird seen; flight bounding and erratic. Voice. Tape-recorded (62–73, 76–89, 105, 110–113, B, C). Common contact call a liquid, tinkling ‘pintsi’ or ‘pintsitit’, often given in flight, and forming part of song when woven into series of trills and twitters; song bustling and continuous, with barely perceptible breaks between sections; common element is liquid ‘pit-pit-pit-pit’, and a few dry, swallow-like burrs may be added. Other calls include nasal ‘djewy’ or ‘zwee’ when feeding in flock, hard ‘jijijijijit’ and softer ‘tikluyu’. For comprehensive repertoire see Cramp and Perrins (1994). General Habits. Well known in Europe, less so in NW Africa. Inhabits cultivation (farms, fields of cereals and sunflowers, hedges, avenues, gardens, orchards, parks, leafy suburbs, plantations, tree nurseries), broad-leaved and conifer copses, scrub, open places with tall weeds; heath and steppe, oases, arid hillsides, and river valleys; in mountains, but commonest in lowlands. Occurs in pairs and small parties, and aggregates into flocks of tens and often hundreds in winter. Feeds mainly in seeding heads of 05–2 m tall annuals, perching acrobatically, sometimes almost upside down. Regularly wipes bill on perch after feeding. Occasionally forages on ground, for instance with European Greenfinches Carduelis chloris around bird tables, but seldom mixes with different finches in other situations,

Carduelis spinus although largest flocks in Morocco often accompanied by Greenfinches, Common Linnets C. cannabina, European Serins Serinus serinus and House Sparrows Passer domesticus. Birds in flock forage and fly together, often several m apart in bushes or trees and in flight, but behaviour is synchronized and structured. Mobile and flighty, flock traveling several km every day to forage. Flight rapid, undulating or bounding. Flock leaves roost in early morning, forages until mid-morning, then rests in tall trees. Sometimes bathes, mainly about mid-day and in late afternoon. Breeding pairs visit neighbours early in morning and sing. Resident, dispersing and wandering within N African breeding range. Partial migrant in Europe and Asia, large proportion of population moving 2000 km or more to south and southwest in late Sept to early Nov, with some onward passage through winter, and return migration in Mar–Apr; NW European birds regularly winter in NW and N Africa. Abundant autumn migrant across Strait of Gibraltar, of NW European as well as Spanish birds (Asensio 1984, 1986), e.g. 58,000 in 1977; main passage across Strait from mid-Oct to mid-Nov and autumn daytime movements sometimes apparent in N Morocco; hundreds move south in Michliffen-Ifrane and Merja Zerga areas in late Oct to mid Nov; influxes into Morocco from Spain can occur in middle of severe winters. Markedly commoner on coast at Settat (7
370 W) on autumn passage, mid-Sept to Nov, than in breeding months. Winter visitors leave W Morocco from mid Feb; main spring passage at Strait of Gibraltar in Mar and 1st week Apr. Commoner in Tafilalet, SE Morocco, in winter than summer. 5 recoveries from Balearic Islands to between Algiers and Tizi-Ouzou, Algeria (1 next day); birds ringed in Tunisia recovered Malta, Italy and former Yugoslavia. Britain to Morocco (3), south to Casablanca; Cyprus to Alexandria, Egypt (1). Food. Small, unripe seeds; in Europe of Compositae, especially thistles Carduus, dandelions Taraxacum, ragworts Senecio and burdocks Arctium and sunflowers Helianthus; also hemp Cannabis sativa (Cannabidaceae); in winter, seeds of pines Pinus and alders Alnus. YY (more than shorter-billed XX) particularly fond of teasel Dipsacus. Occasionally eats small leaves (of e.g. Stellaria), catkins of birch Betula, and pieces of flowers; and caterpillars and other small invertebrates, mainly in breeding season. Breeding Habits. Solitary nester, often in loose groups of 3 or more pairs. Monogamous. Defends small territory

543

around nest; aggressive when nesting, especially X when nest-building and Y thereafter; otherwise not an aggressive bird. Y perches conspicuously on top of bush and occasionally sings; he threatens by crouching at perch and rotating body through c. 45
one way and the other. Y in Song-flight flies above trees, hesitantly, with slow, deep wing-beats; at other times he pursues X vigorously. Y commonly courtship-feeds X. NEST: compact, very neat cup, made of moss, grass, rootlets and spider web; silk is often used profusely, making walls and foundation felt-like; lined with plant down, animal hair and sometimes feathers. Nest occasionally decorated outside with aromatic flowers. SIZE: (n ¼ 3, Azores) ext. diam. 76, int. diam. 43, ext. height 48, int. depth 35. Sited amongst outermost twigs of tree; well concealed; often in fruit trees, also limes Tilia and conifers. Built by X, in 3 bouts of activity (early morning, mid-day, late afternoon), taking 4–14 (av. 7) days (Europe). Y escorts X on collecting trips and may bring small amounts of material. EGGS: 3–5 (av. of 17 clutches 42). Sub-elliptical, smooth, slightly glossy. Very pale bluish white, sparsely scrawled and spotted with purple-brown or reddish, mainly at broad end. SIZE: (parva, n ¼ 18) 140–172  120–130 (166  124), (niediecki, n ¼ 68) 157–190  123–137 (171  130). WEIGHT: parva, av. 134, niediecki, av. 150. LAYING DATES: NW Africa, Mar–July, mainly Apr–May; Egypt (fledglings Apr, May). INCUBATION: by X only. Period (Europe): (n ¼ 41) 9–14 (121) days. DEVELOPMENT AND CARE OF YOUNG: young brooded by X for 7–9 days (Europe); at that stage Y brings food up to 38 times per day, giving it to X by regurgitation; brooding X eats it herself or passes it to nestlings. Later, young fed and cared for by both parents. Eyes begin to open at 5 days and are fully open 2 days later. All faecal sacs swallowed by X for 1st week and some for a few more days; thereafter, young defaecate over and onto rim of nest. Nestling period: (n ¼ 40) 13–18 (147) days. Fledglings out of nest fed by parents, become independent after 7–10 days (Europe). BREEDING SUCCESS/SURVIVAL: trapped for the cage-bird trade in several N African and Mediterranean lands. Birds ringed in NW Europe and recovered in Spain have annual ad. mortality of 65% (Cramp and Perrins 1994). Oldest ringed bird 85 years. Key References Asensio, B. (1986), Cramp, S. and Perrins, C.M. (1994), Finlayson, C. (1992), Isenmann, P and Moali, A. (2000).

Carduelis spinus (Linnaeus). Eurasian Siskin. Tarin des aulnes.

Plate 32

Fringilla Spinus Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 181; Europe.

(Opp. p. 475)

Range and Status. Palearctic; breeds from Scotland to Japan, south to Pyrenees, Alps, Balkans, N Turkey and Caucasus; migrates southwestward to winter throughout Europe, Turkey and Israel. An ‘invasion’ species, wintering sparsely in NW Africa, N Egypt and Sinai, but in some years quite common there.

Morocco, not recorded in every winter but in most years 5–10 records of 1–4 birds and 2 or 3 records of flocks of 5–10, all in N, south to Oulme`s, Aoufouss, Boumalnedu-Dade`s, Errachidia, Gorges du Dade`s and Gorges du Todra; common on Jebel Zerhoun, Mekne`s, and elsewhere in 1991, in flocks of up to 25 (Mar, Sidi-Bouzid) and 55

Carduelis spinus although largest flocks in Morocco often accompanied by Greenfinches, Common Linnets C. cannabina, European Serins Serinus serinus and House Sparrows Passer domesticus. Birds in flock forage and fly together, often several m apart in bushes or trees and in flight, but behaviour is synchronized and structured. Mobile and flighty, flock traveling several km every day to forage. Flight rapid, undulating or bounding. Flock leaves roost in early morning, forages until mid-morning, then rests in tall trees. Sometimes bathes, mainly about mid-day and in late afternoon. Breeding pairs visit neighbours early in morning and sing. Resident, dispersing and wandering within N African breeding range. Partial migrant in Europe and Asia, large proportion of population moving 2000 km or more to south and southwest in late Sept to early Nov, with some onward passage through winter, and return migration in Mar–Apr; NW European birds regularly winter in NW and N Africa. Abundant autumn migrant across Strait of Gibraltar, of NW European as well as Spanish birds (Asensio 1984, 1986), e.g. 58,000 in 1977; main passage across Strait from mid-Oct to mid-Nov and autumn daytime movements sometimes apparent in N Morocco; hundreds move south in Michliffen-Ifrane and Merja Zerga areas in late Oct to mid Nov; influxes into Morocco from Spain can occur in middle of severe winters. Markedly commoner on coast at Settat (7
370 W) on autumn passage, mid-Sept to Nov, than in breeding months. Winter visitors leave W Morocco from mid Feb; main spring passage at Strait of Gibraltar in Mar and 1st week Apr. Commoner in Tafilalet, SE Morocco, in winter than summer. 5 recoveries from Balearic Islands to between Algiers and Tizi-Ouzou, Algeria (1 next day); birds ringed in Tunisia recovered Malta, Italy and former Yugoslavia. Britain to Morocco (3), south to Casablanca; Cyprus to Alexandria, Egypt (1). Food. Small, unripe seeds; in Europe of Compositae, especially thistles Carduus, dandelions Taraxacum, ragworts Senecio and burdocks Arctium and sunflowers Helianthus; also hemp Cannabis sativa (Cannabidaceae); in winter, seeds of pines Pinus and alders Alnus. YY (more than shorter-billed XX) particularly fond of teasel Dipsacus. Occasionally eats small leaves (of e.g. Stellaria), catkins of birch Betula, and pieces of flowers; and caterpillars and other small invertebrates, mainly in breeding season. Breeding Habits. Solitary nester, often in loose groups of 3 or more pairs. Monogamous. Defends small territory

543

around nest; aggressive when nesting, especially X when nest-building and Y thereafter; otherwise not an aggressive bird. Y perches conspicuously on top of bush and occasionally sings; he threatens by crouching at perch and rotating body through c. 45
one way and the other. Y in Song-flight flies above trees, hesitantly, with slow, deep wing-beats; at other times he pursues X vigorously. Y commonly courtship-feeds X. NEST: compact, very neat cup, made of moss, grass, rootlets and spider web; silk is often used profusely, making walls and foundation felt-like; lined with plant down, animal hair and sometimes feathers. Nest occasionally decorated outside with aromatic flowers. SIZE: (n ¼ 3, Azores) ext. diam. 76, int. diam. 43, ext. height 48, int. depth 35. Sited amongst outermost twigs of tree; well concealed; often in fruit trees, also limes Tilia and conifers. Built by X, in 3 bouts of activity (early morning, mid-day, late afternoon), taking 4–14 (av. 7) days (Europe). Y escorts X on collecting trips and may bring small amounts of material. EGGS: 3–5 (av. of 17 clutches 42). Sub-elliptical, smooth, slightly glossy. Very pale bluish white, sparsely scrawled and spotted with purple-brown or reddish, mainly at broad end. SIZE: (parva, n ¼ 18) 140–172  120–130 (166  124), (niediecki, n ¼ 68) 157–190  123–137 (171  130). WEIGHT: parva, av. 134, niediecki, av. 150. LAYING DATES: NW Africa, Mar–July, mainly Apr–May; Egypt (fledglings Apr, May). INCUBATION: by X only. Period (Europe): (n ¼ 41) 9–14 (121) days. DEVELOPMENT AND CARE OF YOUNG: young brooded by X for 7–9 days (Europe); at that stage Y brings food up to 38 times per day, giving it to X by regurgitation; brooding X eats it herself or passes it to nestlings. Later, young fed and cared for by both parents. Eyes begin to open at 5 days and are fully open 2 days later. All faecal sacs swallowed by X for 1st week and some for a few more days; thereafter, young defaecate over and onto rim of nest. Nestling period: (n ¼ 40) 13–18 (147) days. Fledglings out of nest fed by parents, become independent after 7–10 days (Europe). BREEDING SUCCESS/SURVIVAL: trapped for the cage-bird trade in several N African and Mediterranean lands. Birds ringed in NW Europe and recovered in Spain have annual ad. mortality of 65% (Cramp and Perrins 1994). Oldest ringed bird 85 years. Key References Asensio, B. (1986), Cramp, S. and Perrins, C.M. (1994), Finlayson, C. (1992), Isenmann, P and Moali, A. (2000).

Carduelis spinus (Linnaeus). Eurasian Siskin. Tarin des aulnes.

Plate 32

Fringilla Spinus Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 181; Europe.

(Opp. p. 475)

Range and Status. Palearctic; breeds from Scotland to Japan, south to Pyrenees, Alps, Balkans, N Turkey and Caucasus; migrates southwestward to winter throughout Europe, Turkey and Israel. An ‘invasion’ species, wintering sparsely in NW Africa, N Egypt and Sinai, but in some years quite common there.

Morocco, not recorded in every winter but in most years 5–10 records of 1–4 birds and 2 or 3 records of flocks of 5–10, all in N, south to Oulme`s, Aoufouss, Boumalnedu-Dade`s, Errachidia, Gorges du Dade`s and Gorges du Todra; common on Jebel Zerhoun, Mekne`s, and elsewhere in 1991, in flocks of up to 25 (Mar, Sidi-Bouzid) and 55

544

FRINGILLIDAE

Carduelis spinus

(Jan, Rabat). Many in 1993–1994, particularly in mouths of Oued Souss and Oued Massa. Algeria, uncommon, Oct– Apr (rare, May), numbers varying each winter, in coastal plain and in Lesser Kabylie and E Tell (the only area with favourite food tree, alders Alnus glutinosa); near Sahara, singles at Ghardaı¨a and El Gole´a, Jan 1976. Tunisia, rare, Nov–Feb (very rare Mar), mainly in Kroumirie and Cape Bon areas, exceptionally south to Gafsa and Gabe`s. Libya, small flock at Taguira, Tripoli, Oct 1951; 5–6 in Tripoli city, Jan–Mar 1961. Egypt, in most winters an uncommon visitor to Nile Delta and N Nile valley, south to Faiyum, but abundant in Feb–Mar 1907 and abundant near Port Said in winter 1966–1967; very rare elsewhere (1, Wadi el Natrun, Oct 1914; 2, Bahariya, Oct 1983; 3, 90 km west of Safaga, Nov 1993). In Algeria, recoveries from Britain (3), Sweden (2), Germany and Baltic States (6) and Italy (2). Description. ADULT Y: forehead to nape black; hindneck, mantle, back and scapulars olive-green, narrowly streaked black; rump lemon-yellow with narrow dusky streaks; uppertail-coverts dull olive-green. Tail feathers blackish, T2–T6 with base and middle part of outer webs bright yellow and of inner webs paler yellow, T1 and black distal part of T2–T6 with narrow yellowish green fringe to outer web when fresh, and yellowish grey fringe to inner web and around tip. Greenish yellow superciliary stripe prominent behind eye; lores and band through upper ear-coverts olive-green; lower ear-coverts and cheeks yellow, streaked dark olive-green; sides of neck olive-yellow, mottled olive-green. Patch on chin and centre of upper throat black with some yellow mottling; sides of throat, lower throat and breast olive-yellow, merging into white belly, sides of breast mottled dark olivegreen; flanks whitish, tinged olive-yellow, with long blackish streaks; undertail-coverts pale olive-yellow with fine blackish central streaks. Remiges blackish; primaries edged olive-yellow, P1–P6 with pale yellow bases forming small patch on closed wing; secondaries narrowly edged olive-yellow, with pale yellow bases hidden below coverts; tertials edged olive-yellow, becoming

whitish toward tips. Primary coverts and alula blackish; greater and median coverts blackish, broadly tipped bright olive-green, forming 2 prominent wing-bars; lesser coverts olive-green. Underwing-coverts and axillaries and undersides of inner borders of flight feathers pale olive-yellow. Bill thin and sharply pointed, greyish or brownish horn, paler on lower mandible, with blackish culmen; eyes dark brown; legs dark brown to flesh brown. ADULT X: forehead to nape greenish grey, streaked dark olive-grey; bar across hindneck streaked olive-yellow and dark olive-grey; mantle, scapulars and back light olive-grey, merging with pale olive-yellow rump, all boldly streaked blackish; uppertail-coverts olive-grey. Tail feathers blackish, yellow confined to borders of base and middle part of both webs of T2–T5 and inner web of T6. Superciliary stripe poorly marked, yellowish white; lores and upper ear-coverts dusky olive-grey; cheeks and lower ear-coverts olive-grey, streaked pale yellowish; sides of neck olive-yellow streaked olive-grey. Underparts greyish white, tinged oliveyellow, chin sparsely mottled with blackish, breast and flanks broadly streaked blackish and undertail-coverts more narrowly so. Wing as ad. Y, but yellow at bases of secondaries and inner primaries less extensive, more hidden by coverts; greater and median coverts broadly tipped pale olive-yellow to form 2 prominent wingbars; lesser coverts olive-grey. Underwing coverts greyish white, tinged yellow. Bare parts as ad. Y. SIZE (10 YY, 10 XX): wing, Y 70–73 (711), X 68–73 (703); tail, Y 39–47 (435), X 42–46 (435); bill, Y 12–13 (124), X 12–13 (124); tarsus, Y 135– 145 (139), X 135–145 (139). WEIGHT: W Europe, Y (n ¼ 385) 101–185 (135), X (n ¼ 379) 103–18 (130). IMMATURE: in 1st winter like ad., but retained juv. tertials and outer greater coverts have narrower and whiter tips; retained juv. tail feathers more pointed, Y with yellow more confined to basal edges of feathers than in ad.

Field Characters. Length 12 cm. A tiny green and yellow finch, Y with black cap and chin, X greener, less yellow and streakier; juv. brown and more heavily streaked; all ages have black wings with yellow bars forming bands on inner half of wing in flight, and yellow patches at base of tail. Rump greenish yellow, less bright and contrasting than European Serin Serinus serinus, but latter lacks yellow in wings and tail. Further distinguished from serins Serinus spp. by fine pointed bill and tit-like manner of feeding. Voice. Tape-recorded (62–73, 76–89, 105, 110–113, B, C). Calls include nasal ‘zerwy’, shrill liquid ‘pee-yee’, ‘pleep’ or ‘peeo’ and dry ‘reedada’; these and other calls incorporated into song, a long, rapid medley of liquid trills and dry chatters at various speeds and pitches, often ending with long buzzy ‘zheeeee’. General Habits. In NW Africa strongly associated with alders and cedars; also inhabits woods of, and open country with, spruce, fir, pine and larch, but prone to occur on edges of all lowland broadleaved woods, in shrubs along streams and in parks and gardens. Arboreal. Feeds in trees, eating seeds mainly of spruce, alder and birch. Readily comes to bird feeders; does not usually come to ground nor into cereal crops, but in Egypt ‘the 1907 invasion apparently resulted in serious agricultural damage, as a substantial proportion of the barley crop near Maryut was said to have been destroyed’ by siskins (Goodman and Meininger 1989). Gregarious, forming small flocks on its own and consorting loosely with other finches. Flight fast and undulating; often calls in flight.

Carduelis cannabina Occurs irregularly in NW and N Africa in Oct–Apr (–May); however, a regular migrant at Gibraltar in Oct– Nov and Feb to early Apr, scarce in some years and numerous in others; conspicuous invasions in 1949–1950, 1959–1960, 1961–1962, 1965–1966, 1966–1967, 1972–1973, 1988–1989 (Finlayson 1992) and 1993–1994. Bird ringed in Germany recovered Morocco.

545

Food. Seeds, mainly of conifers. Not studied in Africa, but see above.

Key Reference Cramp, S. and Perrins, C.M. (1994).

Carduelis flammea (Linnaeus). Common Redpoll. Sizeron flamme´.

Plate 32

Fringilla flammea Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 182; Europe.

(Opp. p. 475)

Holarctic, breeding between 55
and 70
N; partial migrant, wintering south to about 35
N in E America and to 45
N in W Palearctic. Vagrant south to Spain, Malta, Turkey and Cyprus, and to Bermuda. 2 old records from Morocco and Egypt, the latter rejected long ago (Cramp and Perrins 1994, Goodman and Meininger 1989). A very small, arboreal finch, Y with scarlet forehead, pinkish rump, pink cheeks, throat and breast and white belly, X with scarlet forehead and greyish white breast and belly; both sexes with finely dark-striped brown hindcrown, hindneck, mantle and back, blackish lores, neat black patch on chin, brown-striped flanks, 2 narrow white wing bars and yellow-horn bill. Length 115–145 cm; wing, Y 70–83 (776), X 70–80 (752). Inhabits damp willow, birch, alder and conifer woodland. Fast, steeply undulating, bouncy flight, often high up. Commonest calls, a 3–4 note dry rattle, ‘chu-chu-chu-chut’ and a nasal ‘dzooee’. TAXONOMIC NOTE: Lesser Redpoll C. cabaret (Britain and S Norway to Alps) is now recognized as a species separate from C. flammea (Knox et al. 2001, Sangster et al. 2002). African and

Mediterranean records of redpolls are probably (though not all certainly) of the more migratory flammea. For differences between the 2 species, see Knox et al. (2001).

Carduelis cannabina (Linnaeus). Common Linnet. Linotte me ´lodieuse.

Plate 32

Fringilla cannabina Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 182; Europe (Sweden).

(Opp. p. 475)

Range and Status. Palearctic: Canary Is, N Africa, Iberia and Ireland east to c. 90
E, north to 66
N in Sweden. Summer visitor in N half of range, winter visitor in Transcaucasia, much of Iran, Sinai and Nile Delta, elsewhere resident and partial migrant. Resident and partial migrant within NW Africa, and winter visitor from Europe. Morocco, frequent to common and widespread north of Haut Atlas, in Souss and to Atlantic and Mediterranean coasts; mainly in lowlands but up to 3000 m in Oukaı¨meden in summer and 3400 on Toubkal Massif in autumn; in winter south to Tiznit, Errachidia, Tarda, Bouanane, Erfoud, Merzouga, Tazzarine and L. Khnifiss. Algeria, breeds from coast south to Saharan Atlas, Djelfa and Aure`s; common in Tell; up to 1800 m in Djurdjura; absent from Aı¨n Sefra and from northernmost oases; in winter, south to Biskra, Ghardaı¨a (once 65 birds), Ouargla, El Gole´a, Beni Abbe`s (frequent), and once 100 in Bordj Omar Driss (¼ Temassinine). Tunisia, rather scarce resident, south to central plateau and to Gafsa; scarce to common winter visitor, south to Djerba. Libya, common on Tripoli coast, south locally to Jebel Nafusa in Oct–Mar (uncommon in July–Aug and Apr–May) in flocks of up to 100; evidently a small breeding population near Tripoli. In Cyrenaica, common at Sabratah, Homs, Leptis Magna, Sirt, Benghazi, Al Beyda, Cyrene and Apollonia in Apr (Massa 1999); frequent in

Carduelis cannabina

Carduelis cannabina Occurs irregularly in NW and N Africa in Oct–Apr (–May); however, a regular migrant at Gibraltar in Oct– Nov and Feb to early Apr, scarce in some years and numerous in others; conspicuous invasions in 1949–1950, 1959–1960, 1961–1962, 1965–1966, 1966–1967, 1972–1973, 1988–1989 (Finlayson 1992) and 1993–1994. Bird ringed in Germany recovered Morocco.

545

Food. Seeds, mainly of conifers. Not studied in Africa, but see above.

Key Reference Cramp, S. and Perrins, C.M. (1994).

Carduelis flammea (Linnaeus). Common Redpoll. Sizeron flamme´.

Plate 32

Fringilla flammea Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 182; Europe.

(Opp. p. 475)

Holarctic, breeding between 55
and 70
N; partial migrant, wintering south to about 35
N in E America and to 45
N in W Palearctic. Vagrant south to Spain, Malta, Turkey and Cyprus, and to Bermuda. 2 old records from Morocco and Egypt, the latter rejected long ago (Cramp and Perrins 1994, Goodman and Meininger 1989). A very small, arboreal finch, Y with scarlet forehead, pinkish rump, pink cheeks, throat and breast and white belly, X with scarlet forehead and greyish white breast and belly; both sexes with finely dark-striped brown hindcrown, hindneck, mantle and back, blackish lores, neat black patch on chin, brown-striped flanks, 2 narrow white wing bars and yellow-horn bill. Length 115–145 cm; wing, Y 70–83 (776), X 70–80 (752). Inhabits damp willow, birch, alder and conifer woodland. Fast, steeply undulating, bouncy flight, often high up. Commonest calls, a 3–4 note dry rattle, ‘chu-chu-chu-chut’ and a nasal ‘dzooee’. TAXONOMIC NOTE: Lesser Redpoll C. cabaret (Britain and S Norway to Alps) is now recognized as a species separate from C. flammea (Knox et al. 2001, Sangster et al. 2002). African and

Mediterranean records of redpolls are probably (though not all certainly) of the more migratory flammea. For differences between the 2 species, see Knox et al. (2001).

Carduelis cannabina (Linnaeus). Common Linnet. Linotte me ´lodieuse.

Plate 32

Fringilla cannabina Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 182; Europe (Sweden).

(Opp. p. 475)

Range and Status. Palearctic: Canary Is, N Africa, Iberia and Ireland east to c. 90
E, north to 66
N in Sweden. Summer visitor in N half of range, winter visitor in Transcaucasia, much of Iran, Sinai and Nile Delta, elsewhere resident and partial migrant. Resident and partial migrant within NW Africa, and winter visitor from Europe. Morocco, frequent to common and widespread north of Haut Atlas, in Souss and to Atlantic and Mediterranean coasts; mainly in lowlands but up to 3000 m in Oukaı¨meden in summer and 3400 on Toubkal Massif in autumn; in winter south to Tiznit, Errachidia, Tarda, Bouanane, Erfoud, Merzouga, Tazzarine and L. Khnifiss. Algeria, breeds from coast south to Saharan Atlas, Djelfa and Aure`s; common in Tell; up to 1800 m in Djurdjura; absent from Aı¨n Sefra and from northernmost oases; in winter, south to Biskra, Ghardaı¨a (once 65 birds), Ouargla, El Gole´a, Beni Abbe`s (frequent), and once 100 in Bordj Omar Driss (¼ Temassinine). Tunisia, rather scarce resident, south to central plateau and to Gafsa; scarce to common winter visitor, south to Djerba. Libya, common on Tripoli coast, south locally to Jebel Nafusa in Oct–Mar (uncommon in July–Aug and Apr–May) in flocks of up to 100; evidently a small breeding population near Tripoli. In Cyrenaica, common at Sabratah, Homs, Leptis Magna, Sirt, Benghazi, Al Beyda, Cyrene and Apollonia in Apr (Massa 1999); frequent in

Carduelis cannabina

Carduelis cannabina Occurs irregularly in NW and N Africa in Oct–Apr (–May); however, a regular migrant at Gibraltar in Oct– Nov and Feb to early Apr, scarce in some years and numerous in others; conspicuous invasions in 1949–1950, 1959–1960, 1961–1962, 1965–1966, 1966–1967, 1972–1973, 1988–1989 (Finlayson 1992) and 1993–1994. Bird ringed in Germany recovered Morocco.

545

Food. Seeds, mainly of conifers. Not studied in Africa, but see above.

Key Reference Cramp, S. and Perrins, C.M. (1994).

Carduelis flammea (Linnaeus). Common Redpoll. Sizeron flamme´.

Plate 32

Fringilla flammea Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 182; Europe.

(Opp. p. 475)

Holarctic, breeding between 55
and 70
N; partial migrant, wintering south to about 35
N in E America and to 45
N in W Palearctic. Vagrant south to Spain, Malta, Turkey and Cyprus, and to Bermuda. 2 old records from Morocco and Egypt, the latter rejected long ago (Cramp and Perrins 1994, Goodman and Meininger 1989). A very small, arboreal finch, Y with scarlet forehead, pinkish rump, pink cheeks, throat and breast and white belly, X with scarlet forehead and greyish white breast and belly; both sexes with finely dark-striped brown hindcrown, hindneck, mantle and back, blackish lores, neat black patch on chin, brown-striped flanks, 2 narrow white wing bars and yellow-horn bill. Length 115–145 cm; wing, Y 70–83 (776), X 70–80 (752). Inhabits damp willow, birch, alder and conifer woodland. Fast, steeply undulating, bouncy flight, often high up. Commonest calls, a 3–4 note dry rattle, ‘chu-chu-chu-chut’ and a nasal ‘dzooee’. TAXONOMIC NOTE: Lesser Redpoll C. cabaret (Britain and S Norway to Alps) is now recognized as a species separate from C. flammea (Knox et al. 2001, Sangster et al. 2002). African and

Mediterranean records of redpolls are probably (though not all certainly) of the more migratory flammea. For differences between the 2 species, see Knox et al. (2001).

Carduelis cannabina (Linnaeus). Common Linnet. Linotte me ´lodieuse.

Plate 32

Fringilla cannabina Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 182; Europe (Sweden).

(Opp. p. 475)

Range and Status. Palearctic: Canary Is, N Africa, Iberia and Ireland east to c. 90
E, north to 66
N in Sweden. Summer visitor in N half of range, winter visitor in Transcaucasia, much of Iran, Sinai and Nile Delta, elsewhere resident and partial migrant. Resident and partial migrant within NW Africa, and winter visitor from Europe. Morocco, frequent to common and widespread north of Haut Atlas, in Souss and to Atlantic and Mediterranean coasts; mainly in lowlands but up to 3000 m in Oukaı¨meden in summer and 3400 on Toubkal Massif in autumn; in winter south to Tiznit, Errachidia, Tarda, Bouanane, Erfoud, Merzouga, Tazzarine and L. Khnifiss. Algeria, breeds from coast south to Saharan Atlas, Djelfa and Aure`s; common in Tell; up to 1800 m in Djurdjura; absent from Aı¨n Sefra and from northernmost oases; in winter, south to Biskra, Ghardaı¨a (once 65 birds), Ouargla, El Gole´a, Beni Abbe`s (frequent), and once 100 in Bordj Omar Driss (¼ Temassinine). Tunisia, rather scarce resident, south to central plateau and to Gafsa; scarce to common winter visitor, south to Djerba. Libya, common on Tripoli coast, south locally to Jebel Nafusa in Oct–Mar (uncommon in July–Aug and Apr–May) in flocks of up to 100; evidently a small breeding population near Tripoli. In Cyrenaica, common at Sabratah, Homs, Leptis Magna, Sirt, Benghazi, Al Beyda, Cyrene and Apollonia in Apr (Massa 1999); frequent in

Carduelis cannabina

546

FRINGILLIDAE

coastal zone and parts of Jebel Akhdar in Oct–Apr, in flocks of c. 50 near Tobruk; thought to breed near Barce and Benghazi; elsewhere, large winter flocks near Hun and Sabha, and single bird at Serir. Egypt, fairly common and regular winter visitor from early Sept to early Apr along Mediterranean coast, in Nile Delta and Valley south to New Nubia; records from Suez Canal area, Faiyum, Wadi el Natrun, Bahariya, Dakhla and Safaga; flocks of 200, Kasfarit and Fayid; many freshly caught birds kept by bird dealers in Cairo and Luxor (in 1960s: Mathiasson 1972). Mauritania, irregular: Nouadhibou, up to 10, Mar 1985; Nouakchott, 11, Dec–Jan and 2, June; Tiguent, 1, Jan; Aleg, flocks of 5–10, Dec–Jan (Lamarche 1993). Senegal, 1, Richard-Toll, Mar. Sudan, frequent near 2nd Cataract in Feb–Mar 1964 when up to 7 birds on 10 dates near Sharta and singles or once ‘several’ on 6 dates at Genisab (Mathiasson 1972); records from Wadi Halfa and Khartoum (Nikolaus 1987). Breeding density of 6 pairs per km2 in maquis, Morocco (The´venot 1982). Description. C. c. mediterranea (Tschusi): Iberia to S Italy, Greece and Bulgaria. Not certainly known from N Africa, but certain to occur, at least in N Morocco, in winter, since a common autumn migrant at Strait of Gibraltar (Finlayson and Corte´s 1987). ADULT Y: forehead to hindneck grey-buff, streaked dark grey-brown, feathers on centre of forehead and forecrown with dark red central spot; mantle, back and scapulars chestnut-brown with narrow dusky streaks; rump and short uppertail-coverts pale grey-buff, the latter streaked dusky; long uppertail-coverts blackish with greyish white fringes. Tail feathers blackish, outer webs of T2–T6 with greyish white edges, broad basally, tapering and becoming toward tip. Side of head grey-brown, with short superciliary stripe and narrow stripe below eye pale greyish buff, contrasting with darker grey-brown stripe on lores and behind eye; poorly marked buffish patch on rear of cheek. Side of neck grey-brown with faint buffish mottling. Chin and throat pale buff, plain at sides, marked with short dark greyish streaks centrally; breast creamy white, each feather with partially concealed central rose-red spot; rest of underparts creamy white, flanks washed cinnamon-buff, undertail-coverts whiter with small narrow dusky streaks. In autumn, after moult, grey head and neck tinged buff and red on forehead and breast concealed by buff feather tips; buff feather edges produce more mottled pattern on mantle and scapulars. Flight feathers blackish, primaries with white outer edges, broad basally, secondaries broadly edged cinnamon-brown, tips fringed buffy white; tertials blackish brown, broadly bordered and tipped deep rufous- or chestnut-brown and narrowly fringed buff; primary coverts and alula blackish, fringed grey-brown; greater, median and lesser coverts rufous-brown, narrowly fringed buff. Underwing-coverts and axillaries greyish white. ADULT X: forehead to hindneck streaked blackish brown and buff; mantle, scapulars, back and rump streaked tawny brown, blackish and buffy brown; uppertail-coverts dark brown, edged pale buff. Sides of head as in ad. Y, but browner (less grey). Underparts pale buff, tinged brown on breast and flanks, grading to creamy white on belly and undertail-coverts; centre of chin and throat with short dark grey-brown streaks, breast and flanks streaked dark brown. Tail and wings as in ad. Y, but borders of tertials and greater, median and lesser coverts tawny- or olivebrown rather than rufous. SIZE (10 YY, 6 XX, N Africa, Mar– May): wing, Y 74–80 (774), X 74–78 (760); tail, Y 51–54 (525), X 48–54 (515); bill, Y 115–135 (128), X 125–13 (127); tarsus, Y 145–16 (156), X 15–16 (156). IMMATURE: juv. similar to ad. X, but buffy brown (not greyish) on head, and paler, more buffish above; streaks on breast and flanks shorter and narrower.

NESTLING: down fairly long and plentiful, smoke grey, mainly on head, back, upperwing and thigh; mouth bright rose-red, inside edges of upper mandible tinged light blue-grey to whitish, gape flanges pale yellowish. C. c. cannabina (Linnaeus): Europe and W Siberia, south to Pyrenees, N Italy, Ukraine and central Kazahkstan; winters throughout N Africa. Like mediterranea but slightly darker and larger: wing, Y (n ¼ 75) 78–85 (808). WEIGHT: Sudan, 3, unsexed, 163, 170, 185, W Europe, Y (n ¼ 27) 159–257 (202), X (n ¼ 52) 151–205 (189). Unnamed subspecies (C.S. Roselaar in Cramp and Perrins 1994): Morocco to Libya. Like mediterranea but distinctly paler, and longer- and deeper-billed; bill to skull, Y (n ¼ 7) 127–139 (132), bill to nostril, Y (n ¼ 7) av. 03 mm longer than Y mediterranea. C. c. bella (C.L. Brehm): Crimea, Turkey and Levant to N Iraq, Iran and SW Turkmeniya; also central Asia; winters Egypt. Paler than nominate race, grey of head and neck lighter, mantle and scapulars more cinnamon-brown; lower rump and uppertailcoverts whitish with narrower streaking; bill length and depth as in unnamed NW African race (C.S. Roselaar in Cramp and Perrins 1994). Wing, Y (n ¼ 5) 79–84 (810). WEIGHT: Y (n ¼ 11) 18–22 (198), X (n ¼ 5) 175–210 (192).

Field Characters. Length 13–135 cm. A small finch with grey head and brown back, readily identified overhead by dry twittering flight call; in flight shows white in primaries and at base of tail. Breeding Y has red patches on forehead and at sides of breast, rich chestnut back; in winter lacks red, back and underparts streaked, breast rich pinkish buff. X and juv. like winter Y but duller, rather nondescript except for white primary edges showing on folded wing. Voice. Tape-recorded (62–73, 76–89, 105, 110–113, B, C). Musical song rapid and continuous (up to 17 s), sweet pure notes mixed with dry twitters and churrs, often starting with ‘chita, chita’, like flight call; some notes have twangy, tweaky quality, others shrill, some buzzy; very variable. Flight call a distinctive dry ‘tuk’ or ‘tututuk’, higherpitched and more clipped than similar calls of Common Redpoll C. flammea or European Greenfinch C. chloris. Main contact call a descending whistle, ‘peeo’ or ‘plee-yu’; alarm, plaintive ‘tsooeet’. For full treatment of extensive repertoire, see Cramp and Perrins (1994). General Habits. Inhabits coasts, lowlands, and in Algeria the highest upland meadows; cultivated fields with thickets and hedgerows; sunny scrub, heath, young plantations, orchards, maquis; in winter, salt marshes, foreshores, farmyards and weedy waste ground; avoids dense woods, forest and towns. In pairs in breeding season and mainly small, tight flocks out of it, sometimes mixing with other finches. In winter in Morocco occurs in flocks of dozens or less commonly in hundreds. Roosts communally; several hundred in trees in Foire de Mekne`s and 2000 near Sidi Yahia des Zae¨rs, Morocco, in Oct–Nov. Winter flocks in Libya and Egypt usually of 50 kg can be applied by powerful masseter muscles. Palearctic; migratory. No close relatives in W Palearctic or Africa; perhaps most closely related with grosbeaks Eophona of China and Japan and more distantly with 4 E Palearctic yellow and black grosbeaks Mycerobas and 2 American ones Hesperiphona.

Coccothraustes coccothraustes (Linnaeus). Hawfinch. Gros-bec cassenoyaux.

Plate 33

Loxia coccothrasutes Linnaeus, 1758. Syst. Nat., ed. 10, p. 171; southern Europe.

(Opp. p. 522)

Range and Status. Palearctic, from NW Africa, Portugal and Britain to Manchuria, Sakhalin I. and Japan. Summer visitor in NE Europe and Asia, partial migrant and resident in W Europe, resident and winter visitor in Spain, Italy, Turkey and NW Africa. Accidental Iceland, Faeroes and Madeira. Resident and winter visitor (from Europe). Morocco, uncommon (or uncommonly seen: c. 12 records per year), in wooded valleys in Rif region (Ouezzane and to east), in Moyen Atlas (Ifrane, Azrou, Dayet Afourgah, Taroudant, Azigza, Khe´nifra, Hachlaff, Aguelmane, Afenourir, Jaaba forest) and Haut Atlas (Aı¨t Ourir, Tizi-n-Test, Ourika Valley, Toubkal, Ida-ou-Tenane, Nfis, Asni and Imlil valleys, Souss at Igli); most frequent around Mekne`s, Sidi

Bettache forest and Rabat; once 12, Oulmes, May. Algeria, breeds in Aure`s, Constantine area, El Kala, N slopes of Tell Atlas, Grande Kabylie, Babor (up to 1900 m), Djurdjura (rare), Algiers area, Ouarsenis and Tlemcen Mts; more or less regular winter visitor from Europe; in winter local birds move erratically and descend mountains (Ledant et al. 1981, Isenmann and Moali 2000). Tunisia, scarce resident, high central and W Kroumirie Mts, and rare winter visitor, mainly in N in Oct–Nov and Mar–Apr, very rare south as far as Gabe`s and Djerba. Libya, rare winter visitor to Tripoli coast, Oct–Mar, south to Beni Ulid and Jebel Nafusa where once a flock of 12; Cyrenaica, once, Al Adem. Egypt, rare, although long ago a marked passage noted on Mediterranean coast in Nov–early Dec

Coccothraustes coccothraustes

559

Genus Pyrrhula Brisson

6 species of grey and black Palearctic finches, YY with pink or orange, XX mainly buffy; white rumps; purple-glossed wings with prominent wing bar; 2 spp. with square tails and 4 with short central and progressively lengthy outer rectrices. Bill very short and deep, culmen strongly curved; adapted for feeding on buds. Mainly Himalayan-Chinese; 1 in Philippines; 1 from Azores to Japan, vagrant to NW Africa.

Pyrrhula pyrrhula (Linnaeus). Eurasian Bullfinch. Bouvreuil pivoine.

Plate 33

Loxia pyrrhula Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 171; Europe.

(Opp. p. 522)

Breeds from Azores, N Spain, N Portugal, Italy, Balkans, N Turkey and Caucasus to Kamchatka and Japan. Partial winter migrant south to S Portugal, S-central Spain, S Italy, Greece and S Turkey. Several records Gibraltar. Vagrant in NW Africa. Morocco, 5 records, of 1–4 birds, 1973–1987, south to Settat, 33
040 N, and Mekne`s, 33
530 N (Bergier et al. 2000). Algeria, reported Dec 1859. Tunisia, ‘important number of birds’ in 1886 (Thomsen and Jacobsen 1979). Length 145–16 cm. Stout, with thick neck and flattish head and short, deep bill. Black cap, chin and tail, black wings with white bar, white undertail-coverts. Underparts pinkish red in Y, buffy in X. White rump conspicuous in flight. Immature like X but with no black on head. Rather shy, keeping mainly to thick cover, where best located by call, a distinctive soft melancholy ‘whew’.

Genus Coccothraustes Brisson

Single species of large grosbeak, with large head, massive bill, short white-tipped tail, and inner secondaries with expanded, cleaver-shaped tips (A). Bill adapted for splitting large, woody nuts; seed is held by 2 striated knobs at base of each mandible, through which a force of >50 kg can be applied by powerful masseter muscles. Palearctic; migratory. No close relatives in W Palearctic or Africa; perhaps most closely related with grosbeaks Eophona of China and Japan and more distantly with 4 E Palearctic yellow and black grosbeaks Mycerobas and 2 American ones Hesperiphona.

Coccothraustes coccothraustes (Linnaeus). Hawfinch. Gros-bec cassenoyaux.

Plate 33

Loxia coccothrasutes Linnaeus, 1758. Syst. Nat., ed. 10, p. 171; southern Europe.

(Opp. p. 522)

Range and Status. Palearctic, from NW Africa, Portugal and Britain to Manchuria, Sakhalin I. and Japan. Summer visitor in NE Europe and Asia, partial migrant and resident in W Europe, resident and winter visitor in Spain, Italy, Turkey and NW Africa. Accidental Iceland, Faeroes and Madeira. Resident and winter visitor (from Europe). Morocco, uncommon (or uncommonly seen: c. 12 records per year), in wooded valleys in Rif region (Ouezzane and to east), in Moyen Atlas (Ifrane, Azrou, Dayet Afourgah, Taroudant, Azigza, Khe´nifra, Hachlaff, Aguelmane, Afenourir, Jaaba forest) and Haut Atlas (Aı¨t Ourir, Tizi-n-Test, Ourika Valley, Toubkal, Ida-ou-Tenane, Nfis, Asni and Imlil valleys, Souss at Igli); most frequent around Mekne`s, Sidi

Bettache forest and Rabat; once 12, Oulmes, May. Algeria, breeds in Aure`s, Constantine area, El Kala, N slopes of Tell Atlas, Grande Kabylie, Babor (up to 1900 m), Djurdjura (rare), Algiers area, Ouarsenis and Tlemcen Mts; more or less regular winter visitor from Europe; in winter local birds move erratically and descend mountains (Ledant et al. 1981, Isenmann and Moali 2000). Tunisia, scarce resident, high central and W Kroumirie Mts, and rare winter visitor, mainly in N in Oct–Nov and Mar–Apr, very rare south as far as Gabe`s and Djerba. Libya, rare winter visitor to Tripoli coast, Oct–Mar, south to Beni Ulid and Jebel Nafusa where once a flock of 12; Cyrenaica, once, Al Adem. Egypt, rare, although long ago a marked passage noted on Mediterranean coast in Nov–early Dec

560

FRINGILLIDAE

Coccothraustes coccothraustes

(Lynes 1912); 1, Burg el Arab, 1938, 3 there 1980; 4, Bahig, in 1960s and 1970s; 3, Wadi el Natrun, 1914; 1, Siwa, 1927; Alexandria, 1996. Breeding densities in Morocco of 196 pairs per km2 of broad-leaved forest and 15 pairs per km2 of evergreen oak woodland (The´venot 1982). Description. C. c. buvryi Cabanis: NW Africa. ADULT Y: forehead pale buff, grading through buffy brown forecrown to rich umberbrown hindcrown and nape; hindneck, sides of neck and upper mantle grey; lower mantle and scapulars dark umber-brown; back grey-brown, merging into pale cinnamon-brown rump and uppertail-coverts. Central tail-feathers (T1) pale brown on outer web, pale grey-brown on inner web, tips whitish; T2 grey-brown on outer web, inner web black with broad white tip; T3–T5 blackish with tip of outer web pale grey-brown, and broad white tip on inner web, increasing to almost half feather length on T5; T6 blackish with distal half of inner web white. Lores black, extending as narrow band around mandible base to bib, and connecting with very narrow band at side of maxilla base, and with black eye-ring. Cheeks and ear-coverts buffy brown; chin and centre of throat black, forming small bib, bordered narrowly at sides and below with whitish; breast and flanks pale cinnamongrey; belly, thighs and undertail-coverts white. Primaries blackish brown, tips of P2–P5 glossed violet and cleaver-shaped, tips of P7–P9 glossed greenish; secondaries blackish, broad distal outer edges and tips glossed violet, S1–S4 square-tipped (A). Middle part of inner webs of primaries white, the area larger on inner feathers, extending along inner edge toward base; middle of inner webs of secondaries with smaller white lateral patch, not reaching shaft, but extending along inner edge toward base. Outer tertial blackish, tipped brown; middle and inner tertials umber-brown. Primary coverts and alula black, narrowly edged glossy greenish. Outer greater coverts pale grey on outer web, with buff tip, dark brown on inner web; inners blackish on both webs, with cinnamon-buff tip; median coverts dark brown, broadly tipped

pale grey-buff. Median covert tips and greater covert outer edges form long whitish panel on closed wing. Lesser coverts blackish brown, fringed umber-brown. Primary underwing-coverts blackish brown; rest of underwing-coverts and axillaries white. Bill slate-blue in breeding season, underside of mandible pinkish or flesh; in rest of year, yellowish or greyish horn. Eyes amber, light cinnamon or reddish brown; legs flesh-pink, tinged lilac in breeding season. ADULT X: similarly patterned to ad. Y, but differs in having top of head pale grey, grading to darker greybrown on nape; black of lores and bib tinged brown; back to rump pale grey-brown, tinged buff; white tail spots smaller, with narrow fringe only on T1, white on inner web of T6 occupying distal quarter; underparts pale grey, tinged brown, most strongly on sides and flanks; broad distal outer borders of secondaries pale glossy lilac-grey (not violet-black); pale wing panel greyer, lesser covert tips grey-brown, underwing-coverts greyish white. SIZE (10 YY, 10 XX): wing, Y 95–102 (982), X 95–99 (967); tail, Y 52–58 (550), X 51–55 (528); bill, Y 23–25 (242), X 21–24 (227); tarsus, Y 20–225 (214), X 20–215 (209). WEIGHT: 1, unsexed, Morocco, 550. IMMATURE: juv. mottled brown above, rump buffier; side of head pale brown, flecked buff, with distinct paler fore supercilium and blackish malar stripe; underparts whitish, breast, flanks and belly mottled grey-brown. Y has whole head tinged olive-yellow, most strongly on forehead and malar area; X has slight yellow tinge, confined to forehead and malar area. Tail as ad., but T1 fringed tawny buff and lacks white tip; wing as ad. of same sex, but tertials and inner greater coverts more buff, median covert tips tinged buffish yellow, lesser coverts fringed tawny-buff. NESTLING (nominate race): down white, long and dense on upperparts and upperwing, shorter on belly, thighs and flanks; mouth and tongue red, surrounded by purple-blue, centre of tongue pink with whitish spurs; gape flanges bright yellow (Cramp and Perrins 1994). C. c. coccothraustes (Linnaeus): Eurasia east to W Mongolia; visitor to N Africa. Y differs from buvryi in having forehead yellower, nape more orange-brown, mantle darker umber brown, rump brighter cinnamon or tawny-buff; sides of head richer tawny-brown; underparts deeper cinnamon (less grey). X has top and sides of head browner than in buvryi, mantle darker, rump tinged cinnamon, breast and flanks browner. Larger; Y (n ¼ 10), wing 100–107 (103), bill 24–27 (252). WEIGHT: (central Europe) Y (n ¼ 99) 46–72 (580), X (n ¼ 53) 46–65 (553).

Field Characters. Length 18 cm. A large finch with unmistakable shape – massive bill, heavy head, thick neck, plump body and short tail. Plumage pattern also unique: yellow-brown head, grey nape and hind-collar, black lores and bib, dark brown back, pinkish brown underparts; large white wing-patch visible on black wing at rest; bill dark in summer, pale in winter. Shows broad white tip to tail when flushed, 2 white wing-patches in flight. Flying bird appears

Coccothraustes coccothraustes top-heavy, with whirring wings and undulating motion. Feeds quietly in treetops and easily overlooked, presence often indicated by loud ‘tick’ call, also by split stones of seeds below bushes (Beaman and Madge 1998). Voice. Tape-recorded (62–73, 76–89, 105, 110–113, B, C). Song a rambling series of brief sharp and buzzy notes, e.g. ‘tzit . . . tzeeep . . . tsit . . . tsipit . . . bzzzt . . . tsit . . . tzeep . . . tzi-tzeep . . . tsit’. Subsong (Chappuis 2000) low, rather sweet whistles mixed with grating ‘churk’ and ‘jreeo’. Voice lower-pitched in Morocco than in France (Chappuis 1969). Flight call an explosive ‘zick’ given with each upward bound; contact calls include thin ‘sreee’ or ‘seep’ and quiet ‘tick’ or ‘tack’; alarm a single loud ‘tzic’. For sonagrams and additional repertoire, see Cramp and Perrins (1994). General Habits. Inhabits mature, open, mixed broadleaved woods of oak Quercus, particularly cork oak Q. suber, alder Alnus and hornbeam Carpinus, and associates also with such hard-fruit-bearing trees as beech Fagus, wych elm Ulmus, ash Fraxinus and sycamore Acer. In general avoids pinewoods, although winter migrants sometimes feed in pines Pinus. Ascends mountains up to limit of deciduous forest. Occurs in orchards and large gardens, but avoids artefacts of all kinds (buildings, fences, walls, wires). Keeps in pairs and small flocks; shy, unobtrusive, secretive. Forages equally in tree tops and on ground. On ground stands upright and moves clumsily with heavy hops; in trees, carriage more horizontal, bird moving along branches and through twigs with twisting waddle. Birds in small flock on ground are generally several m apart but act in unison. Extremely wary, nervous and easily disturbed; when one is alerted all stop feeding, look up, sleek head plumage then fly off synchronously, rocketing silently up and away with rapid bursts of wing beats into tree canopy. Single bird or pair flies high (50 m) over trees or very high (300 m) over open space, when flight is bounding then deeply undulating. Drinks regularly at surface puddles. Extracts seeds from fleshy fruits by turning fruit in bill to peel soft parts off against lower mandible. Carefully nibbles large seed in bill to position it before biting to crack it. Picks small snails from leaves and crushes them in bill. Rips scales off growing cone of pine Pinus sylvestris, then lands on ground to eat fallen seeds. NW African breeding birds sedentary or make only local movements, but joined in winter by considerable numbers of migrants; ringing recoveries in N Algeria from France (1), Germany (1), Italy (2) and Switzerland (1). Greatest distance 1950 km (Slovakia to S Spain). Regular daytime migrant across Strait of Gibraltar in Oct/early Nov and Mar/early Apr, although numbers fluctuate from year to year (Finlayson 1992). Food. Large, hard seeds; buds, shoots; quite large insects and some snails in breeding season. In Mediterranean eats seeds of olive Olea europaea, nettle-tree Celtis sp., oaks Quercus spp. (acorns) and sunflowers Helianthus; also eats hornbeam nuts, beech nuts, cherry stones, and small seeds picked from thrush droppings.

Breeding Habits. Solitary nester, sometimes in loose colony (once 5 nests in radius of 27 m); monogamous; defends only immediate surroundings of nest. Y has remarkable display flight, ‘rollercoasting’ over woods then plummeting down at steep angle. Perched Y sings infrequently; song not loud and not given from special song posts. In courtship display, Y waddles with body upright, swivelling from side to side, head retracted, and wings held loosely, drooping, to show white shoulders (the Penguin-walk); he then bows deeply in front of X. Y chases X in flight; he courtship-feeds her. NEST: open cup, with foundation of dry twigs, distinct lining of thin twiglets and thin, dry grass blades, and inner lining of rootlets, soft plant matter, thin grass and occasionally hair. Ext. diam. 200–220, ext. depth 90–110, int. diam. 70–80, int. depth 40–50. Sited mainly in old, shrubby oak trees, often in ivy or honeysuckle covering trunk. Nest-building started by Y, completed by X. EGGS: 3–5, av. (n ¼ 20 clutches, NW Africa) 38. Pale blue or greyish green, sparsely marked with dark spots and scrawls, and sometimes blackish scribbling mainly at broad end. SIZE: (C. c. buvryi, n ¼ 20) 201–245  154– 177 (238  174). WEIGHT: 377. LAYING DATES: NW Africa, late Mar to late May. INCUBATION: by X, starting with 3rd egg. Period 9–14 days, usually 11–13. DEVELOPMENT AND CARE OF YOUNG: young brooded by X only, at night and for spells during day. At first Y passes food to X who distributes it to brood; later Y and X feed young directly. Both parents remove faecal sacs, swallowing them or dropping them up to 50 m away from nest. Nestling period: (n ¼ 19) av. 125 days (Europe). After leaving nest, young cared for by both parents separately or jointly and do not become independent for a long time BREEDING SUCCESS/SURVIVAL: success in colonies much greater than in solitary pairs (Cramp and Perrins 1994); predators (Europe) include European Sparrowhawk Accipiter nisus, Eurasian Jay Garrulus glandarius, Common Magpie Pica pica, Carrion Crow Corvus corone and squirrels Sciurus spp. Oldest ringed bird 11 years 8 months; lives up to 18 years in captivity.

Key References Cramp, S. and Perrins, C.M. (1994), Etche´copar, R.D. and Hu ¨ e, F. (1967), Goodman, S.M. and Meininger, P.L. (1989), Isenmann, P and Moali, A. (2000), Mountfort, G.R. (1957), The´venot, M. (1982).

561

562

FRINGILLIDAE

Genus Loxia Linnaeus

A distinctive, Holarctic cardueline genus: YY red-orange or orange-grey, XX olive or grey, juvs streaked. Bill robust, deep and broad-based, asymmetric, with tips of mandibles crossed, elongated in L. curvirostra and L. scotica, tip of upper mandible closing to left or right of tip of lower mandible. Bill is adapted to husking cones of conifers on which the birds feed: L. curvirostra on Picea and Pinus helepensis, L. scotica on Pinus sylvestris, L. pytyopsittacus on P. sylvestris and Picea, and L. leucopterus on Larix and Picea. 4 species, 1 N European, 1 Scottish and 2 Holarctic. The last 2 are resident but irruptive; one of them breeds south to NW Africa.

Plate 33

Loxia curvirostra Linnaeus. Common Crossbill. Bec-croise´ des sapins.

(Opp. p. 522)

Loxia curvirostra Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 171; Europe (Sweden). Range and Status. Holarctic, breeding widely between 45
and 65
N. In W Palearctic breeds south to NW Africa, Crete and Cyprus, west to N Portugal and N Scotland, and north to 69
in Finland. Dispersive and irruptive, with huge movements from N Asia and N Europe to west and southwest in autumn, every decade or so, birds reaching Iceland, Greenland, Madeira, Malta, Gibraltar and N Africa. Mediterranean and NW African populations resident, wandering locally. 05–50 million in Scotland in 1990 irruption. 5 W Palearctic subspecies, varying clinally: curvirostra in Europe (probably this irrupting into N Africa: see below), 1 in Cyprus, Turkey and eastward, 1 endemic to Balearic Is, 1 to Corsica and poliogyna to NW Africa. Morocco, resident in Haut Atlas (Tizi-n-Test, Ouirgane, Asni, Tahanaout, Ourika valley, Aı¨t Ourir near Marrakech, and Tizi-n-Tichka), Middle Atlas, and pine-clad hills and valleys between Haut Atlas and Mediterranean coast: Imouzer du Kandahar, Mekne`s, Fe`s, Jebel Lakhdar (¼ Lakraa) near Tetouan and east to Oujda. In E, only 4 populations, well separated, at Lalla Mimouna at 1200 m, Foreˆt de Draouch at 800 m, Beni Ourimech at 600 m and Taforalt at 800 m (Brosset 1957). Evidently much given to wandering, and reported generally to be uncommon, irregular and local. Occurs all year in Moulay-IdrissZerhoum region and in Aı¨t Ourir, where numerous. Out of the breeding season, flocks of 10–30–140 in Oujda, Korifla, Beni-Snassen and elsewhere. Up to 20 caged birds at a time for sale, illegally, in Mekne`s (El He´dim) and Me´dina de Fe`s bird markets. Algeria, widespread from Algiers, Grande Kabylie and coastal woodlands southwards; formerly widespread, now seemingly local and uncommon, in Atlas Saharien (Mts des Ksour, Jebel Amour, Aı¨n Sefra, Oulad Naı¨l Mts); in land below 1000 m from Monts de Tlemcen to Massif de l’Ouarsenis; local in Chettaba Forest near Constantine but absent from cedars and pines on Mt Babor; widespread in Aure`s Mts and east to Te´bessa and W Monts de la Medjerda. Status as in Morocco, although sufficiently common for 77 nests to have been found in a 3 year study. Tunisia, uncommon resident; along Algerian border from Jendouba to Bou Chebka; commonest in Thala-Jebel Chamba-Bou Chebka hills.

Loxia curvirostra

Breeding densities unknown in Africa; in pine forests, 4 pairs per km2 in Vosges, France, 34 in Italian Alps and, over 5 years, 14–40 in Pyrenees. Description. L. c. poliogyna Whitaker: Morocco to Tunisia. ADULT Y: top of head grey-brown with broad orange-red feather tips; hindneck, sides of neck, mantle and scapulars grey-brown with narrow orange-red feather fringes; back, rump and uppertailcoverts with broad orange-red feather tips masking grey bases. Tail feathers blackish brown, outer webs with fine red-brown or olive-brown fringes when fresh. Broad band through lores and behind eye dark grey-brown; lower ear-coverts and cheeks greybrown with narrow feather tips forming orange-red barring; below this a darker grey-brown malar band. Chin grey-buff, tinged red and mottled dusky grey; throat to breast and flanks grey-brown, sides of breast with orange-red feather fringes, throat, centre of breast and upper flanks with broad orange-red

562

FRINGILLIDAE

Genus Loxia Linnaeus

A distinctive, Holarctic cardueline genus: YY red-orange or orange-grey, XX olive or grey, juvs streaked. Bill robust, deep and broad-based, asymmetric, with tips of mandibles crossed, elongated in L. curvirostra and L. scotica, tip of upper mandible closing to left or right of tip of lower mandible. Bill is adapted to husking cones of conifers on which the birds feed: L. curvirostra on Picea and Pinus helepensis, L. scotica on Pinus sylvestris, L. pytyopsittacus on P. sylvestris and Picea, and L. leucopterus on Larix and Picea. 4 species, 1 N European, 1 Scottish and 2 Holarctic. The last 2 are resident but irruptive; one of them breeds south to NW Africa.

Plate 33

Loxia curvirostra Linnaeus. Common Crossbill. Bec-croise´ des sapins.

(Opp. p. 522)

Loxia curvirostra Linnaeus, 1758. Syst. Nat., ed. 10, 1, p. 171; Europe (Sweden). Range and Status. Holarctic, breeding widely between 45
and 65
N. In W Palearctic breeds south to NW Africa, Crete and Cyprus, west to N Portugal and N Scotland, and north to 69
in Finland. Dispersive and irruptive, with huge movements from N Asia and N Europe to west and southwest in autumn, every decade or so, birds reaching Iceland, Greenland, Madeira, Malta, Gibraltar and N Africa. Mediterranean and NW African populations resident, wandering locally. 05–50 million in Scotland in 1990 irruption. 5 W Palearctic subspecies, varying clinally: curvirostra in Europe (probably this irrupting into N Africa: see below), 1 in Cyprus, Turkey and eastward, 1 endemic to Balearic Is, 1 to Corsica and poliogyna to NW Africa. Morocco, resident in Haut Atlas (Tizi-n-Test, Ouirgane, Asni, Tahanaout, Ourika valley, Aı¨t Ourir near Marrakech, and Tizi-n-Tichka), Middle Atlas, and pine-clad hills and valleys between Haut Atlas and Mediterranean coast: Imouzer du Kandahar, Mekne`s, Fe`s, Jebel Lakhdar (¼ Lakraa) near Tetouan and east to Oujda. In E, only 4 populations, well separated, at Lalla Mimouna at 1200 m, Foreˆt de Draouch at 800 m, Beni Ourimech at 600 m and Taforalt at 800 m (Brosset 1957). Evidently much given to wandering, and reported generally to be uncommon, irregular and local. Occurs all year in Moulay-IdrissZerhoum region and in Aı¨t Ourir, where numerous. Out of the breeding season, flocks of 10–30–140 in Oujda, Korifla, Beni-Snassen and elsewhere. Up to 20 caged birds at a time for sale, illegally, in Mekne`s (El He´dim) and Me´dina de Fe`s bird markets. Algeria, widespread from Algiers, Grande Kabylie and coastal woodlands southwards; formerly widespread, now seemingly local and uncommon, in Atlas Saharien (Mts des Ksour, Jebel Amour, Aı¨n Sefra, Oulad Naı¨l Mts); in land below 1000 m from Monts de Tlemcen to Massif de l’Ouarsenis; local in Chettaba Forest near Constantine but absent from cedars and pines on Mt Babor; widespread in Aure`s Mts and east to Te´bessa and W Monts de la Medjerda. Status as in Morocco, although sufficiently common for 77 nests to have been found in a 3 year study. Tunisia, uncommon resident; along Algerian border from Jendouba to Bou Chebka; commonest in Thala-Jebel Chamba-Bou Chebka hills.

Loxia curvirostra

Breeding densities unknown in Africa; in pine forests, 4 pairs per km2 in Vosges, France, 34 in Italian Alps and, over 5 years, 14–40 in Pyrenees. Description. L. c. poliogyna Whitaker: Morocco to Tunisia. ADULT Y: top of head grey-brown with broad orange-red feather tips; hindneck, sides of neck, mantle and scapulars grey-brown with narrow orange-red feather fringes; back, rump and uppertailcoverts with broad orange-red feather tips masking grey bases. Tail feathers blackish brown, outer webs with fine red-brown or olive-brown fringes when fresh. Broad band through lores and behind eye dark grey-brown; lower ear-coverts and cheeks greybrown with narrow feather tips forming orange-red barring; below this a darker grey-brown malar band. Chin grey-buff, tinged red and mottled dusky grey; throat to breast and flanks grey-brown, sides of breast with orange-red feather fringes, throat, centre of breast and upper flanks with broad orange-red

Loxia curvirostra feather tips largely masking the grey; belly and undertail-coverts light grey-brown, the latter with dusky grey-brown centres showing as dark mottling. Upperwing feathers blackish brown, slightly glossy; primaries finely edged pinkish buff or grey-buff; tertials and coverts tinged olive-green, with edges of greater coverts, tips of median coverts and broader tips to lesser coverts reddish brown. Underwing-coverts grey, tinged red near carpal edge; axillaries greyish white. Bill with curving mandibles crossing at tip, grey or greyish horn; eyes dark brown; legs dark brown or grey-brown. ADULT X: top of head, hindneck, sides of neck, mantle, back and scapulars pale olive-grey with diffuse dusky mottling; back and rump pale olive-green; uppertailcoverts grey-brown, shorter feathers fringed pale olive-green. Tail feathers dark grey-brown, finely edged olive-green. Side of head light grey-brown, darker on lores, behind eye and in malar area. Underparts light grey-brown, breast and flanks tinged olivegreen, belly and undertail-coverts paler, the latter with dusky mottling. Upperwing feathers dark grey-brown, primaries finely edged pale grey-green, tertials and coverts tinged olive-green. Bare parts as ad. Y. SIZE: wing, Y (n ¼ 12) 95–103 (991), X (n ¼ 16) 93–101 (962); tail, Y (n ¼ 12) 54–62 (582), X (n ¼ 13) 51–57 (546); bill, Y (n ¼ 10) 21–25 (230), X (n ¼ 16) 21–25 (229); tarsus, Y (n ¼ 12) 18–19 (186), X (n ¼ 14) 18–19 (185) (Cramp and Perrins 1994). WEIGHT: X (n ¼ 1) 377. IMMATURE: juv. pale grey-buff top of head, merging with light olive-brown mantle and scapulars and pale yellowish buff rump, the entire upperparts streaked dark brown; side of head and underparts buffy white, breast and flanks washed olive-brown, all boldly and extensively streaked dark brown; tail feathers finely edged buff; primaries edged and tipped buff, tertials narrowly fringed buff and greater and median coverts tipped buff to form 2 narrow wing-bars. 1st-year immature male like female but top of head, mantle and scapulars tinged olive-yellow or orange-yellow, rump predominantly light orange or orange-yellow; underparts washed olive-green (usually with some orange), especially on throat, breast and upper flanks. NESTLING (nominate curvirostra): down very dark grey, on upperparts, upperwing and thigh. (L. c. curvirostra Linnaeus: Europe south to N Spain, central Italy and Balkans; east to Siberia. Strongly irruptive; in 1990, 500,000 birds invaded Scotland, with many in Faeroes, 800 in Iceland, and (presumably this race) 50 in Malta and 78 in Gibraltar; no irruptive birds were noted in NW Africa, but 2 presumed to be this race in Morocco in 1855 and Algeria in 1909, and several in NE Tunisia in Aug 1930–Mar 1931, coinciding with great irruption into Europe southwest to Spain. Y bright red, summer and winter, except for blackish wings and tail; X more olive, less greyish, than X poliogyna. Intergrades with poliogyna in S Italy and Sicily. Bill smaller but wings longer than in poliogyna (Questiau et al. 1999). WEIGHT: (Scotland) Y (n ¼ 10) 37–45 (407), X (n ¼ 11) 37–46 (405).) TAXONOMIC NOTE: taxonomy of the Loxia curvirostra group of species throughout the Holarctic is complex; in relation to it, voices of NW African populations merit further investigation (Robb 2000).

Field Characters. Length 16–17 cm. Plump, with large head and short, deeply forked tail. Crossed bill diagnostic but hard to see as bird feeds high in conifer. Y mainly brick-red, brightest on rump (variable; some more orange, others mottled); X drab-looking, olive-green with yellowish rump conspicuous in flight. Unobtrusive and often out of sight among pine branches; usually first spotted by ‘jipjip’ call, frequently given in flight and in more subdued version while feeding. Voice. Tape-recorded in Europe (62–73, 76–89, 105, 110– 113, B, C); not tape-recorded in Africa and vocalizations of

African birds not described. In Europe, song a pleasant, disjointed medley of whistles, chirps, twitters and scratchy notes, including ‘jip’ calls; in alarm, ‘jip’ repeated to become hard chatter. Both song and call types vary widely in Europe (Robb 2000), and African birds may well have distinctive calls. General Habits. Feeds and nests in crowns of conifers, mainly Aleppo pines Pinus halepensis; inhabits forests, copses and isolated trees of those species that provide plant food, principally in well-wooded foothills and mid altitudes: spruce, juniper, Thuja, firs Abies numidica and Pseudotsuga, cedar and larch and, less commonly, almond, walnut, birch, poplar, alder, hornbeam, beech, oak, hazel and others; at 600–2200 m. Roosts communally in tall, dense conifer. Gregarious all year, feeding in flocks of 10–15 even in breeding season; afterwards family parties gather in flocks of up to 32 birds or more (Morocco). Roosts sometimes in mixed finch flocks; sometimes flocks by day with Eurasian Siskin Carduelis spinus. Very agile and acrobatic when extracting seeds from pine cones; flutters from twig to twig, sidles along branch to its cone-bearing tip, sometimes moving along with help of bill, like a parrot; either nips a cone off and carries it in bill to a more rigid perch or (generally) extracts seeds in situ, hanging under drooping cone, inserting bill between cone’s scales from the side and moving lower mandible from side to side to loosen seed, which is scooped out with tongue (Cramp and Perrins 1994). Entire cone is commonly stripped of its scales and seeds, scales and remains of cone being dropped and littering ground beneath fruiting tree. Bird can carry a cone as large as itself and can hold loose cone against underside of branch with feet and extract seeds whilst working upside-down; but usually carries snipped-off cone to top of horizontal branch or to fork, holding it under foot and standing on top of it to remove seeds. In 115 observations of feeding in central Spain, 96% were on cones, 2% on branches and 2% on ground. Extracts seeds from large fruits such as apples. Forages mainly higher than 16 m up in conifers. In Europe, wanders quite widely in search of food within regular range, so that local abundance fluctuates widely from season to season and year to year; in NW Africa, irregularity of records at many localities suggests similar vagrancy. Food. Seeds of conifers listed above; seeds, fruits and buds of broad-leaved trees; arthropods including bugs, flies, tortricid larvae, hymenopterans and spiders. (Not studied in Africa.) Breeding Habits. Solitary nester; apparently monogamous. Y sings from apex of conifer or sometimes side of tree, usually close to X or to nest. He may shake and flap wings. Y often sings in flight, following X in wide arc back to the perch where she started. Y threatens intruder, head lowered and thrust forward and bill open (A). Y courtshipfeeds X. Over much of range breeding begins in autumn with 2 broods reared over a protracted period and the

563

564

EMBERIZIDAE

population as a whole breeding continuously for up to 9 months. Bird fledging in spring can breed in the autumn. NEST: open cup, made of dead conifer twigs, strips of bark, moss and lichen, lined with fine dry grass, hair, wool, plant down and some feathers and fragments of wood. Built c. 45 m up in Aleppo pine by X, Y sometimes helping, taking 3–5 days. X collects material from nest tree or a tree several hundred m away, when Y accompanies her. SIZE (n ¼ 6, N Italy): ext. diam. 105–150 (127), int. diam. 58–80 (67), ext. depth 65–85 (73), int. depth 35–45 (39). EGGS: 2–5, mainly 3–4, av. (78 clutches Te ´bessa, Algeria) 334. Laid daily; hatch asynchronously over c. 3 days.

Subelliptical, smooth, slightly glossy; cream-white or bluish white, sparingly marked at broad end with purple specks and scrawls. SIZE: (78 clutches, Te´bessa) 205– 240  155–180; also (NW Africa, n ¼ 61) 200–240  146–169 (217  156). LAYING DATES: Morocco, Nov–June; Algeria, Jan–June, mainly (n ¼ 77) Feb–Apr, (nests Oct) (Le D 1935). INCUBATION: by X only, starting with first egg laid, brooding constantly for first few days, when she rarely laves the nest. Period: 13–16, mainly 14–15, days. DEVELOPMENT AND CARE OF YOUNG: growth slow; eyes start to open at 5–8 days; fed and cared for by both parents; for first 7–12 days X broods young and Y brings most of their food. Both parents swallow faecal sacs or carry them away and drop them. Nestling period: 16–28, mainly 20– 25, days. Tips of juv.’s mandibles not crossed when it fledges but become fully crossed in 3–6 weeks, during which time fledglings are still fed by parents, near nest for 10–12 days and further afield thereafter, brood being divided between the parents. BREEDING SUCCESS/SURVIVAL: of 8 pairs in N Italy, 2 failed to produce young and 6 reared 20 young successfully. Oldest ringed bird, 7 years. Key References Brosset, A. (1957), Clement, P. et al. (1993), Cramp, S. and Perrins, C.M. (1994), Le D, R. (1935).

Family EMBERIZIDAE: buntings and American sparrows

A very large family of terrestrial and arboreal granivores with numerous Nearctic and Neotropical genera and in the Old World a mere 7 genera, of which 6 are small or monotypic; radiation has occurred only in the seventh, Emberiza. American emberizids include sparrows Zonotrichia, Ammodramus, Spizella, Aimophila and others, juncos Junco, towhees Pipilo and brush-finches Atlapetes. New World warblers, tanagers, icterids and cardinals were included by Sibley and Ahlquist (1990), who also combined buntings sens. lat. with the New World and old World finches in their vast family Fringillidae. The 7 genera in the Old World are Calcarius and Plectrophenax (Holarctic), the monotypic Melophus, Latoucheornis, Urocynchramus and Miliaria, and the large Emberiza with 8 Afrotropical and 31 Palearctic species.

Genus Emberiza Linnaeus

A large but well-defined genus of streaky seedeaters, YY boldly marked and coloured, XX usually much less so; rather uniform morphologically as well as in proportions and overall size. Bill sharp, conical, sharp-pointed, culmen straight or slightly decurved, upper mandible sharply angled downward at nostrils and in most species with bony palatal protuberance. Culmen markedly decurved in some subspecies of E. schoeniclus with swollen lower mandible. Legs and feet uniform in size, even though some species are largely terrestrial (and rock-dwelling) and others largely arboreal. Mantle and scapular feathers in many species and tertials and upperwing-coverts in all species blackish with sharply-defined pale borders and tips, often rusty. Forage on ground. Songs mainly short, wheezy trills, most not melodious. Nest a deep cup; eggs with dark scribble marks. 4 plump, high-latitude buntings (longspurs, 3 Nearctic, 1 Palearctic) are very similar to Emberiza but have long hind claws and are referred to genus Calcarius.

564

EMBERIZIDAE

population as a whole breeding continuously for up to 9 months. Bird fledging in spring can breed in the autumn. NEST: open cup, made of dead conifer twigs, strips of bark, moss and lichen, lined with fine dry grass, hair, wool, plant down and some feathers and fragments of wood. Built c. 45 m up in Aleppo pine by X, Y sometimes helping, taking 3–5 days. X collects material from nest tree or a tree several hundred m away, when Y accompanies her. SIZE (n ¼ 6, N Italy): ext. diam. 105–150 (127), int. diam. 58–80 (67), ext. depth 65–85 (73), int. depth 35–45 (39). EGGS: 2–5, mainly 3–4, av. (78 clutches Te ´bessa, Algeria) 334. Laid daily; hatch asynchronously over c. 3 days.

Subelliptical, smooth, slightly glossy; cream-white or bluish white, sparingly marked at broad end with purple specks and scrawls. SIZE: (78 clutches, Te´bessa) 205– 240  155–180; also (NW Africa, n ¼ 61) 200–240  146–169 (217  156). LAYING DATES: Morocco, Nov–June; Algeria, Jan–June, mainly (n ¼ 77) Feb–Apr, (nests Oct) (Le D 1935). INCUBATION: by X only, starting with first egg laid, brooding constantly for first few days, when she rarely laves the nest. Period: 13–16, mainly 14–15, days. DEVELOPMENT AND CARE OF YOUNG: growth slow; eyes start to open at 5–8 days; fed and cared for by both parents; for first 7–12 days X broods young and Y brings most of their food. Both parents swallow faecal sacs or carry them away and drop them. Nestling period: 16–28, mainly 20– 25, days. Tips of juv.’s mandibles not crossed when it fledges but become fully crossed in 3–6 weeks, during which time fledglings are still fed by parents, near nest for 10–12 days and further afield thereafter, brood being divided between the parents. BREEDING SUCCESS/SURVIVAL: of 8 pairs in N Italy, 2 failed to produce young and 6 reared 20 young successfully. Oldest ringed bird, 7 years. Key References Brosset, A. (1957), Clement, P. et al. (1993), Cramp, S. and Perrins, C.M. (1994), Le D, R. (1935).

Family EMBERIZIDAE: buntings and American sparrows

A very large family of terrestrial and arboreal granivores with numerous Nearctic and Neotropical genera and in the Old World a mere 7 genera, of which 6 are small or monotypic; radiation has occurred only in the seventh, Emberiza. American emberizids include sparrows Zonotrichia, Ammodramus, Spizella, Aimophila and others, juncos Junco, towhees Pipilo and brush-finches Atlapetes. New World warblers, tanagers, icterids and cardinals were included by Sibley and Ahlquist (1990), who also combined buntings sens. lat. with the New World and old World finches in their vast family Fringillidae. The 7 genera in the Old World are Calcarius and Plectrophenax (Holarctic), the monotypic Melophus, Latoucheornis, Urocynchramus and Miliaria, and the large Emberiza with 8 Afrotropical and 31 Palearctic species.

Genus Emberiza Linnaeus

A large but well-defined genus of streaky seedeaters, YY boldly marked and coloured, XX usually much less so; rather uniform morphologically as well as in proportions and overall size. Bill sharp, conical, sharp-pointed, culmen straight or slightly decurved, upper mandible sharply angled downward at nostrils and in most species with bony palatal protuberance. Culmen markedly decurved in some subspecies of E. schoeniclus with swollen lower mandible. Legs and feet uniform in size, even though some species are largely terrestrial (and rock-dwelling) and others largely arboreal. Mantle and scapular feathers in many species and tertials and upperwing-coverts in all species blackish with sharply-defined pale borders and tips, often rusty. Forage on ground. Songs mainly short, wheezy trills, most not melodious. Nest a deep cup; eggs with dark scribble marks. 4 plump, high-latitude buntings (longspurs, 3 Nearctic, 1 Palearctic) are very similar to Emberiza but have long hind claws and are referred to genus Calcarius.

564

EMBERIZIDAE

population as a whole breeding continuously for up to 9 months. Bird fledging in spring can breed in the autumn. NEST: open cup, made of dead conifer twigs, strips of bark, moss and lichen, lined with fine dry grass, hair, wool, plant down and some feathers and fragments of wood. Built c. 45 m up in Aleppo pine by X, Y sometimes helping, taking 3–5 days. X collects material from nest tree or a tree several hundred m away, when Y accompanies her. SIZE (n ¼ 6, N Italy): ext. diam. 105–150 (127), int. diam. 58–80 (67), ext. depth 65–85 (73), int. depth 35–45 (39). EGGS: 2–5, mainly 3–4, av. (78 clutches Te ´bessa, Algeria) 334. Laid daily; hatch asynchronously over c. 3 days.

Subelliptical, smooth, slightly glossy; cream-white or bluish white, sparingly marked at broad end with purple specks and scrawls. SIZE: (78 clutches, Te´bessa) 205– 240  155–180; also (NW Africa, n ¼ 61) 200–240  146–169 (217  156). LAYING DATES: Morocco, Nov–June; Algeria, Jan–June, mainly (n ¼ 77) Feb–Apr, (nests Oct) (Le D 1935). INCUBATION: by X only, starting with first egg laid, brooding constantly for first few days, when she rarely laves the nest. Period: 13–16, mainly 14–15, days. DEVELOPMENT AND CARE OF YOUNG: growth slow; eyes start to open at 5–8 days; fed and cared for by both parents; for first 7–12 days X broods young and Y brings most of their food. Both parents swallow faecal sacs or carry them away and drop them. Nestling period: 16–28, mainly 20– 25, days. Tips of juv.’s mandibles not crossed when it fledges but become fully crossed in 3–6 weeks, during which time fledglings are still fed by parents, near nest for 10–12 days and further afield thereafter, brood being divided between the parents. BREEDING SUCCESS/SURVIVAL: of 8 pairs in N Italy, 2 failed to produce young and 6 reared 20 young successfully. Oldest ringed bird, 7 years. Key References Brosset, A. (1957), Clement, P. et al. (1993), Cramp, S. and Perrins, C.M. (1994), Le D, R. (1935).

Family EMBERIZIDAE: buntings and American sparrows

A very large family of terrestrial and arboreal granivores with numerous Nearctic and Neotropical genera and in the Old World a mere 7 genera, of which 6 are small or monotypic; radiation has occurred only in the seventh, Emberiza. American emberizids include sparrows Zonotrichia, Ammodramus, Spizella, Aimophila and others, juncos Junco, towhees Pipilo and brush-finches Atlapetes. New World warblers, tanagers, icterids and cardinals were included by Sibley and Ahlquist (1990), who also combined buntings sens. lat. with the New World and old World finches in their vast family Fringillidae. The 7 genera in the Old World are Calcarius and Plectrophenax (Holarctic), the monotypic Melophus, Latoucheornis, Urocynchramus and Miliaria, and the large Emberiza with 8 Afrotropical and 31 Palearctic species.

Genus Emberiza Linnaeus

A large but well-defined genus of streaky seedeaters, YY boldly marked and coloured, XX usually much less so; rather uniform morphologically as well as in proportions and overall size. Bill sharp, conical, sharp-pointed, culmen straight or slightly decurved, upper mandible sharply angled downward at nostrils and in most species with bony palatal protuberance. Culmen markedly decurved in some subspecies of E. schoeniclus with swollen lower mandible. Legs and feet uniform in size, even though some species are largely terrestrial (and rock-dwelling) and others largely arboreal. Mantle and scapular feathers in many species and tertials and upperwing-coverts in all species blackish with sharply-defined pale borders and tips, often rusty. Forage on ground. Songs mainly short, wheezy trills, most not melodious. Nest a deep cup; eggs with dark scribble marks. 4 plump, high-latitude buntings (longspurs, 3 Nearctic, 1 Palearctic) are very similar to Emberiza but have long hind claws and are referred to genus Calcarius.

Emberiza cabanisi

565

At least 39 species, in Palearctic, Africa and S Asia; at least 28 of them fall into groups of closely related species, several embracing superspecies. 21 species in Africa, 14 breeding, 8 endemic. Species-group (1), 7 rock buntings: 4 endemic, capensis, vincenti and impetuani (southern Africa) and socotrana (Socotra); tapahisi widespread in Africa, also in SW Arabia; striolata circumSaharan and east to NW India; and cia, somewhat divergent from the other 5, in NW Africa, S Europe and Asia. Speciesgroup (2), 4 endemic yellow-breasted buntings, flaviventris (widespread), affinis (N tropics) and poliopleura (E Africa) parapatric and little more differentiated than a superspecies, and cabanisi (widespread). The remaining 10 species are Palearctic ones whose breeding ranges extend into NW Africa (cirlus, hortulana, schoeniclus) or which are non-breeding winter visitors (cineracea, caesia) or vagrants (citrinella, leucocephalos, rustica, pusilla, melanocephala). We recognize 3 superspecies: flaviventris/poliopleura, capensis/vincenti and leucocephalos/citrinella.

Emberiza capensis superspecies

Emberiza flaviventris superspecies

1

1

2 1,2 1 E. flaviventris 2 E. poliopleura

1

1 E. capensis 2 E. vincenti

2

1

Emberiza cabanisi (Reichenow). Cabanis’s Bunting. Bruant de Cabanis.

Plate 34

Polymitra (Fringillaria) cabanisi Reichenow, 1875. J. Orn., 23, p. 233, pl. 2, figs 2,3; Cameroons.

(Opp. p. 523)

Range and Status. Endemic resident, tropics between c. 10
N and 20
S. Guinea, Wassou and Mambia areas (Demey 1995), frequent in Haut Niger Nat. Park (Nikolaus 2000), 2 records in E (1 Gekedou), near Liberian border. Sierra Leone, uncommon in N and NE (Nerekoro, Momoria Bedala, Loma Mansa, Tembikunda Mts). Liberia, bird received dead, Voinjama, Jan 1993, is the only record. Mali, known only from Bamako (Y collected 1962), east of Mandjakuy (several singing YY) and Sikasso region. Burkina Faso, recorded (Y. Thonne´rieux in Dowsett 1993). Ivory Coast, scarce in S guinean savannas but quite common north of 9
N, at least in E of country; uncommon to frequent in Comoe´ Nat. Park. Ghana, not uncommon resident, widespread; in Mole, numbers increase in dry season. Togo, not uncommon in guinea savanna zone, from Bismarckburg and Misaho ¨he north to Paio; common along

Bafilo ridge. Benin, rare, Be´te´rou region. Nigeria, common in Kainji Lake Nat. Park and around Enugu, uncommon in Falgore Game Res. and Gashaka-Gumti Nat. Park, elsewhere irregular, local but widespread, south to Ibadan and Port Harcourt. Cameroon, uncommon (55 localities, M. Languy, pers. comm.): Benue´ Plain, Parc Nat. de la Benoue´ Nord, Adamawa Plateau, and in S along forestsavanna interface; west to Essosong, Mt Kupe´. Mbini, record(s) in N. Chad, frequent between Le´re´ and Pala (Vieillard 1972). Central African Republic, occurs in at least 4 regions or localities in N, one in W and several in S including Ouossi R., as shown, but probably far more widespread; known from Ngoundji, Mboko, Landjia and Bo, all close together around 4
300 N, 18
300 E (Germain and Cornet 1994). Sudan, uncommon in S. NE Zaı¨re, frequent. Uganda, uncommon resident and wanderer in

Emberiza cabanisi

565

At least 39 species, in Palearctic, Africa and S Asia; at least 28 of them fall into groups of closely related species, several embracing superspecies. 21 species in Africa, 14 breeding, 8 endemic. Species-group (1), 7 rock buntings: 4 endemic, capensis, vincenti and impetuani (southern Africa) and socotrana (Socotra); tapahisi widespread in Africa, also in SW Arabia; striolata circumSaharan and east to NW India; and cia, somewhat divergent from the other 5, in NW Africa, S Europe and Asia. Speciesgroup (2), 4 endemic yellow-breasted buntings, flaviventris (widespread), affinis (N tropics) and poliopleura (E Africa) parapatric and little more differentiated than a superspecies, and cabanisi (widespread). The remaining 10 species are Palearctic ones whose breeding ranges extend into NW Africa (cirlus, hortulana, schoeniclus) or which are non-breeding winter visitors (cineracea, caesia) or vagrants (citrinella, leucocephalos, rustica, pusilla, melanocephala). We recognize 3 superspecies: flaviventris/poliopleura, capensis/vincenti and leucocephalos/citrinella.

Emberiza capensis superspecies

Emberiza flaviventris superspecies

1

1

2 1,2 1 E. flaviventris 2 E. poliopleura

1

1 E. capensis 2 E. vincenti

2

1

Emberiza cabanisi (Reichenow). Cabanis’s Bunting. Bruant de Cabanis.

Plate 34

Polymitra (Fringillaria) cabanisi Reichenow, 1875. J. Orn., 23, p. 233, pl. 2, figs 2,3; Cameroons.

(Opp. p. 523)

Range and Status. Endemic resident, tropics between c. 10
N and 20
S. Guinea, Wassou and Mambia areas (Demey 1995), frequent in Haut Niger Nat. Park (Nikolaus 2000), 2 records in E (1 Gekedou), near Liberian border. Sierra Leone, uncommon in N and NE (Nerekoro, Momoria Bedala, Loma Mansa, Tembikunda Mts). Liberia, bird received dead, Voinjama, Jan 1993, is the only record. Mali, known only from Bamako (Y collected 1962), east of Mandjakuy (several singing YY) and Sikasso region. Burkina Faso, recorded (Y. Thonne´rieux in Dowsett 1993). Ivory Coast, scarce in S guinean savannas but quite common north of 9
N, at least in E of country; uncommon to frequent in Comoe´ Nat. Park. Ghana, not uncommon resident, widespread; in Mole, numbers increase in dry season. Togo, not uncommon in guinea savanna zone, from Bismarckburg and Misaho ¨he north to Paio; common along

Bafilo ridge. Benin, rare, Be´te´rou region. Nigeria, common in Kainji Lake Nat. Park and around Enugu, uncommon in Falgore Game Res. and Gashaka-Gumti Nat. Park, elsewhere irregular, local but widespread, south to Ibadan and Port Harcourt. Cameroon, uncommon (55 localities, M. Languy, pers. comm.): Benue´ Plain, Parc Nat. de la Benoue´ Nord, Adamawa Plateau, and in S along forestsavanna interface; west to Essosong, Mt Kupe´. Mbini, record(s) in N. Chad, frequent between Le´re´ and Pala (Vieillard 1972). Central African Republic, occurs in at least 4 regions or localities in N, one in W and several in S including Ouossi R., as shown, but probably far more widespread; known from Ngoundji, Mboko, Landjia and Bo, all close together around 4
300 N, 18
300 E (Germain and Cornet 1994). Sudan, uncommon in S. NE Zaı¨re, frequent. Uganda, uncommon resident and wanderer in

566

EMBERIZIDAE

Emberiza cabanisi

? ?

?

?

?

NW, south and east to Murchison Falls Nat. Park, Budongo, Masindi and Acholi. Burundi, scarce, Akagera Nat. Park (Kiyonza, Kitabiri). Gabon, Mt Iboundji. Congo, widespread from Le´fini Res. northwest to Gabon border; in W Zaı¨re lives along Congo R. so doubtless on Congo side as well. In other countries south of Equator as mapped, sparse to locally frequent, probably far more widespread than shown in W and S Zaı¨re, Angola, W Tanzania and Mozambique north of 20
N and south to Cometala. At 300–2000 m but mainly ˆ i, regular on Malaw ˆ i Hill above at 750–1800 m; in Malaw 490 m, elsewhere at 760–1530 m, sparse on Viphya Mts up to 1830 m and on Nyika Plateau up to 2070 m. In S Zimbabwe extends to Bikita, Great Zimbabwe, Sarahuru, Mt Buhwa and Matobo Hills (where rare); in E Highlands, occurs up to 1300 m on Iyanga, in Honde Valley, Mt Gorongo, and at Stapleford at 1400 m, south to Lusitu– Haroni confluence in Chipinga Uplands (Irwin 1981). Description. E. c. cabanisi Reichenow: Sierra Leone to S Sudan, NE Zaı¨re and NW Uganda. ADULT Y: forehead to nape blackish brown, merging with dull grey-brown upperparts; mantle and scapulars with diffuse blackish streaking. Tail feathers blackish brown, T3–T5 with progressively larger white distal area on inner web and small white tip to outer web, T6 with distal half of inner web white and outer web white with black streak along shaft broadening toward tip (A). Narrow white superciliary stripe, extending to below nape; lores, cheeks and ear-coverts blackish brown, well demarcated from ashy grey sides of neck (A). Chin and throat white; sides of breast dark grey; rest of breast, flanks and belly bright yellow; thighs dark grey; undertail-coverts dull white. Remiges dark grey-brown, primaries and secondaries narrowly edged pale grey-buff; primary coverts and alula dark grey brown; greater coverts dark grey-brown, outer feathers tipped white to form narrow wing-bar; median coverts blackish, tipped white to form a broader upper wing-bar; lesser coverts dull grey-brown. Underwing-coverts dark grey-brown with pale grey

fringes; axillaries grey. Bill dark horn or blackish above and at tip below, base of lower mandible pale pinkish or pale horn; eyes brown; legs dull pale brown. ADULT X: similar to ad. Y, but top of head, cheeks and ear-coverts browner, upperparts slightly browner, superciliary stripe and throat tinged buff, upper breast washed tawny-brown. SIZE (10 YY, 10 XX): wing, Y 77–84 (810), X 75–82 (768); tail, Y 66–78 (725), X 65–68 (666); bill, Y 145–16 (150), X 14–15 (145); tarsus, Y 19–21 (197), X 19–21 (198). WEIGHT: Ghana, 1 Y 240, 2 XX 241, 255, unsexed (n ¼ 2) 22, 225. IMMATURE: juv. differs from ad. X in having top of head and upperparts browner, breast browner (less yellow), wing-bars pale buff or pale brown. E. c. orientalis Shelley (includes ‘cognominata’): S Zaı¨re and Tanzania to Angola, Zambia, Zimbabwe and N Mozambique. Y differs from nominate race in having dark-streaked, pale grey central crown stripe; sides of crown and cheek patch blacker; hindneck and sides of neck paler; mantle and scapulars paler grey-brown with more prominent blackish streaking and some rufous feather edging; yellow on underparts extending to throat; and median covert bar broader. X richer brown and more streaked on head and upperparts than in nominate race, with yellow on throat. WEIGHT: Zimbabwe, Y (n ¼ 1) 223, unsexed (n ¼ 7) 137–255 (213). NESTLING: dark brown feathers of upperparts margined with rufous–buff; chin and throat rufous–buff, belly yellow (Bannerman 1948).

Field Characters. Length 16–17.5 cm. A stripe-headed, yellow-breasted bunting of central and W Africa, distinguished from the 3 similar buntings by grey back, solid black face patch without white line below eye, and lack of orange wash on breast. X like Y but head duller, tawny wash on upper breast; juv. like X but head, upperparts and supercilium tawny, face patch brown, underparts less yellow. Voice. Tape-recorded (86, 99, 104, B, C, F, LEM). Song unhurried, sweet and ringing, extremely variable, often with repetition of single or double note: ringing ‘tyowytyowy-tyowy-tyowy-tyowy’; up-slurred ‘chuwee-chuweechuwee’; down-slurred ‘cheewi-cheewi-cheewi’; with introductory note, short, ‘ts wee-wee-wee-wee’, ‘sit-sweeswee-swee’, or long, ‘wee-tya-tya-tya-tya’; 2-speed, ‘tututututu-tsway-tsway-tsway’; 3-part, ‘ps-ps-chewy-chewy-chu’; and a variety of slow trills, 9–11 notes on one pitch. Many songs reminiscent of Coal Tit Parus ater. Call, a rolled, upslurred ‘burrreee’; contact note a clear ‘tsseeoo’; in flight, ‘tsipp’; warning call a high-pitched, drawn-out, sibilant ‘seeeee’ (Byers et al. 1995).

General Habits. Inhabits wooded grassland, guinea savanna with scattered trees and rocky terrain (Togo), farm clearings in orchard bush, farmland and degraded

Emberiza flaviventris savanna (Nigeria), borders of heavy timber where shrubs grow profusely, and second-growth bush dotted with small trees (S Zaı¨re, Vincent 1949); Brachystegia woodland and ˆ i). montane grassland with scattered bushes (Malaw Inhabits Brachystegia with little or no undergrowth in Zimbabwe and may be at its commonest in undisturbed miombo, but in E also occurs in places with scattered trees and bushes, often near borders of lower-altitude evergreen forest. In pairs, sometimes solitary or in family parties; small flocks have occurred near Harare, Zimbabwe. Forages exclusively on ground, sometimes in company of other buntings and finches, although appears to be largely insectivorous. Occasionally a member of mixed-species foraging flocks in miombo woods (Zimbabwe). Waits in a clear space on ground for advancing column of driver ants, and eats small insects that they disturb (R.E. Moreau in Chapin 1954). Uses bushes and small trees freely, flying to one when disturbed on ground, singing from small tree top, and nesting in them. Appears to moult completely after breeding. In Nigeria birds show signs of wear from Mar and wear becomes progressively more marked until Oct; plumage moulted regularly in Nov–Dec, but chin and throat feathers moult irregularly (Serle 1957). In southern Africa moults in Mar–May. Reporting rates in Zimbabwe markedly greater in Nov and somewhat greater in Aug–Feb than in Mar–July; due either to greater conspicuousness of singing YY in southern spring or to partial migration (Harrison et al. 1997). In Birkdale Estate, Mvurwi, Mashonaland highveld, Zimbabwe, present erratically in Aug–Jan in normal years but in drought year of 1992 frequent from Apr onward (Tree 1994) Food. Insects and seeds, including millet and rice (Byers et al. 1995). Of 27 stomachs (Enugu, Nigeria, all months, Serle 1957), insects in 24 and seeds in 3; of 6 in NE Zaı¨re, all contained insects (grasshoppers, beetles) and none held seeds; eats mainly small grasshoppers in S Cameroon. In captivity eats seeds of cereal grasses Rhynchelytrum repens, Panicum natalense and P. maximum (Brickell 1986).

567

Breeding Habits. Solitary nester; monogamous; territorial. NEST: one in Nigeria composed mainly of fine roots and placed in a yam-vine. Built by X, Y perching near nest site and singing; he occasionally brings material. 3 in S Cameroon: shallow, loosely built of dried leaves and small stems, with a few fine fibres inside cup. In S Zaı¨re, a rather bulky cup of old, chaffed grass, fine twiglets and dry, weedy plant stems and a few brown and skeletal leaves, quite neatly lined with soft, fine grass and some small rootlets. Sited in shrub, small tree or sapling, in fork 1–2 m (once 45 m) above ground, well hidden within screen of leaves. Built by X only; or, mainly by X, the Y accompanying her on collecting trips and occasionally bringing material himself (Marchant 1942). EGGS: 2–3. Rather wide, very slightly glossy; dull white or very pale cream, with large, irregular, fine, twirling lines, short wavy pencillings, hair lines and blotches of pale umber-brown and pale grey; markings concentrated in a ring near middle of egg, towards its broad end. A different egg type is finely freckled all over with pale sepia and pale grey, with dull, pale grey clouding. SIZE: (W Africa) 220  155, 210  55; (southern Africa, n ¼ 17) 180–219  135–151 (195  145). LAYING DATES: Ghana, (carrying food, early Sept); Togo, (carrying food, Aug); Nigeria, Sept (and nest being built, July; birds are in fresh plumage in (Dec)–Jan–Feb); Sudan, July; Zaı¨re, Katanga, Dec (thought to breed in Feb–May in ˆ i, Oct–Jan, mainly Nov– NE); Angola, (Sept–Jan); Malaw Dec; Zambia, Oct–Feb; Zimbabwe, Sept–Mar, mainly (67% of 45 clutches) Oct–Nov. INCUBATION: period 13–14 days. By X only, Y feeding her on the nest. DEVELOPMENT AND CARE OF YOUNG: nestling period once 16 days (in captivity). BREEDING SUCCESS/SURVIVAL: parasitized by Klaas’s Cuckoo Chrysococcyx klaas (records in Tanzania and Mozambique). Bird ringed as ad., Zimbabwe, found dead after 37 months. Key References Byers, C. et al. (1995), Chapin, J.P. (1954), Skead, C.J. (1960), Vincent, A.W. (1946).

Emberiza flaviventris Stephens. Golden-breasted Bunting. Bruant a ` poitrine dore´e.

Plate 34

Emberiza flaviventris Stephens, 1815. In Shaw, General Zool., 9, 2, p. 374; Cape of Good Hope and Cayenne.

(Opp. p. 523)

Forms a superspecies with E. poliopleura. Range and Status. Endemic resident and wanderer. Senegal, 1, Sinnthiou-Fissa. Mauritania, locally quite common, from Senegal R. valley north to 17
N or in Tagant region to 17
300 N, e.g. Plateau d’El Aaguer. Mali, locally quite common as mapped (Lamarche 1993), sometimes abundant, between 12
300 N and 17
N; not in W (Boucle du Baoule´ Nat. Park). Burkina Faso, uncommon around Ouagadougou; frequent in NW; frequent in central latitudes, southwest to about 11
400 N, 3
050 W; rare in

Pendjari and Arli Nat. Parks (where may just enter NW Benin). Benin, rare, Be´te´rou (Apr). Niger, ‘W’ Nat. Park; a few records in W, north to about Talcho and Tahoua (both 15
N) and east to E Maradi District, and 2 in E, at Kaadjia (13
540 N) and near Nguigmi. Nigeria, fairly common in N Sokoto State; in N Kano State, occasional around Kano (12
N), but further north frequent at Yashi and common in Daura area on Niger border; record Aliya (11
100 N, 10
550 E), occasional in NE Bornu State, south to Maiduguri

Emberiza flaviventris savanna (Nigeria), borders of heavy timber where shrubs grow profusely, and second-growth bush dotted with small trees (S Zaı¨re, Vincent 1949); Brachystegia woodland and ˆ i). montane grassland with scattered bushes (Malaw Inhabits Brachystegia with little or no undergrowth in Zimbabwe and may be at its commonest in undisturbed miombo, but in E also occurs in places with scattered trees and bushes, often near borders of lower-altitude evergreen forest. In pairs, sometimes solitary or in family parties; small flocks have occurred near Harare, Zimbabwe. Forages exclusively on ground, sometimes in company of other buntings and finches, although appears to be largely insectivorous. Occasionally a member of mixed-species foraging flocks in miombo woods (Zimbabwe). Waits in a clear space on ground for advancing column of driver ants, and eats small insects that they disturb (R.E. Moreau in Chapin 1954). Uses bushes and small trees freely, flying to one when disturbed on ground, singing from small tree top, and nesting in them. Appears to moult completely after breeding. In Nigeria birds show signs of wear from Mar and wear becomes progressively more marked until Oct; plumage moulted regularly in Nov–Dec, but chin and throat feathers moult irregularly (Serle 1957). In southern Africa moults in Mar–May. Reporting rates in Zimbabwe markedly greater in Nov and somewhat greater in Aug–Feb than in Mar–July; due either to greater conspicuousness of singing YY in southern spring or to partial migration (Harrison et al. 1997). In Birkdale Estate, Mvurwi, Mashonaland highveld, Zimbabwe, present erratically in Aug–Jan in normal years but in drought year of 1992 frequent from Apr onward (Tree 1994) Food. Insects and seeds, including millet and rice (Byers et al. 1995). Of 27 stomachs (Enugu, Nigeria, all months, Serle 1957), insects in 24 and seeds in 3; of 6 in NE Zaı¨re, all contained insects (grasshoppers, beetles) and none held seeds; eats mainly small grasshoppers in S Cameroon. In captivity eats seeds of cereal grasses Rhynchelytrum repens, Panicum natalense and P. maximum (Brickell 1986).

567

Breeding Habits. Solitary nester; monogamous; territorial. NEST: one in Nigeria composed mainly of fine roots and placed in a yam-vine. Built by X, Y perching near nest site and singing; he occasionally brings material. 3 in S Cameroon: shallow, loosely built of dried leaves and small stems, with a few fine fibres inside cup. In S Zaı¨re, a rather bulky cup of old, chaffed grass, fine twiglets and dry, weedy plant stems and a few brown and skeletal leaves, quite neatly lined with soft, fine grass and some small rootlets. Sited in shrub, small tree or sapling, in fork 1–2 m (once 45 m) above ground, well hidden within screen of leaves. Built by X only; or, mainly by X, the Y accompanying her on collecting trips and occasionally bringing material himself (Marchant 1942). EGGS: 2–3. Rather wide, very slightly glossy; dull white or very pale cream, with large, irregular, fine, twirling lines, short wavy pencillings, hair lines and blotches of pale umber-brown and pale grey; markings concentrated in a ring near middle of egg, towards its broad end. A different egg type is finely freckled all over with pale sepia and pale grey, with dull, pale grey clouding. SIZE: (W Africa) 220  155, 210  55; (southern Africa, n ¼ 17) 180–219  135–151 (195  145). LAYING DATES: Ghana, (carrying food, early Sept); Togo, (carrying food, Aug); Nigeria, Sept (and nest being built, July; birds are in fresh plumage in (Dec)–Jan–Feb); Sudan, July; Zaı¨re, Katanga, Dec (thought to breed in Feb–May in ˆ i, Oct–Jan, mainly Nov– NE); Angola, (Sept–Jan); Malaw Dec; Zambia, Oct–Feb; Zimbabwe, Sept–Mar, mainly (67% of 45 clutches) Oct–Nov. INCUBATION: period 13–14 days. By X only, Y feeding her on the nest. DEVELOPMENT AND CARE OF YOUNG: nestling period once 16 days (in captivity). BREEDING SUCCESS/SURVIVAL: parasitized by Klaas’s Cuckoo Chrysococcyx klaas (records in Tanzania and Mozambique). Bird ringed as ad., Zimbabwe, found dead after 37 months. Key References Byers, C. et al. (1995), Chapin, J.P. (1954), Skead, C.J. (1960), Vincent, A.W. (1946).

Emberiza flaviventris Stephens. Golden-breasted Bunting. Bruant a ` poitrine dore´e.

Plate 34

Emberiza flaviventris Stephens, 1815. In Shaw, General Zool., 9, 2, p. 374; Cape of Good Hope and Cayenne.

(Opp. p. 523)

Forms a superspecies with E. poliopleura. Range and Status. Endemic resident and wanderer. Senegal, 1, Sinnthiou-Fissa. Mauritania, locally quite common, from Senegal R. valley north to 17
N or in Tagant region to 17
300 N, e.g. Plateau d’El Aaguer. Mali, locally quite common as mapped (Lamarche 1993), sometimes abundant, between 12
300 N and 17
N; not in W (Boucle du Baoule´ Nat. Park). Burkina Faso, uncommon around Ouagadougou; frequent in NW; frequent in central latitudes, southwest to about 11
400 N, 3
050 W; rare in

Pendjari and Arli Nat. Parks (where may just enter NW Benin). Benin, rare, Be´te´rou (Apr). Niger, ‘W’ Nat. Park; a few records in W, north to about Talcho and Tahoua (both 15
N) and east to E Maradi District, and 2 in E, at Kaadjia (13
540 N) and near Nguigmi. Nigeria, fairly common in N Sokoto State; in N Kano State, occasional around Kano (12
N), but further north frequent at Yashi and common in Daura area on Niger border; record Aliya (11
100 N, 10
550 E), occasional in NE Bornu State, south to Maiduguri

568

EMBERIZIDAE

Emberiza flaviventris

?

?

and Dikwa. Cameroon, 4 records: Gagadje´, Garoua, Matoua and 40 km south of Maroua (M. Languy, pers. comm.). Chad, common in sahel zone and frequent in sudan zone, south to 11
N; abundant at Abe´che´ in July; in Ouadi Rime´–Ouadi Achim Faunal Res., restricted to SE quarter where uncommon. Central African Republic, ManovoGounda-Saint Floris Nat. Park and Bamingui area (Carroll 1988) and Vakaga region. Sudan, frequent throughout central belt, common at about 14
N and numerous near El Obeid, and common in valleys of Atbara R., Setit R., and Blue Nile above Wad Medani (flavigaster), uncommon in extreme S (kalaharica). Ethiopia, presumably this species, not E. poliopleura, seen uncommonly in Tekeze (Takkaze) R. valley and around Adwa (Aduwa), Tigray (Cheesman and Sclater 1935–1936). Eritrea, frequent at 1650–1750 m (Zinner 2001; occurs lower, but absent from coastal plains. Uganda and to south, minimal range is as shown. Fairly common in E Africa, up to 2200 m. Widespread in Rwanda, at least in W. Burundi, resident in Bururi Forest. Much commoner than Cabanis’s Bunting E. cabanisi around Lumbumbashi S Zaı¨re. Probably more widespread than shown, in Tanzania and N Mozambique. Angola, common and widespread, east to Zaı¨re and Zambia borders. Common almost throughout Zambia, Zimbabwe, NE South Africa and S Mozambique. Frequent to common in Botswana but absent from SW; locally very common at some seasons in E; Manamodi Pans, July. Av. density in Botswana of 1 bird per 2 ha in 8 types of broad–leaved and mixed open woodland, and max. of 1 bird per 08 ha; in Acacia-dominated habitats, 1 bird per 20 ha (M. Herremans in Harrison et al. 1997). Probably >500,000 birds in S Mozambique, where densities in acacia savanna of 3, 10, 15 and 225 birds per 100 ha (Monadjem 2002).

Description. E.f. kalaharica Roberts (includes ‘carychroa’): SE Sudan, Uganda, Kenya, E and S Zaire and Tanzania south to central and SE Angola, NE Namibia, Botswana, Mozambique (north of Delagoa Bay) and N South Africa (Transvaal, Free State and N Cape). ADULT Y: stripe from centre of forehead to centre of nape white flecked with black, bordered by broad black crown stripe; hindneck grey-brown; mantle and scapulars rufouschestnut, feathers fringed grey-buff when fresh; back, rump and short uppertail-coverts ashy grey; long uppertail-coverts greybrown. Tail feathers blackish brown, T1 narrowly fringed buffish, T3 narrowly tipped white, T4–T5 with distal part of both webs extensively white, T6 with distal white area extending more than half the feather length, but sullied brownish on outer web and with broad black shaft streak near tip (A). Lores white, linked to broad white superciliary stripe which extends back to nape, and to broad white subocular stripe; black mark in front of eye, narrowly black below eye and broad black stripe behind eye, linked though black bar around ear-coverts to black lower cheeks and moustachial stripe (A). Upper chin to malar region and front sides of neck greyish white, grading to ashy grey on rear sides of neck and sides of upper breast; lower chin to centre of belly golden yellow with broad orange-tawny band across upper breast, well demarcated from yellow throat; upper flanks cinnamonbrown, lower flanks grey-brown; sides of belly, vent, thighs and undertail-coverts greyish white. Remiges blackish brown, primaries narrowly edged buff, secondaries broadly edged buff, tertials broadly fringed cinnamon-buff on outer web, the innermost with entire outer web rufous-cinnamon; primary coverts and alula blackish brown; greater coverts blackish brown, edged buff, off-white tips forming prominent lower wing-bar, innermost feathers fringed cinnamon-buff; median coverts and outer lower lesser coverts white with hidden blackish bases, forming broader upper wing-bar; rest of lesser coverts blackish, tipped grey. Axillaries and underwing-coverts whitish, but primary underwing-coverts grey-brown. Bill dark horn above, pinkish below; eyes dark brown; legs pinkish brown. ADULT X: similar to ad. male, but differs in having black parts of head pattern tinged brown and central crown stripe pale buff; sides of neck, hindneck and back to uppertail-coverts more brownish grey; yellow underparts slightly paler, saffron band less intense; lesser coverts tipped brownish grey. SIZE (10 YY, 10 XX): wing, Y 79–86 (830), X 78–83 (806); tail, Y 66–75 (713), X 65-73 (681); bill, Y 13–15 (139), X 13–15 (139); tarsus, Y 175–195 (188), X 175–195 (186). WEIGHT: Uganda and Kenya, unsexed (n ¼ 5) 175–212 (191); Zambia, Y (n ¼ 2) 195, 22, X (n ¼ 2) 205, 228; Zimbabwe, Y (n ¼ 5) 151–205 (185), X (n ¼ 5) 161–185 (176).

IMMATURE: juv. differs from ad. X in having mantle pale tawny brown, streaked dark brown, and scapulars cinnamon-rufous, streaked blackish; back to uppertail-coverts browner; breast band pale tawny-brown, with fine dusky shaft-streaks; yellow on underparts paler, more restricted; pale bar across tips of greater coverts narrower; white bar across median coverts marked with black; tail feathers browner, white distal areas less extensive NESTLING: skin brownish; dense dorsal down, grey-brown; palate yellow-orange; edges of mouth and gape pale yellow (photo

Emberiza flaviventris in Ginn et al. 1989; mouth very different from that of nestling E. capensis, q. v.). E.f. flavigaster Cretzschmar: S edge of Sahara from Mauritania to Eritrea. Paler than kalaharica, mantle and scapulars more rufous-cinnamon (less chestnut), hindneck more buffy, sides of breast tinged more cinnamon (less grey); strong yellow on underparts more restricted to central areas. Smaller; wing, Y (n ¼ 10), 76–82 (792). Weight: Chad, unsexed (n ¼ 5) 14–19 (165). E.f. flaviventris Stephens: South Africa (S and E Cape Prov. to KwaZulu-Natal), E Swaziland and S Mozambique (Maputo District). Slightly darker and browner than kalaharica; central crown stripe tinged rufous-buff, rump to uppertail-coverts tinged olive-buff, hindneck and sides of breast browner. E.f. princeps Clancey and Winterbottom: central, N and NW Namibia to SW Angola. Slightly paler above than kalaharica, rump lighter grey; breast band paler and more diffuse; flanks and sides of belly rather whiter. Larger: wing, Y (n ¼ 7) 86–93 (900), X (n ¼ 4) 83–86 (841) (Clancey and Winterbottom 1960).

Field Characters. Length 15–16 cm. A widespread and attractive bunting with black-and-white striped head, yellow underparts and orange-tawny breast. In flight shows 2 white wing-bars and broad white corners to tail. Differs from Cabanis’s Bunting E. cabanisi in plain rufous mantle and scapulars, white stripe on face below eye and orange breast, and from Brown-rumped Bunting E. affinis in grey rump, broad white wing-bars, broader black stripes on head, white belly and more contrasting orange breastband. In nominate race, black eyeline extends forward across part of lores (lores wholly white in Brown-rumped and Somali Buntings E. poliopleura); for further differences from Somali Bunting, see that species. Juv. browner and duller, with brown and buff head stripes, streaked back, less white in wing. Voice. Tape-recorded (22, 58, 72, 86, 88, 99, 104, B, C, F, LEM, PAY). Variable song pleasant but not melodious, with sibilant quality; typically repetition of single or double notes in short phrase, ‘tswee-tswee-tswee’; ‘chuvee-chuveechu’; tit-like ‘ts-cha, ts-cha, ts-cha, ts-cha’ or ‘p’tseea, p’tseea, p’tseea’; 2-part, ‘tsiu-hee-hee-hee-hee’ or ‘way-wayway-tsyap-tsyap-tsyap-tsyap’; 4-part, with variable speed, ‘tya-tya-tya-tya-tya, woy, tititititi-tyaa-tyaa’; 2-speed, second part faster, ‘tswee-tswee-tsutsutsutsutsu’; sometimes a slow dry trill, ‘trutrutrutrutru . . . ’. Similar but weaker songs given outside breeding season (Skead 1960). Common call, a down-slurred, slightly rolled ‘treeeoo’, rather harder in W Africa, a double ‘plo-jeeea’; often answered by mate with casual ‘sitee’. In flight a single, quiet ‘chip’; contact call between Y and X, quiet ‘tsip, tsip, tsip’; alarm, ‘choochoochee’ and a low grating call (Skead 1960). General Habits. Inhabits combretaceous and sparse or dense Acacia country from Mauritania to Eritrea, and Acacia and other dry woodlands in Angola; around Sokoto, NW Nigeria, thin, open bushland, dry and thorny but with some broad-leaved trees, bordering a cultivated plain. In remainder of range habitat is different: open broad-leaved and mixed woodland and treed and bushy grassland, woody plantations, farmyards and gardens. In E Africa, open woodland, forest edges and suburban gardens between 1400 and 2200 m; further south, does occur in

Acacia but not commonly (though common in low thornveld around Umvuma, Zimbabwe); shuns tall, dense riverine woods (e.g. floodplain-fringing forest in Okavango, and scarce in well-wooded parts of Limpopo and Zambezi Valleys); also near Umvuma in open second-growth bush on high ground, and around Bulawayo sparse in alluvial woodland and richer miombo formations; absent from seaward-facing slopes of E highlands. In Botswana, commonest in open Combretum and Terminalia woodland. Keeps mainly to the ground under canopy of trees and avoids open areas without shade. In southern Africa, from sea level up to 1400 m (Zimbabwe). Usually in pairs; sometimes in flocks of 10–30, and flocks of several hundreds can occur near waterholes in the dry season. Forms mixed foraging flocks with other buntings and canaries. Tame and confiding around dwellings. Occurs alongside Cabanis’s Bunting, without obvious ecological difference (but bill sizes differ, so dietary seeds probably differ accordingly: B). Feeds entirely on ground; gait a one-foot-forward hop, bird crouching and shuffling with short, quick restless steps, stopping now and then to pick up an item of food. When approached, moves quickly, hopping out of the way; flies rather reluctantly, but when pressed, to nearest bush or tree; keeps out of view, then soon resumes foraging on ground. Flight rather more undulating than in other buntings; sometimes deeply undulating, with wings closing between each series of beats; often very like dipping flight of a honeyguide Indicator. Often flicks tail up upon settling. Sings from top of shady side of tree. Drinks regularly and generally occurs near water, but can also occur in arid woodlands well away from water. Reporting rates throughout southern Africa show a substantial seasonal density variations in N Botswana (increase in Nov–Mar) and SE Botswana, S Zimbabwe and Transvaal (decrease in Dec–May), consistent with migration on a WNW–ESE axis (Harrison et al. 1997). A regular migrant at a farm near Mvurwi, Zimbabwe, arriving in Aug and departing in Dec–Jan, and altitudinal movements detected on Inyanga Downs, Zimbabwe (Tree 1994). Food. Seeds, including grass Chloris virgata; flower buds; insects, including ants and termites (Skead 1960); 95% of meals given to young are insects (small noctuid and saturniid larvae, crickets, mantids, termites, flies and nymphal grasshoppers: van Someren 1956). In captivity, eats seeds of Eragrostis tef, E. curvula, E. obtusa, Phalaris canariensis, Panicum maximum and P. laevifolium and fruits of Cestrum laevigatum (Brickell 1986). Breeding Habits. Solitary nester; monogamous, perhaps pairing for life; no elaborate courtship displays; territorial, with some mild rivalry but no serious conflict when a stray Y appears, and 4 adjacent breeding pairs did not squabble over territory (van Someren 1956). 2 YY singing nearby on small trees raised crown feathers, quivered wings, expanded tails, and often fluffed out orange breast feathers; 1 Y then chased the other away, flew to ground next to a X, picked up a grass stem and passed it rapidly across bill several times, with crest raised. He dropped the grass,

569

570

EMBERIZIDAE

around broad end (C) or, occasionally, around middle of egg; black lines can be thin or wide, or both on 1 shell; often 1–2 bold, jet-black blobs or broad, curly lines (Vincent 1949). SIZE: (n ¼ 206, southern Africa) 155– 225  131–153 (203  143); (n ¼ 4, Nigeria) 182–190  129–136 (185  132). LAYING DATES: Mauritania and Mali, June–Sept; Burkina Faso, (carrying nest material, July); Nigeria, June (nestlings, May, nest-building, June); Sudan, July–Sept (flavigaster); Eritrea, (young, July); E Africa: Region A, Feb, May, July, Sept, Region C, Oct–Apr, Region D, Dec– May, July, mainly (13 out of 25 clutches) Apr; Angola, Sept; Botswana, Oct–Jan; E Cape, Sept–Mar (mainly Nov– Dec); Zimbabwe, Sept–Apr (n ¼ 401, Sept 16, Oct 89, Nov 160, Dec 73, Jan 31, Feb 18, Mar 12, Apr 2 clutches); KwaZulu-Natal and Transvaal, Oct–Feb (mainly Oct– Dec).

approached X with his wings quivering, found a caterpillar and offered it to her; she accepted it, then the 2nd Y flew in and the 2 YY fluttered together for a few s before the intruder again retreated. Later, the Y found another, large caterpillar, chewed it in bill until flaccid, flew to treetop, called loudly and quivered wings. The X perched next to him, quivered wings, accepted the caterpillar, and they copulated (van Someren 1956). Often 3 broods per year (Kenya). NEST: a bowl-shaped, rather ragged and loose structure of grey, dry grass and weedy plant stems, lined with very fine dry grass, often yellowish, fine rootlets and usually hair (B, C). Some nests made almost entirely of rootlets and horsehair; others contain plant heads as well as stems. Nests in southern Africa are generally shallow and rather sprawling; some are smaller, more compact and cupshaped; in NW Nigeria, rather deep, regular cups; in Darfur, Sudan, small, very neat, semi-transparent cups. Int. diam. 45–50, int. depth 30–38. In S Zimbabwe placed mainly in thinly foliaged, second-growth saplings or small trees, 45–200 cm above ground, on horizontal fork or cradled on twigs, often on an outer branch and rather open to view. In Kenya, placed between upright twigs on a horizontal branch, or in multiple upright fork, or in creeper. Mainly 13–17 m above ground in Nigeria and up to 3 and 45 m high in tree in S Zaı¨re. In N tropics, placed in thin thorn bush or small thorny tree, often an Acacia. Pair in Arada, Chad, nested every year inside an abandoned house (Newby 1980). EGGS: 1–2 (n ¼ 10 clutches, Nigeria and Darfur, Sudan; av. 18), 2–4 (n ¼ 133 clutches, southern Africa, av. 24). Ovate, smooth, slightly glossy; white, creamy, pale greenish or bluish white, freckled, hair-lined and scrolled with black, blackish brown and ash- or lilac-grey, in band

INCUBATION:

by X only. Period: 125–13 days. young fed by Y and X. Nestling period: 16–17 days. BREEDING SUCCESS/SURVIVAL: 3rd and 4th eggs in a clutch are often infertile. Mortality rates of eggs and nestlings high (van Someren 1956). DEVELOPMENT AND CARE OF YOUNG:

Key References Byers, C. et al. (1943), Skead, C.J. (1960), van Someren, V.G.L. (1956), Vincent, A.W. (1949).

Emberiza poliopleura

571

Emberiza poliopleura (Salvadori). Somali Bunting; Somali Golden-breasted Bunting. Bruant de Plate 34 Somalie. (Opp. p. 523) Fringillaria poliopleura Salvadori, 1888. Ann. Mus. Civ. Genova, 26, p. 269; Sodde`, Shoa, Ethiopia. Forms a superspecies with E. flaviventris. Range and Status. Endemic resident and partial migrant, Horn of Africa and E Africa. Ethiopia, widespread below 2000 m, in Awash Nat. Park and Valley, in SE and S Ethiopia; locally frequent, with about half the abundance of Cinnamon-breasted Bunting E. tahapisi; common on arid plains at 900–1400 m around Yabelo, Mega and Arero, occurs north to Arba Minch and Negele areas. Records in Tigray Prov. (Cheesman and Sclater 1936) were probably misidenties for E. flavigaster. Somalia, common and widespread where shown; absent from most of N, east of 45
E, from most of centre north of 4
N and from most of S south of 2
N except for Jubba valley; absent just north of Kenya border although common on Kenyan side; in lowlands in S and at 600–1500 m in N. Kenya, fairly common resident in Commiphora/Acacia bush below 1200 m in L. Turkana basin, near Lokichokio and Sudan border; almost throughout, north of 2
N, except between Moyale and Wajir; south to edge of central highlands; also in eastern plateau from Meru Nat. Park to Tsavo Nat. Parks and Mkomazi, where common; coast north of Malindi; common in Tsavo East Nat. Park; a few records in Rift Valley near Kedong, Olorgesailie and L. Magadi. Sudan, uncommon (3 records) in extreme SE. Uganda, Mt Moroto. Tanzania, restricted to NE, west to Rift Valley, south to 4
400 S, east to 38
300 E near Kenya border; common and widespread, e.g. in Mkomazi Game Res.; a record from 33
300 E may be of a vagrant. Density, Tsavo East Nat. Park, Kenya, av. 5 birds per 10 ha (varying monthly from 3 to 7: Lack 1985). Description. ADULT Y: stripe from centre of forehead to centre of nape white, bordered by broad black lateral crown stripe; hindneck pale grey-buff, marked with chestnut; mantle chestnut, feathers broadly fringed pale grey-buff; scapulars blackish, tinged chestnut, broadly fringed pale buff; back grey, black feather bases showing as dusky mottling; rump and uppertail-coverts grey, narrowly fringed whitish. Central tail feathers dark brown, edged buffish; rest of tail feathers blackish brown, T3 tipped white, T4–T5 with extensive white distal areas on both webs, T6 with distal half of inner web and most of outer web white, but black streak along shaft broadening near tip (A). Lores white, linked to broad white superciliary stripe, extending back to below side of nape, and to broad white stripe under eye; black stripe behind eye, linked narrowly to black on side of nape, and by black band around ear-coverts to black lower cheeks and narrow black moustachial stripe. Upper chin and malar area whitish; lower chin and throat to centre of lower breast yellow, extending as broad yellow stripe down centre of belly, upper breast washed orange; sides of neck pale ashy grey, whitish anteriorly; sides of breast and flanks pale grey; sides of lower breast and belly, vent, thighs and undertail-coverts white. Remiges dark grey-brown, primaries narrowly edged grey-buff (whiter proximal to emarginations), secondaries edged greyish white, tertials with outer webs tinged rufous sub-laterally and fringed pale buff, much of outer web of inner feather rufous; primary coverts and alula blackish brown; greater coverts blackish, broadly tipped white, but innermost feathers rufous tipped and fringed buff; median

Emberiza poliopleura

coverts blackish, broadly tipped white; lesser coverts blackish, fringed grey. Underwing-coverts and axillaries whitish with dusky bases. Bill greyish horn above, pinkish white below, culmen slightly raised or markedly curved (A): eyes brown; legs pale pale brown to pale grey. ADULT X: similar to ad. Y, but has dark parts of head pattern brown streaked with black; crown stripe and face stripes tinged buff; mantle feathers usually with distinct dark shaft streaks; grey of hindneck, sides of breast, back and rump tinged olive; lesser covert fringes browner. SIZE (10 YY, 10 XX): wing, Y 71–78 (749), X 66–76 (716); tail, Y 63–72 (668), X 58–72 (662); bill, Y 125–145 (137), X 135–145 (139); tarsus, Y 175–19 (183), X 175–19 (182). WEIGHT: Kenya, unsexed (n ¼ 22) 141–179 (158), 1 Y 164, 1 X 174. IMMATURE: juv. differs from ad. X in having lateral crown stripe, eye stripe and lower cheek mainly brown, pale head stripes strongly tinged grey-buff; mantle streaked tawny brown and blackish; rump and breast sides more buff; yellow restricted below to tinge from throat to upper breast, with band of sharp

572

EMBERIZIDAE

dark brown streaks across upper breast; white tail areas tinged buff; greater covert tips buffy white; median coverts with narrower white tips and blackish tooth along shaft; lesser coverts tipped buffy brown (not grey). NESTLING: at c. 6 days skin dark salmon pink, head blackish, mouth orange with cream-white gape; well-marked lines of black feathers shafts appearing down centre of back, on scapulars and fore-edge of wing.

Field Characters. Length 15 cm. A bunting of arid scrubby country in NE Africa, allopatric from very similar Golden-breasted Bunting E. flaviventris, which inhabits moister woodlands at higher elevations; differs in pale streaking and mottling on rufous mantle, dark, pale-edged scapulars, black-spotted lower back, paler grey rump; pale grey sides of breast and white flanks more extensive, yellow of underparts narrower and more uniform (breast less orange); white of lores almost meets over base of bill (black crown stripes broadly meet bill in Golden-breasted). Juv. like juv. Golden-breasted but paler, mantle stripes more contrasting, breast spotted. Voice. Tape-recorded (B, GREG, LEM, McVIC, PAY, STJ). Song shorter and less variable than Golden-breasted Bunting, typically 4–6 single or double notes repeated at same speed and pitch, often beginning or ending with shorter note: ringing ‘wee-chiwee-chiwee’; slower, deliberate ‘tsit, tsiwee, tsiwee, tsiwee’; softer ‘siwoo, siwoo, siwoo, siwoo, tsi’; sibilant ‘ps’yaa, ps’yaa, ps’yaa, ps’yaa’ or harder ‘s’chaa, s’chaa, s’chaa, s’chaa, s’chaa, s’chaa’; tit-like in form but less forceful. Calls not noted, but probably similar to Golden-breasted Bunting. General Habits. Inhabits thick, dry thornbush and acacia scrub country; often in dry scrub at foot or on sides of isolated kopjes; in Tsavo East Nat. Park, mainly bushland

and wooded bushland, also woody and bushy grassland and woodland (Lack 1985). Often in 2s and 3s, sometimes in small flocks. Uses bushes and small trees freely. Y sings from small tree top and similar eminence. Forages on ground, ‘‘walking around like a lark’’ (Lack 1985). Described as both shy and tame. Sings ‘tizekk’ song up to 5 times from a perch, bird’s head thrown back and bill wide open; a similar note can be uttered in flight, through closed bill of a bird carrying food to nest (Moreau and Moreau 1939). A few birds have been caught at night in Ngulia, SE Kenya, evidently on migration; ‘extralimital’ records in S Kenya and N-central Tanzania also suggest migration. Food. Said to be seeds and insects. Breeding Habits. Solitary nester. NEST: one was a rough cup made of stiff, dead grass stems, unlined; ext. diam. 100; sited in a fork c. 70 cm up in a bush (Moreau and Moreau 1939). EGGS: 2–3, av. (5 clutches) 26. White, with a ring of blackish dots and scrawls (Moreau and Moreau 1939) or scribbled and smudged with sepia and ash-grey (Archer and Godman 1961). SIZE: (n ¼ 3) 182–191  128–136 (187  132). LAYING DATES: Ethiopia, Apr–June, Sept, mainly June and Sept; Somalia, May (n ¼ 4); Tanzania, Nov; E Africa, Region D, Apr–May. In coastal and inland S Kenya breeding seems to peak in Nov–Jan and in N Kenya in Apr–June (Byers et al. 1995). Nothing further known. Key References Byers, C. et al. (1995), Moreau, R.E. and Moreau, W. (1939).

Plate 34

Emberiza affinis Heuglin. Brown-rumped Bunting. Bruant a` ventre jaune.

(Opp. p. 523)

Emberiza affinis Heuglin, 1867. J. Orn., 15, p. 297; no locality – Sennar, Sudan, designated by Heuglin, 1868, J. Orn., 16, p. 76. Range and Status. Endemic resident, Gambia to Ethiopia. Mauritania, uncommon, south of Guidimaka and in upper Senegal R. valley. Senegal, frequent in S and E, absent from NE, rare in N. Gambia, frequent to common in Kiang West Nat. Park, otherwise uncommon: resident and perhaps a wet-season visitor. Guinea-Bissau, several old records in SE (Hall and Moreau 1970). Guinea, only Gaoual area, in NW. Mali, frequent, local, east at least to 8
W; resident in Mandingo Mts but does not occur in Boucle du Baoule´ area (Lamarche 1993); 3 old records on R. Niger east to 8
W (Hall and Moreau 1970). Ivory Coast, 10 records, mainly N Comoe´ Nat. Park, west to Ouangolodougou and M’Bingue´. Burkina Faso, 2 or 3 records, Ouagadoudou area; in 1988 transect, seen only once (Holyoak and Seddon 1989). Ghana, widespread and not uncommon in N; common in Mole Nat. Park in wet season but apparently absent at end of dry season (Taylor and

Macdonald 1978). Togo, 5 records, in extreme N (Mango, Paio and Naboulgou). Niger, records in ‘W’ Nat. Park (May, July) and Tillabe´ri (Nov). Nigeria, fairly frequent in Borgu, north to Shagunu-Luma region (about 10
220 N, 4
270 E); frequent or common in Yankari Nat. Park and frequent southwest to Pandam; elsewhere uncommon to rare (Zaria and Samaru, 5 records in 5 years; Katsina and Kano east to Potiskum, Maiduguri and Yola). Cameroon, uncommon, c. 31 localities (M. Languy, pers. comm.), nigeriae north of 9
N (and once at Mbakaou, Apr, ‘probably migrated out of its normal range’: Louette 1981), and vulpecula on and around Adamawa Plateau in centre, at 750–1050 m. Chad, uncommon; 7 records, north to 10
200 N (Bongor, Dumtar). Central African Republic, records in W from Sarki on Cameroon border and Bouzoum, and in N in Bamingui-Bangoran Nat. Park (where uncommon: 4 records or more), northeast of Nde´le´,

572

EMBERIZIDAE

dark brown streaks across upper breast; white tail areas tinged buff; greater covert tips buffy white; median coverts with narrower white tips and blackish tooth along shaft; lesser coverts tipped buffy brown (not grey). NESTLING: at c. 6 days skin dark salmon pink, head blackish, mouth orange with cream-white gape; well-marked lines of black feathers shafts appearing down centre of back, on scapulars and fore-edge of wing.

Field Characters. Length 15 cm. A bunting of arid scrubby country in NE Africa, allopatric from very similar Golden-breasted Bunting E. flaviventris, which inhabits moister woodlands at higher elevations; differs in pale streaking and mottling on rufous mantle, dark, pale-edged scapulars, black-spotted lower back, paler grey rump; pale grey sides of breast and white flanks more extensive, yellow of underparts narrower and more uniform (breast less orange); white of lores almost meets over base of bill (black crown stripes broadly meet bill in Golden-breasted). Juv. like juv. Golden-breasted but paler, mantle stripes more contrasting, breast spotted. Voice. Tape-recorded (B, GREG, LEM, McVIC, PAY, STJ). Song shorter and less variable than Golden-breasted Bunting, typically 4–6 single or double notes repeated at same speed and pitch, often beginning or ending with shorter note: ringing ‘wee-chiwee-chiwee’; slower, deliberate ‘tsit, tsiwee, tsiwee, tsiwee’; softer ‘siwoo, siwoo, siwoo, siwoo, tsi’; sibilant ‘ps’yaa, ps’yaa, ps’yaa, ps’yaa’ or harder ‘s’chaa, s’chaa, s’chaa, s’chaa, s’chaa, s’chaa’; tit-like in form but less forceful. Calls not noted, but probably similar to Golden-breasted Bunting. General Habits. Inhabits thick, dry thornbush and acacia scrub country; often in dry scrub at foot or on sides of isolated kopjes; in Tsavo East Nat. Park, mainly bushland

and wooded bushland, also woody and bushy grassland and woodland (Lack 1985). Often in 2s and 3s, sometimes in small flocks. Uses bushes and small trees freely. Y sings from small tree top and similar eminence. Forages on ground, ‘‘walking around like a lark’’ (Lack 1985). Described as both shy and tame. Sings ‘tizekk’ song up to 5 times from a perch, bird’s head thrown back and bill wide open; a similar note can be uttered in flight, through closed bill of a bird carrying food to nest (Moreau and Moreau 1939). A few birds have been caught at night in Ngulia, SE Kenya, evidently on migration; ‘extralimital’ records in S Kenya and N-central Tanzania also suggest migration. Food. Said to be seeds and insects. Breeding Habits. Solitary nester. NEST: one was a rough cup made of stiff, dead grass stems, unlined; ext. diam. 100; sited in a fork c. 70 cm up in a bush (Moreau and Moreau 1939). EGGS: 2–3, av. (5 clutches) 26. White, with a ring of blackish dots and scrawls (Moreau and Moreau 1939) or scribbled and smudged with sepia and ash-grey (Archer and Godman 1961). SIZE: (n ¼ 3) 182–191  128–136 (187  132). LAYING DATES: Ethiopia, Apr–June, Sept, mainly June and Sept; Somalia, May (n ¼ 4); Tanzania, Nov; E Africa, Region D, Apr–May. In coastal and inland S Kenya breeding seems to peak in Nov–Jan and in N Kenya in Apr–June (Byers et al. 1995). Nothing further known. Key References Byers, C. et al. (1995), Moreau, R.E. and Moreau, W. (1939).

Plate 34

Emberiza affinis Heuglin. Brown-rumped Bunting. Bruant a` ventre jaune.

(Opp. p. 523)

Emberiza affinis Heuglin, 1867. J. Orn., 15, p. 297; no locality – Sennar, Sudan, designated by Heuglin, 1868, J. Orn., 16, p. 76. Range and Status. Endemic resident, Gambia to Ethiopia. Mauritania, uncommon, south of Guidimaka and in upper Senegal R. valley. Senegal, frequent in S and E, absent from NE, rare in N. Gambia, frequent to common in Kiang West Nat. Park, otherwise uncommon: resident and perhaps a wet-season visitor. Guinea-Bissau, several old records in SE (Hall and Moreau 1970). Guinea, only Gaoual area, in NW. Mali, frequent, local, east at least to 8
W; resident in Mandingo Mts but does not occur in Boucle du Baoule´ area (Lamarche 1993); 3 old records on R. Niger east to 8
W (Hall and Moreau 1970). Ivory Coast, 10 records, mainly N Comoe´ Nat. Park, west to Ouangolodougou and M’Bingue´. Burkina Faso, 2 or 3 records, Ouagadoudou area; in 1988 transect, seen only once (Holyoak and Seddon 1989). Ghana, widespread and not uncommon in N; common in Mole Nat. Park in wet season but apparently absent at end of dry season (Taylor and

Macdonald 1978). Togo, 5 records, in extreme N (Mango, Paio and Naboulgou). Niger, records in ‘W’ Nat. Park (May, July) and Tillabe´ri (Nov). Nigeria, fairly frequent in Borgu, north to Shagunu-Luma region (about 10
220 N, 4
270 E); frequent or common in Yankari Nat. Park and frequent southwest to Pandam; elsewhere uncommon to rare (Zaria and Samaru, 5 records in 5 years; Katsina and Kano east to Potiskum, Maiduguri and Yola). Cameroon, uncommon, c. 31 localities (M. Languy, pers. comm.), nigeriae north of 9
N (and once at Mbakaou, Apr, ‘probably migrated out of its normal range’: Louette 1981), and vulpecula on and around Adamawa Plateau in centre, at 750–1050 m. Chad, uncommon; 7 records, north to 10
200 N (Bongor, Dumtar). Central African Republic, records in W from Sarki on Cameroon border and Bouzoum, and in N in Bamingui-Bangoran Nat. Park (where uncommon: 4 records or more), northeast of Nde´le´,

Emberiza affinis

Emberiza affinis

and in Vakaga region where uncommon (also recorded in S, in Lobaye area: confirmation needed). Sudan, fairly common north to 9
N (affinis, and omoensis near Ethiopian border), affinis probably far more widespread than shown; also (presumably omoensis) about 11
N in Sobat and Blue Nile Districts. Ethiopia, frequent, at least locally in areas shown in NW and SW. NE Zaı¨re, grasslands south to Kibali-Uele R.; Faradje, common around Garamba. Uganda, uncommon at medium elevations in N, south to Murchison Falls Nat. Park, Budongo, Mubende, Nyakwai Hills and Mt Elgon. Kenya, recorded in past only at foot of Mt Elgon, base of Kongelai Escarpment and in N Kerio R. valley. Density of 8 singing birds on 1 km of track, Gambia (Bull. Afr. Bird Club 1995, 2, 124). Description. E. a. affinis Heuglin: S Sudan, N Uganda and adjacent areas of Zaı¨re. ADULT Y: stripe from centre of forehead to centre of nape greyish white, bordered by broad black stripe; hindneck grey-brown; mantle, back and scapulars dull chestnut, with a few narrow dark central feather streaks; rump and uppertail-coverts grey-brown. Tail blackish brown, (T3) T4–T5 with white area on distal part of inner web, T6 with distal half of inner web and edge of outer web white (A). Lores, long superciliary stripe, and below eye to upper cheeks and ear-coverts greyish white; stripe behind eye black, linked by narrow border arround ear-coverts to black lower cheeks and narrow black moustachial stripe; sides of neck grey-brown. Upper chin whitish; lower chin to flanks and belly bright yellow, upper breast tinged rusty; vent and undertail-coverts white. Remiges blackish brown, primary bases narrowly edged buff, outer webs of tertials broadly fringed chestnut; primary coverts and alula blackish brown; greater coverts dark brown, inner feathers broadly fringed chestnut; median and lesser coverts grey-brown. Axillaries greyish white; underwing-coverts greyish white with dusky bases. Bill relatively slender (A), blackish or slate above, pale bluish grey below; eyes dark brown; legs greenish grey to flesh brown. Sexes alike. SIZE (7 YY, 7 XX): wing, Y 70–76 (743),

X 70–75 (720); tail, Y 57–66 (606), X 55–60 (573); bill, Y 12–13 (126), X 12–13.5 (127); tarsus, Y 18–185 (181), X 175–19 (184). IMMATURE: juv. head pattern brown and buffy white, yellow underparts much paler, and less white in outer tail feathers. E. a. nigeriae Bannerman and Bates: Gambia to Nigeria and N Cameroon. Mantle slightly paler than in nominate race, feathers more extensively buff-edged; underparts slightly paler yellow, with whitish submoustachial stripe. WEIGHT: (n ¼ 4, Ghana) unsexed 145–160 (1535). E. a. vulpecula Grote: Cameroon (Adamawa plateau), S Chad and N Central African Republic. Mantle deeper reddish brown, more uniform than in nominate race. E. a. omoensis Neumann: S Ethiopia. Mantle duller brown than in nominate race, with more pronounced streaking. NOMENCLATURAL NOTE: formerly called Emberiza forbesi; for reasons for change, see Dowsett and Dowsett-Lemaire 1993b, p. 379).

Field Characters. Length 125–145 cm. Smallest of the yellow-breasted buntings, differing from Golden-breasted and Somali Buntings E. flaviventris and E. poliopleura in brown rump, wing-bars faint or lacking, and yellow extending to flanks and belly (only chin and undertail white); bill grey, not pinkish brown. Narrow black head stripes and extensive white preorbital region and under eye make head appear whiter than Golden-breasted Bunting. Juv. has rufous upperparts and wings, chestnut lateral crown stripes, unspotted underparts dull yellow with rusty wash on breast. Voice. Tape-recorded (104, B, CHA, MOR, STJ). Song brief (1–15 s) rapid jingle of sharp and tinkling notes, usually ending with short burry trill; shorter version, ‘chachawit, chiwaaa’; for other renditions, see Barlow et al. (1997). Song-like contact (?) call from perched birds, a rapid, liquid and melodious ‘pidru-e-driliwi’, like miniature bulbul (Byers et al. 1995); in flight, short ‘chip’.

General Habits. An uncommon and poorly-known bird. Inhabits dry savanna woodlands, farmed areas and old fields and scrub; degraded, overgrazed and overbrowsed grassy and shrubby savannas. In some regions seems to be loosely associated with human habitation and edges of villages, and in others with riverside vegetation or river floodplains (Gambia, R. Niger, R. Chari, Nile). Said to occur in areas with tall termite mounds, and has been seen inspecting cavities in termite hills at start of nesting season (Byers et al. 1995). Forages on ground but perches freely in bushes and small trees. Y sings from a stump or tree. Unobtrusive; approachable; singing YY prominent. Most W African records appear to be in wet season, about Apr–Sept, because of migration (Byers et al. 1995) or of YY being conspicuous when singing (Barlow et al.

573

574

EMBERIZIDAE

1997); apparent seasonal change in abundance in Mole, N Ghana, (see above) does suggest partial migration. Food. 9 stomachs contained only insects, including termites (Chapin 1954).

Breeding Habits. Barely known. One nest found, in Sudan, but not described. LAYING DATES: Gambia, (singing YY in early rains; accompanied fledgling in Dec); Sudan, Mar.

Plate 34

Emberiza tahapisi A. Smith. Cinnamon-breasted Bunting; African Rock Bunting. Bruant cannelle.

(Opp. p. 523)

Emberiza tahapisi A. Smith, 1836. Rep. Exped. Central Africa, p. 48; source of Vaal R., Transvaal. Range and Status. Resident and partial migrant, subsaharan Africa, SW Arabia and Socotra. E. t. goslingi: Senegal, widespread in rocky habitats in E in dry season and in woodland in N in wet season; uncommon in S (e.g. only 2 records in Parc Nat. de Basse Casamance). Gambia, local but not uncommon breeding visitor, Nov–Mar; mainly in Lower and Middle River areas; occasional on coast. Mauritania, locally common in S; moves north in rains to 17
N, sometimes to 20
N in Adrar. Mali, widespread, quite common, north to 17
N; numerous east of Mopti; moves to north of Tessalit, 20
N, in Aug– Sept. Guinea, occurs in Gaoual area in NW; frequent on Konossou peak in N; frequent in Haut Niger Nat. Park (Nikolaus 2000); rare, Kounounkan. Sierra Leone, small parties mainly of young birds found at 1200 m in Bintumane Peak region and Kulikoro in NE in Mar 1930, thought to be migrants. Liberia, rare: c. 8, Mar, Mt Nimba. Ivory Coast, common around Korhogo, south to Niangbo; otherwise rare: records in W at Mt Touba and Vavoua, in Scentre at Lamto (Salewski and Go ¨ken 1999) and in NE in Comoe´ Nat. Park. Burkina Faso, quite common; in NW recorded only in May–Dec. Ghana, fairly common resident in N (e.g. Bongo Hills, Lawra, Gambaga Escarpment), breeding dry-season visitor at Mole. Togo, common dryseason visitor to N guinean and sudan savannas, Oct–Apr; resident at e.g. Pe´wa, Kara area and Defale. Benin, frequent, south to Be´te´rou area. Niger, frequent in S, north to Tahoua (where common in June, not breeding), Tanout (15
N), and Dillia de Lagane (130 km north of L. Chad; once 27 in Aug); frequent in ‘W’ Nat. Park although uncommon in Niger R. valley north of it; in SE, near L. Chad, noted in Kaadjia and common at Maine´-Soroa, in wet season only; frequent all year in N Aı¨r. Nigeria, in N a widespread, locally common resident and locally abundant migrant; in sahel zone, a wet-season visitor to Malamfatori, June–Oct, with strong passage in Sept and 1st week Oct; at Katsina, small flocks of migrants occur widely from mid May, splitting up in July, birds disappearing in mid Nov; at Zaria, a very common resident and from mid Apr to late May an abundant migrant; in Borgu, frequent in rains, and a marked autumn influx with arrival dates from late Sept to late Oct (5 years); in S, occurs south in dry season to Kabba and to Lagos, Burutu and Enugu (once each, Sept), Port Harcourt (once, Oct), Mambilla and Obudu Plateaux (small parties mainly of young birds in Mar, thought to be migrants). Cameroon, common north of 10
N, frequent south to about 8
N; records in W (Bamenda at 1675– 1830 m; Kounden; Nyasoso, Mt Kupe´). Chad, common east

Emberiza tahapisi

? ? ?

? ? ?

? ? ?

Migrations: see text.

of 19
E in Biltine and Ouaddaı¨; in Ouadi Rime-Ouadi Achim Faunal Res. resident but uncommon in southernmost wadis, but in wet season, when it breeds, an influx as far north as Wadi Haddat; absent from Ennedi; 400 caught in Abe´che´ in a year (Salvan 1968). Central African Republic, occasional, Manovo-Gounda-Saint Floris Nat. Park; Vakaga Pre´f., May–June. Sudan, locally common resident in Darfur, Kordofan and Bahr al Ghazal; barely known in Western Equatoria Prov., but in Eastern Equatoria occurs at Kajo-Kaji on Bahr el Jebel (White Nile), and sure to be on land above 500 m to the west, since abundant at Aba, Zaı¨re, 10 km from Sudan border (at least in Dec, when breeding, Chapin 1954). Racial identity of any birds that may reside in land below 500 m west of Nile between 10
and 16
N requires further study. NE Zaı¨re (besides Aba), records of E. tahapisi in highlands west of L. Albert, south to Semliki (Hall and Moreau 1970), probably refer to goslingi. E. t. septemstriata: E Sudan, west to about Nile, north to SE Kassala Prov. and Blue Nile valley, south to SE Sobat Prov. Ethiopia, frequent in N; southern limits unclear. Eritrea, common near coast and in Mareb R. catchment at

574

EMBERIZIDAE

1997); apparent seasonal change in abundance in Mole, N Ghana, (see above) does suggest partial migration. Food. 9 stomachs contained only insects, including termites (Chapin 1954).

Breeding Habits. Barely known. One nest found, in Sudan, but not described. LAYING DATES: Gambia, (singing YY in early rains; accompanied fledgling in Dec); Sudan, Mar.

Plate 34

Emberiza tahapisi A. Smith. Cinnamon-breasted Bunting; African Rock Bunting. Bruant cannelle.

(Opp. p. 523)

Emberiza tahapisi A. Smith, 1836. Rep. Exped. Central Africa, p. 48; source of Vaal R., Transvaal. Range and Status. Resident and partial migrant, subsaharan Africa, SW Arabia and Socotra. E. t. goslingi: Senegal, widespread in rocky habitats in E in dry season and in woodland in N in wet season; uncommon in S (e.g. only 2 records in Parc Nat. de Basse Casamance). Gambia, local but not uncommon breeding visitor, Nov–Mar; mainly in Lower and Middle River areas; occasional on coast. Mauritania, locally common in S; moves north in rains to 17
N, sometimes to 20
N in Adrar. Mali, widespread, quite common, north to 17
N; numerous east of Mopti; moves to north of Tessalit, 20
N, in Aug– Sept. Guinea, occurs in Gaoual area in NW; frequent on Konossou peak in N; frequent in Haut Niger Nat. Park (Nikolaus 2000); rare, Kounounkan. Sierra Leone, small parties mainly of young birds found at 1200 m in Bintumane Peak region and Kulikoro in NE in Mar 1930, thought to be migrants. Liberia, rare: c. 8, Mar, Mt Nimba. Ivory Coast, common around Korhogo, south to Niangbo; otherwise rare: records in W at Mt Touba and Vavoua, in Scentre at Lamto (Salewski and Go ¨ken 1999) and in NE in Comoe´ Nat. Park. Burkina Faso, quite common; in NW recorded only in May–Dec. Ghana, fairly common resident in N (e.g. Bongo Hills, Lawra, Gambaga Escarpment), breeding dry-season visitor at Mole. Togo, common dryseason visitor to N guinean and sudan savannas, Oct–Apr; resident at e.g. Pe´wa, Kara area and Defale. Benin, frequent, south to Be´te´rou area. Niger, frequent in S, north to Tahoua (where common in June, not breeding), Tanout (15
N), and Dillia de Lagane (130 km north of L. Chad; once 27 in Aug); frequent in ‘W’ Nat. Park although uncommon in Niger R. valley north of it; in SE, near L. Chad, noted in Kaadjia and common at Maine´-Soroa, in wet season only; frequent all year in N Aı¨r. Nigeria, in N a widespread, locally common resident and locally abundant migrant; in sahel zone, a wet-season visitor to Malamfatori, June–Oct, with strong passage in Sept and 1st week Oct; at Katsina, small flocks of migrants occur widely from mid May, splitting up in July, birds disappearing in mid Nov; at Zaria, a very common resident and from mid Apr to late May an abundant migrant; in Borgu, frequent in rains, and a marked autumn influx with arrival dates from late Sept to late Oct (5 years); in S, occurs south in dry season to Kabba and to Lagos, Burutu and Enugu (once each, Sept), Port Harcourt (once, Oct), Mambilla and Obudu Plateaux (small parties mainly of young birds in Mar, thought to be migrants). Cameroon, common north of 10
N, frequent south to about 8
N; records in W (Bamenda at 1675– 1830 m; Kounden; Nyasoso, Mt Kupe´). Chad, common east

Emberiza tahapisi

? ? ?

? ? ?

? ? ?

Migrations: see text.

of 19
E in Biltine and Ouaddaı¨; in Ouadi Rime-Ouadi Achim Faunal Res. resident but uncommon in southernmost wadis, but in wet season, when it breeds, an influx as far north as Wadi Haddat; absent from Ennedi; 400 caught in Abe´che´ in a year (Salvan 1968). Central African Republic, occasional, Manovo-Gounda-Saint Floris Nat. Park; Vakaga Pre´f., May–June. Sudan, locally common resident in Darfur, Kordofan and Bahr al Ghazal; barely known in Western Equatoria Prov., but in Eastern Equatoria occurs at Kajo-Kaji on Bahr el Jebel (White Nile), and sure to be on land above 500 m to the west, since abundant at Aba, Zaı¨re, 10 km from Sudan border (at least in Dec, when breeding, Chapin 1954). Racial identity of any birds that may reside in land below 500 m west of Nile between 10
and 16
N requires further study. NE Zaı¨re (besides Aba), records of E. tahapisi in highlands west of L. Albert, south to Semliki (Hall and Moreau 1970), probably refer to goslingi. E. t. septemstriata: E Sudan, west to about Nile, north to SE Kassala Prov. and Blue Nile valley, south to SE Sobat Prov. Ethiopia, frequent in N; southern limits unclear. Eritrea, common near coast and in Mareb R. catchment at

Emberiza tahapisi 50–1750 m (Zinner 2001). E. t. insularis: Socotra, locally common in mountains between 100 m and 850 m and can be very common in plains down to sea level. E. t. tahapisi: Sudan, fairly common in SE, from about Nile northeast to Boma Hills. Ethiopia, resident, south of septemstriata, mainly above 900 m. Djibouti, scarce to frequent resident; up to 50 in a day (A. Laurent per G. Walsh, pers. comm.). Somalia, locally common breeding resident in NW, east to about Berbera and hinterland up to 1800 m, and single record from hills behind Maydh. Uganda, West Nile Prov. and L. Albert; Kidepo Valley Nat. Park and Mt Moroto; and uncommon but widespread in S and SW. Kenya, widespread but local resident at 400–3000 m: Lokichokio, W and E sides of L. Turkana, W and central highlands, Nairobi Nat. Park, Mara, Amboseli and Tsavo West Nat. Park; mainly a breeding visitor in Dec–Feb in Tsavo East. In Africa south of Equator reaches western coasts but avoids eastern ones; as mapped, but probably much more widespread in W Tanzania, Zaı¨re and Angola. Mainly resident; prone to wandering; may be a breeding visitor in ˆ i, seasonally common at 500–1500 m, S Tanzania. In Malaw uncommon below 500 m, once at 2380 m on Nyika Plateau; marked influx in late Mar. Botswana, recent unusual records from Nossob and 200 in Modipe valley. Zimbabwe, occurs up to 1700 m in Chimanimani Mts, once at 2200 m on Inyanga Downs. In southern Africa, restricted to summer-rainfall regions, except for N Namibia where it occurs in late-summer and autumn rainfall region. Density of 29 pairs (i.e. nests) in c. 100 ha, Zaria, Nigeria (Gartshore 1975); once, 3 breeding pairs in 02 ha (Serle 1940); 9 pairs (i.e. nests) in 60 ha, Cape (Skead 1960) and 20 nests in 6–7 ha, S Zimbabwe (Cumming and Steyn 1966). Description. E. t. tahapisi Smith (includes ‘nivenorum’): S Ethiopia, E Africa and S Zaı¨re to South Africa. ADULT Y: stripe from centre of forehead to centre of nape white with narrow black streaks, bordered by broad black lateral crown stripe; mantle, scapulars and back warm buffy brown, streaked blackish brown; rump and uppertail-coverts buffy brown, mottled darker brown. Tail blackish brown, edges of T1, outer edge of T6 and all feather tips narrowly fringed rich buff when fresh. A white superciliary stripe extends back to below nape, bordered by broad black stripe through lores to upper ear-coverts; below this a white stripe across upper cheek, a black moustachial stripe, broadening on lower cheek, and a narrow white sub-moustachial stripe. Chin and throat sooty black, feathers of chin fringed whitish; rest of underparts cinnamon-rufous, upper breast invaded by short blackish streaks. Flight feathers blackish brown; primaries narrowly edged pale buff, inner feathers with variable rusty border toward base of inner web; secondaries edged cinnamon buff, inner webs often mainly rufous except at tip. Primary coverts and alula blackish brown, narrowly fringed buff; tertials and rest of upperwing-coverts blackish brown, broadly fringed cinnamon-buff. Underwing-coverts and axillaries cinnamon. Upper mandible dark brown; lower mandible yellowish or pinkish; eyes dark brown or dark chestnut; legs pale brown or straw-coloured. ADULT X: similar to ad. Y, but black parts of head browner, white stripes tinged buff; top of head streaked buffy brown and dark brown, paler streaks concentrated along central crown stripe: throat mottled blackish and grey; streaks on mantle, scapulars and back paler and browner. SIZE (10 YY, 10 XX): wing, Y 76–81 (790), X 74–80 (764); tail, Y 60–66 (639), X 60–64 (616); bill, Y 12–13 (127), X 115–13 (123); tarsus, Y 175–19 (181), X 17–19 (180). WEIGHT: Kenya, unsexed (n ¼ 2) 151, 161, X (n ¼ 1) 170; Zimbabwe, Y (n ¼ 1) 132,

X (n ¼ 1) 145; Transvaal, unsexed (n ¼ 1) 142; southern Africa, unsexed (n ¼ 44) 12–22 (148) (Maclean 1993). IMMATURE: juv. similar to ad. X, but pale head stripes buffier; throat paler and browner; upperparts buffier with streaking browner, less distinct; underparts paler, tawny-buff, breast often with faint streaks. NESTLING: E. t. tahapisi: at day 1 skin flesh pink, covered on dorsal surface with tufts of fine grey down; dark line down centre of wings where quills will emerge; bill blackish, gape orangeyellow; eyes closed; legs whitish (Cumming and Steyn 1966). E. t. goslingi (Alexander): Gambia to Sudan west of Nile, and extreme NE Zaı¨re. Throat plain grey in both sexes. Base of outer primaries and all except tips of inner primaries and secondaries rufous; primary coverts and alula fringed rufous; outer webs of tertials, greater coverts and median coverts rufous, and lesser coverts wholly rufous. WEIGHT: N Nigeria, Zaria, (Oct–Dec, breeding) 8 YY and 8 XX, combined av. 147; (non-breeding, n ¼ 44, Nov–Mar) 116–156 (135), (non-breeding, n ¼ 128, Apr–July) 111–150 (131). E. t. septemstriata Ru ¨ ppell: Sudan east of Nile, and Ethiopia (except S). Intermediate between tahapisi and goslingi. Amount of rufous in wing varies, but typically differs from nominate race in having flight feathers more rufous at base and with rufous outer edges; outer edges of primary, greater and median coverts cinnamon-rufous, but less broad than in goslingi. Throat of Y mottled black and grey. E. t. insularis (Ogilvie-Grant and Forbes): Socotra. Like tahapisi but paler above and below, underwing-coverts whiter, black bib more restricted, and inner webs of flight feathers without rufous. TAXONOMIC NOTE: YY of goslingi (N tropics east to R. Nile) have grey throats and rufous wings; YY of other populations (septemstriata, Nile to Red Sea), arabica (SW Arabia), insularis (Socotra) and tahapisi (central Ethiopia to Angola and E South Africa) have black throats and brown wings. Ranges of goslingi and tahapisi appear to meet at W end of the Sudan/Uganda border without hybridization, which suggests that the birds may behave as separate, parapatric species there. Further north, the boundary between goslingi and septemstriata is not precisely known; septemstriata has variably brown or rufous wings and ‘‘seems to be a hybrid population between goslingi and tahapisi’’ (Byers et al. 1995). A case can thus be made for raising goslingi to species rank, so in parts of this account we deal with subspecies separately.

Field Characters. Length 13–15 cm. Widespread in rocky habitats. Darker than House Bunting E. striolata; shares rich cinnamon underparts, but head heavily striped black and white, throat and upper breast unstreaked, black in E and S Africa, grey in W; rufous upperparts streaked dark (plain or lightly streaked in W African House Buntings, but equally streaked in E forms); wing-coverts dark with pale edges (plain rufous in House Bunting); W race goslingi has rufous panel on flight feathers, visible on folded wing and in flight. X has dark brown and buff face stripes, narrow short streaks on crown, smudgy mottling on dark throat; juv. like X but throat pale with fine streaking, face stripes duller, less contrasting. Edges to tail feathers rufous (white in yellow-breasted buntings). Voice. Tape-recorded (22, 86, 88, 99, 104, B, F, LEM, PAY). Short and high-pitched song of hurried, tuneless phrases, including short dry trills, rather disjointed: ‘jajaja-titi’; ‘ti-jaa, yo-tsi’; ‘ti-trrrr-titi-ya-tsi’; ‘ti-jaa, yaasisisi’; in W Africa (104), more chattering quality, ‘chacha-cha-si-cha-cha’; where sympatric with House Bunting, song shorter, e.g. ‘tissy-chaa-tsee’. Call a curious nasal, rolling, ‘j’waa-waah’, like starling Lamprotornis. Other calls

575

576

EMBERIZIDAE

include feeble, seedeater-like ‘pay-way’ or ‘pee-wee’, and in alarm, musical ‘tchrer-ree’ or long-drawn ‘sweeee’ (Skead 1960). General Habits. Inhabits inselbergs, rocky and lateritic ridges and outcrops, boulder-strewn hillsides with open bush, short-grassed open ground with scattered bushes, farmbush (Guinea), bare stony areas and wadis (Chad). Throughout range, particularly favours patches of bare soil: well-used footpaths, dirt roads and gravelly verges of surfaced roads, sheet erosion and bare or sparsely vegetated patches of ground in woodland. Also erosion gullies and dry watercourses with scrub-covered earthen cliffs (Nigeria); dry, rock-strewn areas, irrigated fields, often near running water (Somalia); rocky and eroded hillsides and open woodland with short grass and stony patches (Angola, Dean 2000); dry water courses and river beds in wooded savannas of all types, especially where degraded and ground overgrazed or bare, and abandoned quarries, borrow-pits, mines and copper-workings (Zimbabwe); sometimes forages in cornfields and suburban streets (southern Africa, Skead 1960). Usually in pairs, often in flocks of 3–4, sometimes 6–8, migrants in flocks of up to 40. Single bird or pair can be confiding; large flock less so. Unobtrusive. Forages on ground, on patches of bare soil between rocks or under small, leafless bushes. Gleans fallen grains in standing cereal crops, and flutters at or alights on head of millet in order to take its seeds in situ. Gait on ground a shuffling hop; often moves little more than 1 m per min, pecking at dusty surface of soil for minute seeds. When flushed from ground, often flies no further than nearest bush. Flight undulating, rather weak-looking. On alighting, bird often turns around or shuffles sideways. Sings from top of low bush or from small eminence on ground; occasionally sings from well up on side of large tree. Regularly comes to water to drink, and in Nigeria spends much time foraging on damp sand in erosion gulleys and river beds. Can spend up to 30 min preening and loafing at edge of leafy tree or large, leafless one. A popular cage bird in S Nigeria. In N Nigeria (Zaria), moults remiges after breeding, from mid Apr (latest starting in late May), taking c. 63–86 days (earliest completed in mid June: Fry 1971). Further data on moult regimes, derived from skins of goslingi and tahapisi combined, in Byers et al. (1995). In N tropics, west of 15
E, a classic ‘hump-backed bridge’-pattern migrant (Elgood et al. 1973). Appears to be a year-round resident in most of W African range (red on map) and a dry-season breeding visitor further south (pink on map). Strong passage in mid-W and in NE Nigeria in late Sept/early Oct, of birds presumably moving south and about to breed. Occurs in Koundan, Guinea, in Nov–Dec, in Bintumane Peak region, Sierra Leone, in early Mar, and in S and SE Nigeria in Sept–Mar; whether birds breed there is unknown. In Haut Niger Nat. Park, Guinea, frequent in Mar, otherwise absent (Nikolaus 2000). Strong passage, inferred to be northward, in N-central Nigeria in May. In N, a wet-season visitor to 20
N in Mauritania and Mali in Aug (–Sept), and to S-central Niger northwest of L. Chad, evidently non-breeding (grey on map). Occurs all year in Aı¨r Massif, north to 20
N, but not yet shown to

breed there. Partial migrant in Ouadi Rime-Ouadi Achim Res., Chad, resident in wadis in S and extending into N to breed in rains. Whether resident in or a visitor to S Sudan and NE Zaı¨re is not known; common at Aba in Dec, breeding. Thought to migrate on a broad front, by day and night. In Ethiopia and much of E Africa tahapisi appears to be essentially sedentary but in SE Kenya mainly a visitor, in ˆ i and N Zambia a dry-season visitor, Dec–Feb. In Malaw Apr–Oct, a few birds remaining all year; in S Zambia more are year-round residents, and visitors arrive earlier, in Feb–Mar. In SE Africa it has been regarded as resident but nomadic when not breeding, and in drier regions may quit an area when surface water dries up; but in addition to such local movements it is clearly a regular, long-distance migrant. Bird ringed in N Cape Prov., July, recovered next May 1037 km to northeast in Mwenezi District, Zimbabwe. No obvious seasonal change in abundance in South Africa south of 26
S (nor in Namibia) but in N Transvaal, E Botswana and S quarter of Zimbabwe markedly commoner in Nov–Mar than in May–Sept and in rest of Zimbabwe commoner in Jan–June than in July–Dec (Harrison et al. 1997). In Zimbabwe very large numbers arrive in early Dec to breed in e.g. Bulawayo and Esigodini area; concentrations can occur outside breeding season, but whether of migrants or local nomads fleeing arid conditions is not clear. Food. Small seeds and some insects. Seeds of Eragrostis sp.; in captivity, seeds of grasses E. tef, Chloris virgata, Setaria appendiculata, Hyparrhenia cymbaria and H. hirta. Chicks given grass seeds and insects including small caterpillars; melolonthid beetles in stomachs. In captivity, eats mealworms, termites, gnats, fly larvae, wasp larvae, thrips nymphs, caterpillars, spiders, millipedes and small slugs (Brickell and Arnold 1990, Brickell and Konigkramer 1997). Breeding Habits. Solitary nester; monogamous; territorial (Gartshore 1975). Courtship behaviour almost unknown in this retiring, unobtrusive species, except that Y sometimes chases X in air. NEST: a shallow cup, in shady place on ground. E. t. goslingi: one, on bare, sloping rock slab overhung by tuft of grass, at base of huge granite outcrop, consisted mainly of delicate rootlets and finely divided grass-heads, with pieces of stout grass stems up to 60 mm long arranged around the outside; base of nest solidified and stuck to rock substrate with mud. Another, in crevice in a pile of stones by tin mine, had slight foundation of dead leaves, twigs, grass-heads and short lengths of grass stem, the cup made of fine grass roots with a scant lining of grass heads c. 75 mm long. A third was in small depression in the side of a furrow under large tree in guinea-corn field. A fourth, on open ground at base of small tree, was shallow saucer made of grass stems, fine ones inside, coarser ones outside. Size: ext. depth 15, int. depth 10, int. diam. 75 (another 55). E. t. tahapisi: base of nest made of twigs, small sticks, pieces of bark, thick coarse grass, weedy plant stems and occasional small seed pods; cup is shallow, a thin, neat network of pieces of dry, yellowish grass or fine rootlets;

Emberiza striolata nests in a smooth rock hollow can have little or no base and are sometimes sparsely lined. Sited often around mine dumps, on open gravelly ground, in rock niche 2–3 (–6) m up fissure or cliff face, in earthen banks, old earthenware pots, under boulder, next to brick or stone on ground. Adjacent nests often only 20–30 m apart. SIZE (n ¼ 19): width of base 90–140 (121), width of cup 50–71 (59), depth of cup 19–28 (24). Built by X only in 4–13 days. EGGS: goslingi: 2–3 (n ¼ 36 clutches, Nigeria), av. 264. Ovate, glossy or only slightly so, smooth. Pale greyish green, heavily spotted with dark brown, or with bold spots and blotches coalescing at large end, with pale grey-purple or violet undermarkings, or spotting rather uniform. SIZE: (n ¼ 8) 166–232  121–134 (188  1295); also (n ¼ 5) 160–175  125–130 (166  127). E. t. tahapisi: 2–4 ˆ i), av. 286; and 3–4 (n ¼ 30 (n ¼ 43 clutches, Malaw clutches, Zimbabwe), av. 327. White or pale bluish or greenish, evenly speckled, or blotched dark brown or redbrown with or without a dark cap at broad end; usually with underlying greyish speckles. SIZE: (n ¼ 16) 181– 200  130–142 (189  136), also (n ¼ 157) 160–197  122–140 (179  133). LAYING DATES: goslingi: Gambia, Nov; Mauritania, (breeds July–Nov); Mali, (breeds July–Dec); Ghana, Jan; Nigeria, (Kano) July, (Jos Plateau) Dec, Jan, (Zaria, n ¼ 29) late Sept to early Dec: in 1974 most nests built soon after last rainfall in mid Oct; fledglings numerous from early Dec; only a few fledglings early Jan; a few ads still in breeding condition until mid Feb (Gartshore 1975); Sudan, Nov–Jan; Zaı¨re (Aba), Dec; septemstriata: no breeding records; insularis: Socotra, Jan; tahapisi: Somalia, June; E Africa, Region B, Mar; Region C, Apr, May, July (mainly May; breeds at end of rains and early in dry season), Region D, Jan–June (breeds mainly in rains); Zaı¨re

577

ˆ i, Apr–July; Zimbabwe, (Lubumbashi), May–June; Malaw Nov–June (12 Nov–Dec, 37 Jan, 44 Feb, 73 Mar, 20 Apr, 2 May–June); South Africa, Free State, Dec–Apr, KwaZuluNatal, Nov–Feb, Transvaal, Oct–Mar (mainly Jan). INCUBATION: by X in goslingi; Y generally stays nearby whilst X on nest. By X (mainly) and Y in tahapisi. Period: (n ¼ 3) 12 days (Nigeria), 12–14 days (Zimbabwe). DEVELOPMENT AND CARE OF YOUNG: goslingi: nestling period (n ¼ 3) 12–15 days, fledglings being able to fly well. For growth curves of young in broods of 2 and of 3, see Gartshore (1975); young reach 90% of ad. weight in 12 days; 55 fledglings of various ages weighed 120–165 (135) g. Young continue to feed within parental territory for 3 weeks after leaving nest, still soliciting food at the end of that time but generally unsuccessfully. A ringed juv. retrapped within 150 m of nest site 61 days after fledging. E. t. tahapisi: for table of development see Cumming and Steyn (1966); young fed only by X, Y often perching nearby and singing strongly. BREEDING SUCCESS/SURVIVAL: goslingi: 39% of eggs laid were taken by predators (or were infertile or abandoned); of nestlings, 29% taken by predators, 7% killed by ants and 64% survived; of 16 eggs in clutches of 2, 10 produced fledglings (625% success; 125 fledglings per nest); of 63 eggs in clutches of 3, 23 produced fledglings (364% success; 109 fledglings per nest) (Zaria, Nigeria: Gartshore 1975). E. t. tahapisi: in one study very high nest failure rate was attributed to small predators such as mongooses which followed human scent trail to nests; of 48 young hatched, only 15 fledged successfully (Cumming and Steyn 1966). Key References Byers, C. et al. (1966), Fry, C.H. (1971), Gartshore, M.E. (1975), Harrison, J.A. et al. (1997), Serle, W. (1940, 1943b), Vincent, A.W. (1949).

Emberiza striolata (Lichtenstein). House Bunting. Bruant striole´.

Plate 35

Fringilla striolata Lichtenstein, 1823. Verz. Doubl. Zool. Mus. Berlin, p. 24; Ambukol, Nubia, Berber, Sudan.

(Opp. p. 538)

Range and Status. Deserts of NW, S and E borders of Sahara, Red Sea hills, N Somalia, Rift Valley in SW Ethiopia and deserts in N Kenya, Sinai, S Levant, W Arabia and parts of central, S and E Arabia, and Iran to NW India. Resident, nomadic or disperses widely in some regions and moves to lower ground in winter in others. Vagrant, Canary Is (sahari), S Spain, Cyprus, Senegal and Gambia. Morocco, resident, almost throughout, north to Rabat, Fe`s and Oujda where frequent, east to Figuig; in S, very common in towns and villages, south at least to Oued Dra and Tan-tan; occurs in N Western Sahara (where may well be widespread, although data are lacking). At Errachidia, local breeding population supplemented by visitors in winter. Has spread northward in Morocco, by 200 km along coast in 35 years until 1902, reaching Casablanca in 1965, Rabat in 1983 and Sale´ by 1992; reached Fe`s in 1985 and Oujda in 1980 (maps in Courteille and The´venot 1988). Records at Ceuta and Tangier. From coast up to

2000 m in Haut Atlas. Algeria, locally common or very common in towns and villages in a narrow band along southern slopes of Atlas Saharien and Plateau de Tademait from Aı¨n Sefra to Laghouat thence northeast to Kalaa des Beni Hammad (80 km southwest of Se´tif); commoner around El Kantara, Biskra and in villages along the Djemorah; in N Sahara occurs in Berriane, Ghardaı¨a and Metlili Chaamba and abundant in M’zab oasis, but does not occur in El Gole´a, Hassi Fahl, Guerrara, Ouargla, Touggourt or El Oued; in central Sahara in Tassili N’Ajjer and Hoggar Mts; abundant in Tassili, widespread for most of year but concentrating near oases when cereals and legumes are harvested in Apr–May and millet in Oct–Nov (Laferre`re 1968); common around Tamanrasset and in SW Hoggar Mts at 1350–1600 m. Tunisia, spread to several new localities in 20th century; numerous in oases of Nefta, Douz, Tozeur, El Hamma and Kebili; rather scarce in desert villages in Nefta-Kairouan-Tathouine region; occurs

Emberiza striolata nests in a smooth rock hollow can have little or no base and are sometimes sparsely lined. Sited often around mine dumps, on open gravelly ground, in rock niche 2–3 (–6) m up fissure or cliff face, in earthen banks, old earthenware pots, under boulder, next to brick or stone on ground. Adjacent nests often only 20–30 m apart. SIZE (n ¼ 19): width of base 90–140 (121), width of cup 50–71 (59), depth of cup 19–28 (24). Built by X only in 4–13 days. EGGS: goslingi: 2–3 (n ¼ 36 clutches, Nigeria), av. 264. Ovate, glossy or only slightly so, smooth. Pale greyish green, heavily spotted with dark brown, or with bold spots and blotches coalescing at large end, with pale grey-purple or violet undermarkings, or spotting rather uniform. SIZE: (n ¼ 8) 166–232  121–134 (188  1295); also (n ¼ 5) 160–175  125–130 (166  127). E. t. tahapisi: 2–4 ˆ i), av. 286; and 3–4 (n ¼ 30 (n ¼ 43 clutches, Malaw clutches, Zimbabwe), av. 327. White or pale bluish or greenish, evenly speckled, or blotched dark brown or redbrown with or without a dark cap at broad end; usually with underlying greyish speckles. SIZE: (n ¼ 16) 181– 200  130–142 (189  136), also (n ¼ 157) 160–197  122–140 (179  133). LAYING DATES: goslingi: Gambia, Nov; Mauritania, (breeds July–Nov); Mali, (breeds July–Dec); Ghana, Jan; Nigeria, (Kano) July, (Jos Plateau) Dec, Jan, (Zaria, n ¼ 29) late Sept to early Dec: in 1974 most nests built soon after last rainfall in mid Oct; fledglings numerous from early Dec; only a few fledglings early Jan; a few ads still in breeding condition until mid Feb (Gartshore 1975); Sudan, Nov–Jan; Zaı¨re (Aba), Dec; septemstriata: no breeding records; insularis: Socotra, Jan; tahapisi: Somalia, June; E Africa, Region B, Mar; Region C, Apr, May, July (mainly May; breeds at end of rains and early in dry season), Region D, Jan–June (breeds mainly in rains); Zaı¨re

577

ˆ i, Apr–July; Zimbabwe, (Lubumbashi), May–June; Malaw Nov–June (12 Nov–Dec, 37 Jan, 44 Feb, 73 Mar, 20 Apr, 2 May–June); South Africa, Free State, Dec–Apr, KwaZuluNatal, Nov–Feb, Transvaal, Oct–Mar (mainly Jan). INCUBATION: by X in goslingi; Y generally stays nearby whilst X on nest. By X (mainly) and Y in tahapisi. Period: (n ¼ 3) 12 days (Nigeria), 12–14 days (Zimbabwe). DEVELOPMENT AND CARE OF YOUNG: goslingi: nestling period (n ¼ 3) 12–15 days, fledglings being able to fly well. For growth curves of young in broods of 2 and of 3, see Gartshore (1975); young reach 90% of ad. weight in 12 days; 55 fledglings of various ages weighed 120–165 (135) g. Young continue to feed within parental territory for 3 weeks after leaving nest, still soliciting food at the end of that time but generally unsuccessfully. A ringed juv. retrapped within 150 m of nest site 61 days after fledging. E. t. tahapisi: for table of development see Cumming and Steyn (1966); young fed only by X, Y often perching nearby and singing strongly. BREEDING SUCCESS/SURVIVAL: goslingi: 39% of eggs laid were taken by predators (or were infertile or abandoned); of nestlings, 29% taken by predators, 7% killed by ants and 64% survived; of 16 eggs in clutches of 2, 10 produced fledglings (625% success; 125 fledglings per nest); of 63 eggs in clutches of 3, 23 produced fledglings (364% success; 109 fledglings per nest) (Zaria, Nigeria: Gartshore 1975). E. t. tahapisi: in one study very high nest failure rate was attributed to small predators such as mongooses which followed human scent trail to nests; of 48 young hatched, only 15 fledged successfully (Cumming and Steyn 1966). Key References Byers, C. et al. (1966), Fry, C.H. (1971), Gartshore, M.E. (1975), Harrison, J.A. et al. (1997), Serle, W. (1940, 1943b), Vincent, A.W. (1949).

Emberiza striolata (Lichtenstein). House Bunting. Bruant striole´.

Plate 35

Fringilla striolata Lichtenstein, 1823. Verz. Doubl. Zool. Mus. Berlin, p. 24; Ambukol, Nubia, Berber, Sudan.

(Opp. p. 538)

Range and Status. Deserts of NW, S and E borders of Sahara, Red Sea hills, N Somalia, Rift Valley in SW Ethiopia and deserts in N Kenya, Sinai, S Levant, W Arabia and parts of central, S and E Arabia, and Iran to NW India. Resident, nomadic or disperses widely in some regions and moves to lower ground in winter in others. Vagrant, Canary Is (sahari), S Spain, Cyprus, Senegal and Gambia. Morocco, resident, almost throughout, north to Rabat, Fe`s and Oujda where frequent, east to Figuig; in S, very common in towns and villages, south at least to Oued Dra and Tan-tan; occurs in N Western Sahara (where may well be widespread, although data are lacking). At Errachidia, local breeding population supplemented by visitors in winter. Has spread northward in Morocco, by 200 km along coast in 35 years until 1902, reaching Casablanca in 1965, Rabat in 1983 and Sale´ by 1992; reached Fe`s in 1985 and Oujda in 1980 (maps in Courteille and The´venot 1988). Records at Ceuta and Tangier. From coast up to

2000 m in Haut Atlas. Algeria, locally common or very common in towns and villages in a narrow band along southern slopes of Atlas Saharien and Plateau de Tademait from Aı¨n Sefra to Laghouat thence northeast to Kalaa des Beni Hammad (80 km southwest of Se´tif); commoner around El Kantara, Biskra and in villages along the Djemorah; in N Sahara occurs in Berriane, Ghardaı¨a and Metlili Chaamba and abundant in M’zab oasis, but does not occur in El Gole´a, Hassi Fahl, Guerrara, Ouargla, Touggourt or El Oued; in central Sahara in Tassili N’Ajjer and Hoggar Mts; abundant in Tassili, widespread for most of year but concentrating near oases when cereals and legumes are harvested in Apr–May and millet in Oct–Nov (Laferre`re 1968); common around Tamanrasset and in SW Hoggar Mts at 1350–1600 m. Tunisia, spread to several new localities in 20th century; numerous in oases of Nefta, Douz, Tozeur, El Hamma and Kebili; rather scarce in desert villages in Nefta-Kairouan-Tathouine region; occurs

578

EMBERIZIDAE

Emberiza striolata

?

? ?

? ?

?

in hills and plains from Gafsa and across Feriana and Sbeitla to Jebel Cherahil; absent from central towns, e.g. Maknassy and Mezzouna. Libya, local along higher parts of Jebel Nafusa escarpment and in adjacent villages, from Nalut east to Gharyan; in Sahara, common around Ghat (24
580 N, 10
110 E) on Jebel Akakus, an eastern extension of Algerian Tassili N’Ajjer highlands, but absent from Serdeles, 120 km to northeast. Egypt, in extreme SW corner occurs and probably breeds on Jebel Uweinat (and occurs on Sudan side of the hills: see below); in SE, common in Jebel Abraq area, locally common in Jebel Elba, and occurs in Jebel Shellal. Mauritania, absent from Zemmour region in NW; occurs in villages and near habitations on ground above 500 m in Adrar, Tagant and Assaba regions and on Massif du Rkiz and at Ayoun el Atrouss; a considerable population along wadi systems of Dhar Tichitt and Dhar Oualata, east to Oualata. Gambia and Senegal, a pair at Albadarr (Albreda) in Dec 1926 seemed ‘at home’ around a house, roosting in a garden hut, and might have nested had the Y not been caught and kept caged; 1 at Barra in Feb 1976 and 1 on cliffs at Popenguine, Apr 1983, were presumably vagrants. Mali, not uncommon but local, between 14
and 17
N, in inland delta of Niger and in sahel zone, generally in rocky areas; frequent around Tombouctou (Timbuktu); north to 20
300 N in Adrar des Ifoghas and the Time´trine. Niger, common in N and central Aı¨r Massif (Aguellal, Igouloulouf to Tebernit) and numerous in SW around Agade`s; common in Zinder region and at Termit Kaoboul. Chad, widespread in Tibesti; another population widespread and locally common at about 700 m in Ennedi and Biltine, south to Abe´che´ and west to between Ouadi Sofaya and Elela (where perhaps only a non-breeding visitor: Newby 1980). Sudan, in extreme NW on Jebel Uweinat; fairly common

in Darfur plateau country, Jebel Marra and Berti Hills north of El Fasher, and above 500 m in S Kordofan, between El Obeid and Talodi; records in Nile valley including one in extreme S; seasonally very common in Red Sea hills. Eritrea, coastal plains and foothills below 650 m, from Sudan border to at least Massawa; records from Denakil coast. Djibouti, common on coast from Tadjoura to Sagallou (38 counted in a day); record inland at Bankoule´; 8 in N at Khor Angur. Somalia, uncommon and local, presumed resident (no breeding records), on N coastal plain from 47
E to Raas Caseyr and along Wadi Dharoor/Wadi Dhuudo, but locally common (e.g. at Iskushuban) or very common (e.g. hot springs near Boosaaso); records at Berbera and west of Dan Gorayo; occurs from sea level up to 1650 m. Ethiopia, striolata frequent in lowlands in NE and S, saturatior uncommon in SW (Urban and Brown 1971); several birds collected in Rift Valley, but not found by Benson (1946) and absent from e.g. Nechisar Nat. Park. Adjacently in N Kenya, occurs at 400–800 m, locally but sometimes commonly, from E shores of L. Turkana to Turbi and N edge of Dida Galgalu Desert, east to Moyale, south to Loiengalani and base of Mt Marsabit; record from Ferguson’s Gulf; common around Kamathia near Sudan border. In west, sahari and sanghae are parapatric with Cinnamonbreasted Rock Bunting E. tahapisi, but in east striolata and saturatior contact E. tahapisi only locally (and in SW and S Arabia the 2 species are sympatric: Jennings 1995 and as mapped here). Density of 2 pairs per ha, Marrakech; 10 singing YY in 1 km transect, Casablanca. Densities in rural and desert areas much less. Description. E. s. sahari Levaillant (includes ‘theresae’): Morocco to NW Libya, south to N Niger and N Chad; perhaps this race in Mauritania; vagrant NW Egypt. ADULT Y: forehead to hindneck pale grey-buff, sharply streaked blackish brown; upper mantle buffy brown with a few long blackish streaks; lower mantle, back and scapulars tawny brown with shorter, less prominent dark brown streaks; rump and uppertail-coverts plain cinnamonrufous. Tail feathers dark brown, narrowly fringed rufous, T6 with rufous outer web and tip. Supercilium poorly marked, greyish white flecked with black, bordered by dark grey streak through lores and more broadly through upper ear-coverts; below this a grey-buff patch across upper cheek, a dark grey moustachial stripe broadening on lower cheek, a pale grey-buff submoustachial stripe, and a narrow, poorly defined dusky malar stripe; sides of neck pale grey-buff, mottled dusky. Upper chin whitish; lower chin to upper breast pale grey, streaked blackish; rest of underparts cinnamon-rufous. Remiges blackish brown, secondaries and primaries proximal to emarginations broadly edged cinnamon-buff, tertials fringed rufous-cinnamon; alula, primary coverts and greater coverts blackish brown, broadly fringed rufous-cinnamon; median and lesser coverts wholly rufous-cinnamon. Underwing-coverts and axillaries cinnamon, axillaries with dusky bases; undersides of inner borders of flight feathers broadly rufous. Birds from S edge of Sahara are slightly darker than those of N Africa. Bill blackish brown above, light brown or yellowish below; eyes brown or orange-brown; feet pale brown. ADULT X: forehead to hindneck pale tawny or buffy brown, sharply streaked dark brown; head pattern less distinct than in ad. Y, eye-stripe and moustachial stripe dusky brown, pale areas buff; chin to upper breast buffy brown with faint fine streaks; otherwise as ad. Y. SIZE (10 YY, 10 XX): wing, Y 73–80 (770), X 72–77 (746): tail, Y 61–67 (641), X 57–67 (619); bill, Y

Emberiza striolata 125–135 (130), X 12–135 (127); tarsus, Y 17–19 (179), X 165– 18 (175). WEIGHT (Algeria and Chad): Y (n ¼ 10) 12–18 (151), X (n ¼ 7) 13–16 (145); (Niger) Y (n ¼ 3) 148, 152, 168 (Cramp and Perrins 1994). IMMATURE: juv. similar to ad. X, but browner, face pattern less distinct; streaks on crown, breast and upperparts more diffuse. NESTLING: dense whitish grey down on head and upperparts; mouth and tongue deep pink, tongue with prominent pale pink spurs distally; mouth with pale yellow borders, whiter at gape (P. Castell, Bull. Afr. Bird Club 7, 2000, 104–106, photo). E. s. sanghae Traylor: S Mali. Darker, more chestnut than all populations of sahari; ground colour of crown and breast darker grey. The species is variable in any one region, and validity of this race was doubted by Lamarche (1981) E. s. striolata Lichtenstein: NE Sudan, Eritrea and N Somalia. Head pattern bolder in both sexes than in sahari; upperparts more buffish and distinctly streaked; underparts paler cinnamon. Y has eye-stripe and moustachial stripe black, superciliary stripe and sub-moustachial stripe white, and grey chin to upper breast more boldly mottled. Slightly smaller than sahari: wing (11 YY) 75–78 (770). E. s. saturatior (Sharpe): central Sudan, SW Ethiopia and NW Kenya. Y with grey face stripes like sahari, but both sexes well streaked on crown, upperparts and chin to upper breast; darker and browner than sahari above and below, with narrower rufous fringes on tertials and primary and greater coverts. E. s. jebelmarrae Lynes: Jebel Marra (W Sudan), intergrading with sahari in NE Chad and NW Sudan. Like saturatior, but more richly coloured and more heavily streaked; underparts deep rufous. Larger: wing (10 YY) 80–84 (821).

Field Characters. Length 13–14 cm. A northern bunting of semi-arid regions. Head less strikingly marked than Cinnamon-breasted Bunting; race sahari (N and W Africa) has grey head and breast with fine dark streaks, face stripes light and dark grey, rest of body, including wings, mainly cinnamon-rufous, with few if any streaks on mantle; at a distance appears uniform grey and rufous (Barlow et al. 1997); tail dark with rufous edges. Sympatric Cinnamonbreasted Bunting has plain grey throat, streaked upperparts, wing feathers dark with pale edges except for rufous panel on flight feathers. X and juv. browner and plainlooking, with inconspicuous face stripes. Rock Bunting E. cia has cinnamon body but plain grey head with triangular black face pattern, prominent white supercilium, much white in tail. In E Africa nominate striolata is paler and less rufous than Cinnamon-breasted, with streaked upperparts; face stripes well marked, but lacks the black-bordered white central crown stripe; X and juv. paler and more washed-out than Y, face stripes less distinct. Voice. Tape-recorded (92, 105, 112, B, HAZ). Song variable, usually short, mixing high-pitched whistles and scratchy notes, typically including up-slurred ‘chi-wee’ and/or down-slurred ‘wee-chu’: monotonous ‘chi-wee, chiwee, chi-wee, chyu-wee’; more varied ‘chi-chu-chee cheew, chu-cheep’; longer ‘wree-wree, chacha-wree-chew, chi-chiwa-cheew’; some longer versions have warbling quality and end with down-slurred ‘cheeer’, like N American House Finch Carpodacus mexicanus, ‘cherchercher, chichi-werchee, chawee, cha-cheeer’. Common call a nasal ‘djer-zherwy’ or ‘ja-jaanh’, like Lamprotornis starling; also harsh ‘jaw’ or ‘jeeaw’; clear, down-slurred whistle, ‘deeeeooo’; short ‘dui’;

and hard ‘juk’ like quailfinch Ortygospiza. For further songs and calls see Cramp and Perrins (1994).

General Habits. Inhabits oases, rocky wadis with sparse growth of acacias, euphorbias and tamarisks, arid, stony plains, sandy wastes and sparsely vegetated country, windpolished rock levels and outcrops with plant growth only in cracks and crevices, and bare rock hillsides in Saharan massifs. Western races strongly associated with human habitations: desert encampments, hamlets at livestock enclosures, dairy farms, fuel dumps, food stores, wells, water holes and water-bottling plants; in NW Africa inhabits even towns and cities; nests in centre of Marrakech (for instance, on Lyce´e Victor Hugo) and Rabat (on Exhibition Hall of ‘Les Oiseaux du Maroc et du Monde’ (1990)); frequents ruins, mud walls, yards, rubbish tips, dung heaps, buildings under construction and noisy thoroughfares, and in many towns more familiar than House Sparrows Passer domesticus. Enters houses, shops, restaurants and mosques, to feed and nest; even known to sing indoors; welcomed and treated with respect by Arab peoples. Frequents settlements in N Kenya desert and villages and towns in N Somalia (saturatior), but striolata is not associated with human habitations at all, rural or urban, in Eritrea (nor in Arabia). In pairs or family parties. Sometimes feeds in loose association with small party of Cinnamon-breasted Buntings E. tahapisi, Desert Larks Ammomanes deserti or Trumpeter Finches Bucanetes githagineus, and can gather in flocks of 20 or 30 at watering places or at spilled grain. Not shy, and sahari can become very tame – ‘fearlessly entered the hotel dining room through the kitchen and picked up crumbs at our feet’ (Bannerman and Priestley 1952); striolata is very shy, however. Forages entirely on ground, on sandy and rocky surfaces with thin herb layer and scattered shrubs. Pecks into small rock crevices; stands upright to pull seeds off grass heads. Jumps up to pull down stem, then stands on it and extracts seeds from inflorescence (A, Fry and Eriksen 1994). Oftens comes to drink at puddle or seepage. Sings from low eminence, rooftop, wall, or rock, or from the ground.

579

580

EMBERIZIDAE

Food. Seeds of wild grasses, barley and other crops in spring and millet in autumn (NW Africa, W Sahara); spiders, ants and other small insects when breeding. Seeds, flowers and fruits of Salvadora persica, Ziziphus, Heliotropium, Phragmites, Phalaris and Bromus. Breeding Habits. Solitary nester; monogamous; territorial when breeding. Semi-colonial: in e.g. Marrakech, breeds numerously in some districts but absent from others, apparently suitable, only a km away. Pair remains together all year. Territories either well separated or adjacent; nests occasionally only 10–30 m apart. Nest or nest site often reused; one nest in Casablanca used 11–12 times in 5 years. 2–3 broods per season, occasionally 4; usually 3 in Marrakech, Morocco, from beginning of Mar to end of Aug (Roux et al. 1990); 2 around Djanet, Tassili, Algeria (Laferre`re 1968). Y defends territory by singing, from 1 preferred site (house wall or garden fence in Morocco, rock in E Sudan) and 2–3 subsidiary ones. Sings all year, but mainly in Nov–May (Morocco), in mornings, for several hours. Y pursues others in flight; they sometimes fight and fall to ground. On ground, rival YY face each other, give shrill calls, vibrate wings and move sideways. Y and X together vibrate wings and press body close to ground, perhaps a prelude to copulation, which takes place on ground (N. Chakir in Cramp and Perrins 1994). NEST: a small cup on a rough foundation of twigs, roots, straw and grass stems, lined with wool, hair, man-made materials and plant down. In Casablanca, Morocco, 16 out of 40 nests sited inside building on ledge, beam or in crevice, 10 on window-sill or balcony, 10 in hole in rockface or wall, and 4 in odd places including a mail-box and a fuse-box. Always in a sheltered place; usually 2–3 m above ground, once on ground under a bush; once on a palm trunk. In Tafilalet, 22 out of 31 nests in buildings or on sea cliffs, and 9 in clefts and crannies in rocks. SIZE (n ¼ 4, Algeria): ext. diam. 140–160 (150), int. diam. 70–80 (73), ext. height 50–60 (56), int. depth 15–20 (18). Built by X, Y helping at start (N. Chakir in Cramp and Perrins 1994); takes 4–7 days to restore old nest and 7–10 days or (another study) 20 days to build new one. EGGS: 1–5 (NW Africa, n ¼ 156 clutches, av. 313); 2–3 (Morocco, n ¼ 35 clutches, av. 269); also Morocco, n ¼ 84

clutches, av. 313 (Courteille and The´venot (1988). Laid daily, in morning. Sub-elliptical, smooth, faintly glossy; whitish, tinged blue or green, speckled with purplish brown forming a ring around large end, with faint undermarkings of purplish grey smears, or rufous-brown with mauve patches. SIZE: (sahari, n ¼ 14) 168–222  137– 146 (195  141), (n ¼ 44) 190–203  130–160; (striolata, n ¼ 15) 180–210  135–150 (193  142). LAYING DATES: Morocco, Feb–Oct, mainly late Feb to early June; Algeria, Mar (Ghardaı¨a), Oct (Djanet, Tassili, and nestlings Apr, Oct, Dec: Laferre`re 1968); Tunisia, Apr; Egypt, Jebel Abraq (fledglings Apr); Mali, (breeds throughout the year); Niger, June (Aguellal), Sept (Kori Tarare and Aoude´ras), (accompanied young, Agade`s, Feb); Sudan, Darfur, Feb, Oct, Nov; Eritrea, (gonads active, Jan). INCUBATION: by X alone; begins as soon as 1st egg laid. Period: 12–16 (13–14) days, or 14–16 days (Roux et al. 1990). In one nest all 3 eggs hatched on same day, but usually hatching is on consecutive days. DEVELOPMENT AND CARE OF YOUNG: nestlings fed by both parents, on regurgitated food. Data on feeding and growth rates in Courteille and The´venot (1988). Young brooded by X constantly for first 4 days and nights; X no longer sleeps in nest from 8th night. Eggshells and faecal sacs eaten by X; later, faecal sacs carried away and dropped. Nestling period: 17–19 days. In brood of 3, 2 leave nest on one day and the 3rd, the youngest, on next day. Young return to nest to sleep for first few nights. Out of nest, young fed by both parents, mainly Y, for 5–20 days (since X soon occupied with a new clutch) or 8 days (Roux et al. 1990). 2 young roosted with parents for >25 days after leaving nest. BREEDING SUCCESS/SURVIVAL: in Morocco (n ¼ 32 nests) 56% of 86 eggs hatched and 50% produced fledglings, also (n ¼ 58 nests) 66% of 169 eggs hatched and 57% gave rise to fledglings; mortality mainly due to human disturbance and cats. Also Morocco (n ¼ 174 clutches), hatching success 657%, rearing success 868%, overall fecundity 57% (Courteille and The´venot 1988). Key References Bannerman, D.A. (1948), Bannerman, D.A. and Priestley, J. (1952), Courteille, C. and The´venot, M. (1988), Cramp, S. and Perrins, C.M. (1994), Laferre`re, M. (1968), Roux, P. et al. (1990), Skead, C.J. (1960).

Plate 35

Emberiza socotrana (Ogilvie-Grant and Forbes). Socotra Bunting. Bruant de Socotra.

(Opp. p. 538)

Fringillaria socotrana Ogilvie-Grant and Forbes, 1899. Bull. Liverpool Mus., 2, p. 2; Adho Dimellus, Socotra. Range and Status. Endemic resident, Socotra. Nowhere common; mainly at 500–1500 m in Hagghier (Hajhir) and Maggheer Mts. Records outside and presumptive breeding season from foothills south of Rookib and Qaysula (near Kallansiya). In 1903, 3 singing YY and a pair encountered in one week at 1200 m; in 1964 common enough for 17 birds to be collected in 15 weeks, though less common than Cinnamon-breasted Bunting E. tahapisi (which has slightly different habitat on Socotra: Ripley and Bond 1966). In 1993 survey of Socotra, only 2 birds found in 8 weeks (although

coverage of high altitude locations was rather limited: Morton 1966). In Nov 1993 and Nov 1997, several individuals and pairs and a flock of 5–6 between 550 and 900 m on Hagghier massif, with 2 singing YY only 200 m apart (Clouet et al. 1998). In Dec 2000, 2 single birds seen in a 3week survey (Jennings 2001). One of the scarcest endemic birds in Socotra, classified as Vulnerable (Collar et al. 1994). Description. ADULT Y: narrow central stripe from forehead to nape buffy white, streaked blackish brown. Broad lateral crown

580

EMBERIZIDAE

Food. Seeds of wild grasses, barley and other crops in spring and millet in autumn (NW Africa, W Sahara); spiders, ants and other small insects when breeding. Seeds, flowers and fruits of Salvadora persica, Ziziphus, Heliotropium, Phragmites, Phalaris and Bromus. Breeding Habits. Solitary nester; monogamous; territorial when breeding. Semi-colonial: in e.g. Marrakech, breeds numerously in some districts but absent from others, apparently suitable, only a km away. Pair remains together all year. Territories either well separated or adjacent; nests occasionally only 10–30 m apart. Nest or nest site often reused; one nest in Casablanca used 11–12 times in 5 years. 2–3 broods per season, occasionally 4; usually 3 in Marrakech, Morocco, from beginning of Mar to end of Aug (Roux et al. 1990); 2 around Djanet, Tassili, Algeria (Laferre`re 1968). Y defends territory by singing, from 1 preferred site (house wall or garden fence in Morocco, rock in E Sudan) and 2–3 subsidiary ones. Sings all year, but mainly in Nov–May (Morocco), in mornings, for several hours. Y pursues others in flight; they sometimes fight and fall to ground. On ground, rival YY face each other, give shrill calls, vibrate wings and move sideways. Y and X together vibrate wings and press body close to ground, perhaps a prelude to copulation, which takes place on ground (N. Chakir in Cramp and Perrins 1994). NEST: a small cup on a rough foundation of twigs, roots, straw and grass stems, lined with wool, hair, man-made materials and plant down. In Casablanca, Morocco, 16 out of 40 nests sited inside building on ledge, beam or in crevice, 10 on window-sill or balcony, 10 in hole in rockface or wall, and 4 in odd places including a mail-box and a fuse-box. Always in a sheltered place; usually 2–3 m above ground, once on ground under a bush; once on a palm trunk. In Tafilalet, 22 out of 31 nests in buildings or on sea cliffs, and 9 in clefts and crannies in rocks. SIZE (n ¼ 4, Algeria): ext. diam. 140–160 (150), int. diam. 70–80 (73), ext. height 50–60 (56), int. depth 15–20 (18). Built by X, Y helping at start (N. Chakir in Cramp and Perrins 1994); takes 4–7 days to restore old nest and 7–10 days or (another study) 20 days to build new one. EGGS: 1–5 (NW Africa, n ¼ 156 clutches, av. 313); 2–3 (Morocco, n ¼ 35 clutches, av. 269); also Morocco, n ¼ 84

clutches, av. 313 (Courteille and The´venot (1988). Laid daily, in morning. Sub-elliptical, smooth, faintly glossy; whitish, tinged blue or green, speckled with purplish brown forming a ring around large end, with faint undermarkings of purplish grey smears, or rufous-brown with mauve patches. SIZE: (sahari, n ¼ 14) 168–222  137– 146 (195  141), (n ¼ 44) 190–203  130–160; (striolata, n ¼ 15) 180–210  135–150 (193  142). LAYING DATES: Morocco, Feb–Oct, mainly late Feb to early June; Algeria, Mar (Ghardaı¨a), Oct (Djanet, Tassili, and nestlings Apr, Oct, Dec: Laferre`re 1968); Tunisia, Apr; Egypt, Jebel Abraq (fledglings Apr); Mali, (breeds throughout the year); Niger, June (Aguellal), Sept (Kori Tarare and Aoude´ras), (accompanied young, Agade`s, Feb); Sudan, Darfur, Feb, Oct, Nov; Eritrea, (gonads active, Jan). INCUBATION: by X alone; begins as soon as 1st egg laid. Period: 12–16 (13–14) days, or 14–16 days (Roux et al. 1990). In one nest all 3 eggs hatched on same day, but usually hatching is on consecutive days. DEVELOPMENT AND CARE OF YOUNG: nestlings fed by both parents, on regurgitated food. Data on feeding and growth rates in Courteille and The´venot (1988). Young brooded by X constantly for first 4 days and nights; X no longer sleeps in nest from 8th night. Eggshells and faecal sacs eaten by X; later, faecal sacs carried away and dropped. Nestling period: 17–19 days. In brood of 3, 2 leave nest on one day and the 3rd, the youngest, on next day. Young return to nest to sleep for first few nights. Out of nest, young fed by both parents, mainly Y, for 5–20 days (since X soon occupied with a new clutch) or 8 days (Roux et al. 1990). 2 young roosted with parents for >25 days after leaving nest. BREEDING SUCCESS/SURVIVAL: in Morocco (n ¼ 32 nests) 56% of 86 eggs hatched and 50% produced fledglings, also (n ¼ 58 nests) 66% of 169 eggs hatched and 57% gave rise to fledglings; mortality mainly due to human disturbance and cats. Also Morocco (n ¼ 174 clutches), hatching success 657%, rearing success 868%, overall fecundity 57% (Courteille and The´venot 1988). Key References Bannerman, D.A. (1948), Bannerman, D.A. and Priestley, J. (1952), Courteille, C. and The´venot, M. (1988), Cramp, S. and Perrins, C.M. (1994), Laferre`re, M. (1968), Roux, P. et al. (1990), Skead, C.J. (1960).

Plate 35

Emberiza socotrana (Ogilvie-Grant and Forbes). Socotra Bunting. Bruant de Socotra.

(Opp. p. 538)

Fringillaria socotrana Ogilvie-Grant and Forbes, 1899. Bull. Liverpool Mus., 2, p. 2; Adho Dimellus, Socotra. Range and Status. Endemic resident, Socotra. Nowhere common; mainly at 500–1500 m in Hagghier (Hajhir) and Maggheer Mts. Records outside and presumptive breeding season from foothills south of Rookib and Qaysula (near Kallansiya). In 1903, 3 singing YY and a pair encountered in one week at 1200 m; in 1964 common enough for 17 birds to be collected in 15 weeks, though less common than Cinnamon-breasted Bunting E. tahapisi (which has slightly different habitat on Socotra: Ripley and Bond 1966). In 1993 survey of Socotra, only 2 birds found in 8 weeks (although

coverage of high altitude locations was rather limited: Morton 1966). In Nov 1993 and Nov 1997, several individuals and pairs and a flock of 5–6 between 550 and 900 m on Hagghier massif, with 2 singing YY only 200 m apart (Clouet et al. 1998). In Dec 2000, 2 single birds seen in a 3week survey (Jennings 2001). One of the scarcest endemic birds in Socotra, classified as Vulnerable (Collar et al. 1994). Description. ADULT Y: narrow central stripe from forehead to nape buffy white, streaked blackish brown. Broad lateral crown

Emberiza socotrana

Emberiza socotrana

X and juv. Cinnamon-breasted Bunting also pale, but uniform light cinnamon below, with streaked throat, and no rufous in wing. Voice. Not tape-recorded. Song described as ringing and metallic whistled ‘hue-heee hu-hey’, recalling Socotra Golden-winged Grosbeak Rhynchostruthus socotranus (Ogilvie-Grant and Forbes 1903); call described by A.D. Forbes-Watson as a high thin whistle (sometimes repeated 2–3 times) followed by a soft gurgle, ‘tseep . . . guruguruguru’ (Ripley and Bond 1966).

stripes blackish brown; hindneck pale buff, streaked brown; mantle and back pale brown with narrow blackish streaks; scapulars cinnamon-rufous, streaked blackish; rump grey-buff, mottled blackish; uppertail-coverts buffy brown. Tail feathers dark brown, narrowly edged pale rufous, T6 with pale rufous outer web and pale buff tip. Broad white superciliary stripe, extending back to below nape; indistinct whitish orbital ring; black stripe through lores and behind eye links with black moustachial stripe, encircling buffy white patch on cheeks and ear-coverts; sub-moustachial stripe buffy white, bordered by faint dusky malar stripe; sides of neck grey. Chin buffy white, grading through pale cinnamon on throat to rufous-cinnamon on upper breast; flanks blotched with cinnamon; rest of underparts creamy, tinged cinnamon. Flight feathers dark brown, primaries narrowly edged pale buff, secondaries edged cinnamon-buff, P5–P8 emarginated; primary coverts and alula dark brown, fringed rich buff; tertials and greater coverts dark brown, broadly fringed rufous-cinnamon; median and lesser coverts rufouscinnamon. Underwing-coverts and axillaries buffy white. Bill dark brown above, orange-yellow or yellowish buff below with small dark tip; eyes dark brown; legs yellowish flesh. ADULT X: similar to ad. Y, but black head stripes tinged brown; cinnamon breast paler, less contrasting. SIZE: wing Y 68–735, X 635–69; tail, Y 555–605, X 50–585; bill to skull, Y 103–124, X 111– 117; tarsus, sexes combined 164–172 (n ¼ ?: Byers et al. 1995); also (1 Y, 1 X) wing, Y 68, X 68, tail, Y 48, X 54, bill Y 13, tarsus, Y 8, X 7. WEIGHT: 13–16 (Ripley and Bond 1966).

Field Characters. Length 13 cm. A stripe-headed bunting (A), distinguished from Cinnamon-breasted Bunting E. tahapisi (only other bunting on Socotra) by white underparts with rufous breast-band, solid rufous wingscoverts without pale fringes, and whitish band across lower back, conspicuous in flight, X similar but colours muted.

General Habits. Inhabits bleak and rugged high ground with granite and limestone peaks, boulders and steep cliffs, and dry, rocky hillsides with thickets, scrub and a few scattered small trees. Often on precipices. Once on a grassy, boulder-strewn plateau with scattered patches of low shrubs. Occasionally in thick scrub, foraging on the ground. Outside breeding season occurs in small flocks on lower ground, so may disperse or migrate altitudinally. 2 birds associated loosely with 15 Cinnamon-breasted Buntings (Kirwan et al. 1996); no association with other species noted in 15 weeks of observation (Ripley and Bond 1966). Forages on narrow rock-face ledges and stony slopes and in thickets; enters interstices and holes among rocks, probably searching for food. Perches on low vegetation rather more readily than does Cinnamon-breasted Bunting, and less prone to make lengthy escape flights when flushed. Sings from low shrub and in flight. Food. Very small grass seeds. Breeding Habits. Barely known; no nests found. LAYING DATES: (X with well-developed brood patch, Feb; sings in Nov and 1st half of Feb; judging from timing of moult, breeds in Dec–Apr). Key References Byers, C. et al. (1995), Clouet, M. et al. (1998), Collar, N.J. et al. (1994), Morton, K.M. (1996), Ogilvie-Grant, W.R. and Forbes, H.O. (1903), Ripley, S.D. and Bond, G.M. (1966).

581

582

EMBERIZIDAE

Plate 35

Emberiza capensis Linnaeus. Cape Bunting. Bruant du Cap.

(Opp. p. 538)

Emberiza capensis Linnaeus, 1766. Syst. Nat., ed. 12, 1, p. 310; Cape of Good Hope. Forms a superspecies with E. vincenti.

Emberiza capensis

Range and Status. Endemic resident and partial migrant, Africa south of 13
S. Angola, locally common from SW Namibe to Serra da Chela in Huı´la; record from near Benguela. Botswana, very uncommon resident from Tswapong Hills to Francistown. Mozambique, only in Chimanimani Massif, where occurs throughout, on border between Manica e Sofala and Zimbabwe. Namibia, Zimbabwe, South Africa and Lesotho, as mapped (Harrison et al. 1997); most abundant in W Cape on western escarpment and interior plateau of Karoo, S and E Cape Prov., and Lesotho. Up to 2200 m in Zimbabwe. In Transvaal, locally common, commonest in hilly parts of Highveld but scarce on Escarpment and in Lowveld. Lesotho, common in lowlands and the Maluti, and at all altitudes up to 2800 m; especially common in Sanqubeta Valley. Rivals Common Stonechat Saxicola torquata as being the commonest bird in Lesotho highlands. In KwaZulu-Natal common in Drakensberg, generally above 1200 m. Down to sea level in W Cape. Description. E. c. capensis Linnaeus: SW and W Cape Prov. to Namibia (north to about Tiraz Mts), and Gordonia, N Cape Prov.; intergrades with cinnamomea. ADULT Y: forehead to hindneck grey-brown, streaked blackish, most heavily along sides of forehead and crown. Rest of upperparts drab brown, mantle and scapulars sharply streaked blackish brown, back, rump and uppertail-coverts plain. Tail blackish brown, T6 with narrow whitish outer edge. Broad white superciliary stripe, extending back to side of nape. Broad black stripe through lores and behind eye; this joining broad black band behind ear-coverts and black moustachial stripe, to enclose buffy white patch on upper cheeks and lower ear-coverts. Chin and throat to sub-moustachial area buffy white; rest of underparts buffy grey, whiter on centre of belly. Flight feathers dark grey-brown, primaries edged buff, secondaries edged rufous; primary coverts and alula dark greybrown, fringed buff; tertials and greater coverts dark grey-brown, broadly edged and tipped rufous. Median and lesser coverts rufous. Underwing-coverts and axillaries greyish white; inner borders of undersides of flight feathers greyish white. Bill slate black; eyes brown; legs dark horn. ADULT X: like ad. Y, but black head stripes more often tinged brown. SIZE (10 YY, 9 XX): wing, Y 76–83 (787), X 71–79 (761); tail, Y 64–71 (673), X 60–67 (636); bill, Y 13–14 (135), X 125–145 (132); tarsus, Y 19–21 (202), X 195–21 (202). WEIGHT: (subspecies?) Y (n ¼ 3) 182– 236 (209), 1 X 225, unsexed (n ¼ 31) 17–24 (207). IMMATURE: juv. similar to ad., but more heavily streaked above; somewhat buffier below, with fine streaks on breast and flanks. NESTLING: at about a week, skin dark pink, dorsal down thick and dark grey, palate crimson or dark pink, gape yellow outside and inside, and mandible and maxilla edged yellow inside; outside, pale yellow of gape extends forward along side of mandible more than half way to its tip. E. c. cinnamomea (Lichtenstein): east of capensis in Cape Prov. (Karoo regions northeast to Lower and Middle Vaal R.), W Free State and adjacently in Transvaal. Warmer brown above than capensis; buffier below, especially on throat. E. c. reidi (Shelley): Lowland N Lesotho, E Free State, highland areas of KwaZulu-Natal, S and SE Transvaal, W Swaziland. Darker, more richly coloured above than capensis, rufous on wing

feathers more intense; deep olive-buff below. Larger: wing, Y (n ¼ 5) 84–87 (850). E. c. basutoensis (Vincent): highlands of Lesotho; seasonally in adjacent East Griqualand and KwaZulu-Natal. Darker brown above than capensis (but not warm brown like reidi); olive-grey below. E. c. limpopoensis (Roberts): central, W and SW Transvaal and SE Botswana. Like reidi but still richer rufous above. E. c. bradfieldi (Roberts): Namibia north of capensis. Slightly warmer brown above than capensis, and paler below. Bill longer: Y (n ¼ 5) 14–15 (147). E. c. nebularum (Rudebeck): SW Angola. Paler and greyer above than bradfieldi; paler and slightly greyer below. Bill longer than in bradfieldi. E. c. vinacea Clancey: N Cape (N Asbestos Mts and Kaap Plateau). Like capensis but underparts, including throat, vinaceous-buff. E. c. plowesi (Vincent): S Zimbabwe (Matobo Hills to Umvukwes and Inyanga Highlands) and NE Botswana. Lighter brown above than capensis, with bold sharp streaking; wing feather edging cinnamon-buff rather than rufous; paler and greyer below, like nebularum. WEIGHT: Y (n ¼ 1) 182. E. c. smithersii (Plowes): E Zimbabwe (Chimanimani Mts) and adjacent Mozambique. Dark olive-brown above, heavily streaked, with less rufous on secondary and tertial fringes; deep olive-grey below. Like basutoensis, but greyer above, and darker below, especially on throat.

Field Characters. Length 15–16 cm. The only bunting in its range combining striped face, unmarked white throat and rufous wings. Variable, from pale grey back and white underparts (SW Angola) to cinnamon-brown above and below (E South Africa); latter told from Cinnamonbreasted Bunting E. tahapisi by rufous wings and lack of black and white coronal stripes. Lark-like Bunting E. impetuani has some rufous in wings but plain, unstriped head.

Emberiza vincenti

583

Voice. Tape-recorded (72, 88, 99, B, F, PAY, WALK). Jerky song of 8–12 sharp chirping notes, lasting c. 3 s, typically beginning and ending with slower ‘chip, chop’, e.g. ‘chip, chop, chertsichee, chop, chertsichee, chop, chip, chop’; for other renditions see Skead (1960). Call a rolling, nasal ‘wer-wer-wi-wi’, rising in pitch. General Habits. Inhabits rocky, mountainous regions and sparse shrublands and grasslands on high mountains; rocky ridges and plains, around farmsteads and in village gardens (Karoo); rocky hills, open grassland and wooded habitats (Transvaal); on Korannaberg, Free State, breeds in rocky hillside bush habitats, and in non-breeding season occurs around farmyards and garden rockeries and in all habitats except forest; boulder-strewn slopes with short grass and scrub, on kranses and in gorges (KwaZuluNatal); escarpments and hillsides with rocky and stony ground, and rocky gorges and river canyons (Botswana); rugged quartzite and schist mountain ranges and granite shield region, confined to outcrops in kloofs and among boulders, overgrown with bushes, also exposed rocks on hillsides covered with Phillippia heath and montane grassland (Zimbabwe); grassy hillsides and in thick undergrowth (Mozambique Chimanimanis); bleak mountain summits in Lesotho, where at lower altitudes breeds within thick scrub. Generally near water. An unusual habitat is dry sandy soils near the sea in SW Western Cape Prov., with low, thin, sparse weeds and scrub, interspersed with coarse rushes (Vincent 1949). Generally in pairs; sometimes singly or in family parties. Forages on ground, progressing with short, shuffling walkhops. Flies low, for only short distances; on alighting often makes a quick about-turn. Usually unobtrusive; can be shy, but becomes tame in farmsteads and very tame at public picnic sites and campsites in W Cape, also in Lesotho. Sings from top of low bush or rock, sometimes opening and closing wings as it does so. Resident, not known as a migrant, but increase in reporting rates in winter in SW Western Cape Prov. suggest that there is an influx of birds then (Harrison et al. 1997).

A

Food. Mainly seeds; in captivity, eats seeds of grasses Chloris virgata, Eragrostis capensis, E. racemosa, E. bicolor, E. curvula, E. tef, Panicum maximum and P. laevifolium (Brickell 1986); arils of Acacia cyclops. Some insects, including grasshoppers, beetles and larvae, fly larvae and termites; also spiders.

courting another with their tails fanned, the third quivering wings and spreading tail (Skead 1960). NEST: neat cup made of fine rootlets, grass, hair and woolly plant down, on untidy foundation of twigs, roots and grass (A). Sited c. 10 cm above ground or sometimes on it, against a plant or single-stemmed stunted bush on the sloping bank of sheltered erosion gully or quarry; sometimes c. 25 cm up in creeper or on fork in bush, or cradled among twiglets near top of bush; often by rock surface; also in thick tuft of grass in dense scrub, in driftwood caught up in tree roots, and bushy vegetation clinging to cliff-face, up to 10 m from foot. Mostly open to view, sometimes partly screened. Int. diam. av. 62, int. depth av. 57. EGGS: 2–5, usually 2–3, av. (n ¼ 23 clutches) 26. Ground colour white or cream, pale blue or pale green; speckled and heavily blotched with reddish brown, fawn, grey and purple, mainly at broad end where can form a ring. SIZE: (n ¼ 62) 179–232  136–164 (203  150). LAYING DATES: W Cape, July–Jan (mainly Sept–Oct); E Cape, Nov–Apr; Free State, Oct–Mar, (young under parental care late Sept to Jan); Transvaal, Oct–Nov; Zimbabwe, Nov–June (mainly Dec–Apr). INCUBATION: no information. DEVELOPMENT AND CARE OF YOUNG: young fed by both parents. No further information. BREEDING SUCCESS/SURVIVAL: occasionally parasitized by Diederik Cuckoo Chrysococcyx caprius (R. Martin in Ginn et al. 1989).

Breeding Habits. Solitary nester; monogamous; territorial. Evident courtship behaviour includes flight from treetop to treetop with slow, deliberate wingbeats; 2 singing birds

Key References Byers, C. et al. (1995), Ginn, P.J. et al. (1989), Harrison, J.A. et al. (1997), Irwin, M.P.S. (1981), Maclean, G.L. (1993), Skead, C.J. (1960), Vincent, A.W. (1949).

Emberiza vincenti (Lowe). Vincent’s Bunting. Bruant de Vincent.

Plate 35

Fringillaria capensis vincenti Lowe, 1932. Bull. Br. Orn. Club, 52, p. 144; Zobrue´ (Zobue´), Mozambique.

(Opp. p. 538)

Forms a superspecies with E. capensis. ˆ i. Range and Status. Endemic resident in and near Malaw ˆ i, S Mzimba District (Pembe Hill, S Viphya Mts) Malaw

south to Ntchisi (Chipata Mts), Dedza (Chongoni Mts), Cape Maclear Peninsula, Mangochi Mts, W Mwanza and

Emberiza vincenti

583

Voice. Tape-recorded (72, 88, 99, B, F, PAY, WALK). Jerky song of 8–12 sharp chirping notes, lasting c. 3 s, typically beginning and ending with slower ‘chip, chop’, e.g. ‘chip, chop, chertsichee, chop, chertsichee, chop, chip, chop’; for other renditions see Skead (1960). Call a rolling, nasal ‘wer-wer-wi-wi’, rising in pitch. General Habits. Inhabits rocky, mountainous regions and sparse shrublands and grasslands on high mountains; rocky ridges and plains, around farmsteads and in village gardens (Karoo); rocky hills, open grassland and wooded habitats (Transvaal); on Korannaberg, Free State, breeds in rocky hillside bush habitats, and in non-breeding season occurs around farmyards and garden rockeries and in all habitats except forest; boulder-strewn slopes with short grass and scrub, on kranses and in gorges (KwaZuluNatal); escarpments and hillsides with rocky and stony ground, and rocky gorges and river canyons (Botswana); rugged quartzite and schist mountain ranges and granite shield region, confined to outcrops in kloofs and among boulders, overgrown with bushes, also exposed rocks on hillsides covered with Phillippia heath and montane grassland (Zimbabwe); grassy hillsides and in thick undergrowth (Mozambique Chimanimanis); bleak mountain summits in Lesotho, where at lower altitudes breeds within thick scrub. Generally near water. An unusual habitat is dry sandy soils near the sea in SW Western Cape Prov., with low, thin, sparse weeds and scrub, interspersed with coarse rushes (Vincent 1949). Generally in pairs; sometimes singly or in family parties. Forages on ground, progressing with short, shuffling walkhops. Flies low, for only short distances; on alighting often makes a quick about-turn. Usually unobtrusive; can be shy, but becomes tame in farmsteads and very tame at public picnic sites and campsites in W Cape, also in Lesotho. Sings from top of low bush or rock, sometimes opening and closing wings as it does so. Resident, not known as a migrant, but increase in reporting rates in winter in SW Western Cape Prov. suggest that there is an influx of birds then (Harrison et al. 1997).

A

Food. Mainly seeds; in captivity, eats seeds of grasses Chloris virgata, Eragrostis capensis, E. racemosa, E. bicolor, E. curvula, E. tef, Panicum maximum and P. laevifolium (Brickell 1986); arils of Acacia cyclops. Some insects, including grasshoppers, beetles and larvae, fly larvae and termites; also spiders.

courting another with their tails fanned, the third quivering wings and spreading tail (Skead 1960). NEST: neat cup made of fine rootlets, grass, hair and woolly plant down, on untidy foundation of twigs, roots and grass (A). Sited c. 10 cm above ground or sometimes on it, against a plant or single-stemmed stunted bush on the sloping bank of sheltered erosion gully or quarry; sometimes c. 25 cm up in creeper or on fork in bush, or cradled among twiglets near top of bush; often by rock surface; also in thick tuft of grass in dense scrub, in driftwood caught up in tree roots, and bushy vegetation clinging to cliff-face, up to 10 m from foot. Mostly open to view, sometimes partly screened. Int. diam. av. 62, int. depth av. 57. EGGS: 2–5, usually 2–3, av. (n ¼ 23 clutches) 26. Ground colour white or cream, pale blue or pale green; speckled and heavily blotched with reddish brown, fawn, grey and purple, mainly at broad end where can form a ring. SIZE: (n ¼ 62) 179–232  136–164 (203  150). LAYING DATES: W Cape, July–Jan (mainly Sept–Oct); E Cape, Nov–Apr; Free State, Oct–Mar, (young under parental care late Sept to Jan); Transvaal, Oct–Nov; Zimbabwe, Nov–June (mainly Dec–Apr). INCUBATION: no information. DEVELOPMENT AND CARE OF YOUNG: young fed by both parents. No further information. BREEDING SUCCESS/SURVIVAL: occasionally parasitized by Diederik Cuckoo Chrysococcyx caprius (R. Martin in Ginn et al. 1989).

Breeding Habits. Solitary nester; monogamous; territorial. Evident courtship behaviour includes flight from treetop to treetop with slow, deliberate wingbeats; 2 singing birds

Key References Byers, C. et al. (1995), Ginn, P.J. et al. (1989), Harrison, J.A. et al. (1997), Irwin, M.P.S. (1981), Maclean, G.L. (1993), Skead, C.J. (1960), Vincent, A.W. (1949).

Emberiza vincenti (Lowe). Vincent’s Bunting. Bruant de Vincent.

Plate 35

Fringillaria capensis vincenti Lowe, 1932. Bull. Br. Orn. Club, 52, p. 144; Zobrue´ (Zobue´), Mozambique.

(Opp. p. 538)

Forms a superspecies with E. capensis. ˆ i. Range and Status. Endemic resident in and near Malaw ˆ i, S Mzimba District (Pembe Hill, S Viphya Mts) Malaw

south to Ntchisi (Chipata Mts), Dedza (Chongoni Mts), Cape Maclear Peninsula, Mangochi Mts, W Mwanza and

584

EMBERIZIDAE

Emberiza vincenti

(654), X 62–70 (641); bill, Y 12–135 (131), X 125–14 (131); tarsus, Y 18–19 (184), X 18–195 (186). IMMATURE: juv. differs from ad. in having pale head stripes, chin and throat pale buff, black head bands tinged brown; upperparts rather browner, underparts paler, more buffy grey. TAXONOMIC NOTE: J. Vincent wrote in Ibis, 1939, p. 121: ‘‘There is a strongly marked discrepancy in size in the entire series of smaller birds from Mirrote. My firm conviction is that time will prove the birds of the type-locality [Zobrue´] to be specifically distinct from the far distant F[ringillaria] capensis of South Africa, and that the Mirrote birds will be subspecifically separable from those west of the Shire´ Valley’’. The Mirrote birds are indeed different from capensis, in our view specifically so, but we can find no reason to separate them subspecifically from the remainder of E. vincenti (and nor could Benson 1941). Differences in appearance between E. vincenti and E. capensis are much greater than those between the many races of E. capensis sens. str. and are at least as great as those within other bunting pairs (e.g. E. striolata/E. tahapisi, E. socotrana/E. capensis, E. buchanani/ E. caesia). The habitat of E. vincenti is distinctive and its song appears to be too.

Field Characters. Length 155 cm. The only bunting in its range combining striped face, unmarked pale throat and rufous wings; differs from Cinnamon-breasted Bunting E. tahapisi in pale throat, uniform dark grey underparts and longer tail. Mulanje; mainly at about 915–1830 m, but down to 500 m on Cape Maclear (where one seen in 1957, perhaps only a vagrant). Zambia, only in Eastern Prov., on land above 1000 m from Nyanje to Chadiza. Tanzania, shown in regions of Mbinga and Songea highlands in extreme SW ˆ i border (Byers et al. 1995). Mozambique, Zobue´ (on Malaw west of Blantyre); records in Mangochi Mts on Mozambique ˆ i side of border (Hall and Moreau side as well as Malaw 1970); numerous at Mirrote (13
500 S, 39
340 E, near Lurio R.) at 275 m; searched for unsuccessfully in likely ˆ i and Mirrote habitat in Niassa Prov. between Malaw (Vincent 1936). Description. ADULT Y: stripe down centre of crown to hindneck dark grey-brown with a few fine black streaks; broad band from side of forehead to side of nape black, with dark grey-brown streaking medially on crown and nape sides. Rest of upperparts dark grey-brown, with poorly defined darker streaks on mantle, back and scapulars. Tail blackish brown. Broad white superciliary stripe, extending back to side of nape; broad black stripe through lores and broad black band through upper ear-coverts, joining black band behind ear-coverts, continuous with broad black band across lower cheeks and black moustachial stripe, enclosing broad greyish white band from base of bill to cheeks and lower earcoverts. Chin, upper throat and sub-moustachial area greyish white; side of neck, lower throat, breast and flanks brownish grey, becoming paler on centre of belly; undertail-coverts brownish grey, tipped greyish white. Remiges dark grey-brown, secondaries and tertials with narrow tawny-brown edges when fresh; primary coverts and alula dark grey-brown with fine tawny fringes; greater coverts dark grey-brown, edged and broadly tipped deep rufous; median and lesser coverts deep rufous. Underwingcoverts and axillaries ashy grey; inner borders of undersides of flight feathers pale grey. Upper mandible slate black, lower mandible pale grey or bluish grey with blackish tip; eyes dark brown; legs dark grey or dark brownish grey. Sexes alike. SIZE (10 YY, 8 XX): wing, Y 75–82 (782), X 72–80 (750); tail, Y 62–70

Voice. Song a ‘shrill, jolly, syncopated, canary-like whistle, ‘tree-re-reet, tree-re-reet’, the second syllable the highest’ and call ‘a small, almost sparrow-like chirp’ (Vincent 1936). General Habits. Inhabits bare, rocky mountain summits (Mt Kambironjo and others in SW Ntcheu District at ˆ i), granite cliff-faces and their wooded 1000–1500 m, Malaw bases where there are large, tumbled boulders (Zambia); in Mozambique, almost bare rocky mountain slopes studded with tree-lilies (Velloziaceae), with a few cacti, aloes, stunted bushes and dry tufts of grass; also ‘a large flat rock just below the mountain summit, beneath woodland trees where big boulders poked up everywhere out of long grass’ (Vincent 1936). Forages on ground; gait, stance and movements like congeners; bird is larger and plumper than Cinnamon-breasted Bunting E. tahapisi, takes to wing more readily and flies further. Y sings from top of rock or bush. Food. Not known. Breeding Habits. Solitary nester. NEST: not described. EGGS: 3 (1 clutch). ˆ i, Apr (and gonads active Apr– LAYING DATES: Malaw June); Zambia, (gonads active Mar); Mozambique, (Y had bred in Apr; many accompanied juvs in June). Key References Benson, C.W. (1941)., Benson, C.W. and Benson, F.M. (1977), Vincent, J. (1936).

Emberiza impetuani

585

Emberiza impetuani A. Smith. Lark-like Bunting. Bruant des rochers.

Plate 35

Emberiza impetuani A. Smith, 1826. Report Exped. Central Africa, p. 48; ‘‘country between Nu. Gariep and Tropic’’, restricted to Bechuanaland, Macdonald, 1957, Contrib. Orn. Western South Africa, p. 169.

(Opp. p. 538)

Range and Status. Endemic resident and irruptive nomad, SW Africa; probably one of the most abundant birds in SW Arid Zone. Breeding range mapped is notional in parts: eastern breeding limits in Namibia and northern ones in Northern Cape Prov. not well known. Angola, Luanda and coastal plain south to SW Namibe; and isolated population at L. Cameia in Moxico. Namibia, common except in NE. Moves in relation to Karoo rainfall; enters W Namaqualand in Sept–Dec and Namib Desert and whole of S Namibia in Feb–Apr. In South Africa common and widespread west of 26
E but absent from coast east of Cape Peninsula. In Transvaal, commonest west of Wolmaransstad and Ottosdal, appearing sporadically in flocks of hundreds, and in many records in central bushveld and N and central lowveld, particularly in 1984; common in Langjan Nat. Res. in Dec 1979 but rare there by next Aug; only 1 Transvaal breeding record (Wyliespoort). In Free State, fairly common to locally abundant nomad west of 26
E; in dry years occurs north to Vaal R. and east to Heilbron; an outlying population resident in Korannaberg area (max. 1890 m, west of Clocolan), abundant in breeding season (Farkas 1988). In Botswana, widespread south of 21
S, sparse to fairly common but occurrence unpredictable and sporadic; common in Nossob valley, irrupts into N woodlands in some years, e.g. 1992, a drought year (Brewster 1994), also 1981, 1987 and 1988; in E, an influx into Gaborone area in 1998, when several thousand birds around Ruretse and nearly 1900 at a waterhole at a time (Tyler and Tyler 2001); none in E in 2000 (Tyler and Brewster 2000); not known to breed in Botswana. In Zimbabwe, makes irruptive or nomadic movements, entering SW in unusually cold or dry winters; major incursions in 1958 and 1964 in late May and June. The 1958 invasion thought to involve millions of birds over large part of NW Matabeleland. In 1964, reached Harare and N bank of Kariba in Nov. Present in Limpopo Valley from Ramaquabane R. to Tuli Circle and Sentinel Ranch; records east to Bubye R. where has bred and may occur regularly (Irwin 1981). Vagrant in Zambia (4 records), SW Zaı¨re (2), Lesotho, Natal and E Cape Prov. coast (1 each). Description. E. i. sloggetti (Macdonald): W, S and central Cape Prov. to W Free State, Griqualand West and adjacent S Transvaal. ADULT Y: top of head and upperparts buffy brown, streaked dark brown, streaks sharp and narrow on head, more diffuse on rump and uppertail-coverts. Tail blackish brown, both webs of T1 and outer web of T2–T6 fringed buff, the fringe on T6 broad and extending around tip. Broad buff superciliary stripe. Lores, cheeks and ear-coverts buffy brown, with slightly darker stripe behind eye and darker moustachial stripe; sub-moustachial stripe buff, bordered faintly below by narrow blackish malar streak; sides of neck buffy brown. Underparts pale buff, greyer on chin and throat, whiter on belly, tinged cinnamon on breast and flanks. Remiges blackish brown, primaries narrowly edged buff, secondaries and tertials edged rufous-buff; primary coverts, alula, and greater, median and lesser coverts dark brown, all broadly fringed rufousbuff. Underwing-coverts and axillaries pale cinnamon-buff.

Emberiza impetuani

Undersides of inner borders of flight feathers cinnamon-buff. Upper mandible dark brown, lower mandible pale brown; eyes blackish brown; feet pinkish brown. ADULT X: like ad. Y, but breast often rather paler (contrasting slightly less with belly) and with faint streaks on sides. SIZE (10 YY, 10 XX): wing, Y 74–80 (784), X 71–77 (741); tail, Y 58–64 (606), X 55–61 (582), bill, Y 11–125 (121), X 11–125 (116); tarsus, 175–19 (182), X 175– 185 (181). Also, wing, Y (n ¼ 10) 77–805 (785), X (n ¼ 5) 75– 775 (757) (Clancey 1989). WEIGHT: Y (n ¼ 8) 138–169 (151), X (n ¼ 7) 133–172 (144), unsexed (n ¼ 31) 137–166 (151). IMMATURE: juv. similar to ad., but less sharply streaked above; breast slightly browner (less cinnamon), faintly streaked or spotted; upperwing-coverts more uniform dull rufous, dark centres less well defined. E. i. impetuani A. Smith: N and NE Cape Prov., W, S and E Botswana and adjacent Namibia, W Transvaal and S Zimbabwe. Slightly more tawny above than sloggetti, more cinnamon-buff below, light mid-ventral area more restricted. Smaller, sexes similar in wing length: wing, Y (n ¼ 8) 73–75 (739), X (n ¼ 12) 70–75 (732) (Clancey 1989); unsexed (n ¼ 325) 70–83 (767). WEIGHT: unsexed (n ¼ 330, SE Botswana) 102–200 (149), av. c. 08 g heavier in Apr than Nov. (Tyler and Tyler 2001). E. i. eremica Clancey: NW Cape Prov., S, central and NW Namibia and SW Angola. Slightly more tawny above than sloggetti, with heavier and blacker streaking; more deeply coloured below with pale mid-ventral area restricted as in impetuani. Size as sloggetti: wing, Y (n ¼ 19) 75–80 (773), X (n ¼ 15) 72–78 (753) (Clancey 1989).

Field Characters. Length 125–14 cm. Looks like a lark, acts like a bunting. Almost featureless, sandy brown with paler supercilium and submoustachial stripe and indistinct brown face patch, and rufous wing-edgings, forming panel

586

EMBERIZIDAE

on folded secondaries. Juv. Cinnamon-breasted Bunting E. tahapisi has distinctly striped face, streaked throat and pale cinnamon underparts. Told from larks by bunting shape (short conical bill, small head, long tail), no white in tail or streaks on breast, and uniform appearance (in most larks upperparts are darker than underparts). Voice. Tape-recorded (38, 88, 99, B, F, ROC). Song hurried and tuneless, lasting 1–2 s, a mixture of harsh chirps and dry rattles, often ending with high-pitched metallic trill reminiscent of Corn Bunting Miliaria calandra. Call a hard, nasal ‘jaa’ or ‘jeea’; sharp ‘chut’ when flushed. General Habits. Inhabits arid and semi-arid savanna, rocky hills with sparse grass and shrubs, sparse perennial grassland on schists and tilted shales (Harrison et al. 1997), sparse woodland, sparse bush along dry drainage lines, heavily grazed rangelands and sparsely vegetated ground in general: dunes, dune grassland, dune troughs, escarpments in dry river valleys, rock or stone outcrops, stony hillsides (Penry 1994); in Botswana, often around water at cattle stations, boreholes and human habitations in arid areas; in Korannaberg area, Free State, breeds in karroid habitats transitional to rocky and grassveld types and wanders throughout all habitat types except forest; often around farmyards in winter. Occurs in pairs, family parties, small, medium and large flocks; often flocks of 5–15, occasionally 50–100, rarely up to thousands of birds (mainly at water). Occurs in mixed flocks with other buntings and Black-headed Canaries Alario alario and forages close to flocking Grey-backed and Black-eared Sparrow-Larks Eremopterix verticalis and E. australis (W.R.J. Dean in Harrison et al. 1997). Forages on ground, walking and running like a lark (although Bunting is shorter-legged than most larks); jumps up to take seeds from grass heads. Restless and somewhat wary, not permitting close approach by a person, and easily disturbed. When approached, crouches down and looks alert but flies reluctantly, and then not distantly; similarly, a flock put to flight soon settles on ground, rocks or bush tops. Flight finch-like, jinking and undulating. Drinks regularly, from ground puddles, pools and dams, and often occurs within a few km of water. In Ruretse, Botswana, commonest at waterhole in July, at 08h00 and in morning, less common in afternoon, uncommon in evening (Tyler and Tyler 2001). Sings from low eminence: top of small bush or rock; also sings from flat ground and top of a bank. Food. Seeds; in rains, some green grass shoots and germinating seeds (Byers et al. 1995); some insects, including small caterpillars. In captivity, takes seeds of Chloris virgata, Rhynchelytrum repens, Panicum maximum, Eragrostis curvula, Paspalum sistichum, Setaria pallidefusca, and mealworms, caterpillars, thrips nymphs, termites, gnats, millipedes and spiders (Brickell and Konigkramer 1997). Breeding Habits. Solitary nester; presumably monogamous. Courtship not described. During nest-building, pair often flies in and settles on ground, then Y flies up onto low bush to sing. Often double-brooded.

NEST: loosely-built foundation of twigs (some quite thick and heavy) and short woody plant stems, with neat, compact, shallow cup upon or within it, made of finest yellowish grass and sometimes fine rootlets. Placed on ground on stony slope or, less frequently, on level stony or sandy ground, in depression amongst sparse growth of weeds and stunted shrubs, in cavity partly under rock or at side of stone, or under bush, generally on SE side; sometimes in lee of tuft of grass or on completely bare ground (A). When nest built against plant stem or low earthen ridge, outer structure is often no more than a loose collection of twigs piled against grass cup on its open side (Vincent 1949). Int. diam. (n ¼ 8) 53–67 (58), depth of cup 24–39 (28). Built by X, Y keeping nearby and accompanying her on collecting trips. Visits to nest under construction are at rather long intervals. Material often gathered some distance away, birds flying in swiftly low over scrub. Near Cape Town, nest-building on upper slopes of low hills starts in early Oct, eggs being laid from late Oct to mid Nov. EGGS: 2–4, usually 3–4 (Cape Town), av. 29. White, palest greenish or pale bluish, finely freckled all over, more densely at large end, and sometimes blotched, with violetgrey, rust and brown. SIZE: (n ¼ 165) 155–196  121–153 (178  132) (Maclean 1993). LAYING DATES: breeds opportunistically and irregularly, in response to rainfall, mainly in spring in winter rainfall regions and in late summer and autumn in late-summer rainfall regions; breeds probably bimodally in Northern Cape Prov.; Namibia, Mar–May; W Cape, Sept–Nov;

Emberiza cia Karoo, Sept–Oct and Mar–May; SW Kalahari, Feb–Mar; Transvaal, (nest-building, July); Zimbabwe, Mar–May. INCUBATION: by X. Period: 13 days. DEVELOPMENT AND CARE OF YOUNG: nestling period c. 12–13 days.

587

BREEDING SUCCESS/SURVIVAL: particularly prone to desert nest before completion (Vincent 1949).

Key References Byers, C. et al. (1995), Clancey, P.A. (1989), Harrison, J.A. et al. (1997), Maclean, G.L. (1993), Skead, C.J. (1960), Tyler, J. and Tyler, L. (2001), Vincent, A.W. (1949).

Emberiza cia Linnaeus. Rock Bunting. Bruant fou.

Plate 35

Emberiza cia Linnaeus, 1766. Syst. Nat., ed. 12, p. 310; Lower Austria.

(Opp. p. 538)

Range and Status. Resident, central and S Europe, NW Africa, Turkey to Caucasus and Iran, and central Asia south to Himalayas. Breeds in hills and mountains in N and W Morocco, N Algeria and N Tunisia; sedentary or dispersive; partly leaves high ground in winter. Morocco, Djebala, Rif and NE highlands to Moyen and Haut Atlas; and W slopes of Anti-Atlas, toward Anja and near Abeino; locally common; Algeria, coast to Atlas Tellien, Aure`s Mts, Atlas Saharien; locally common, but scarce in Oranie, absent from Mitidja and apparently from Hauts Plateaux. Tunisia, near massifs from Tunis south to Gafsa, uncommon. Breeds mainly at 600–1500 m, but up to 2000 m in Algeria, and at 2200– 2300 m (exceptionally above 2700 m) in Moroccan Atlas. In winter, some occur in non-breeding areas and descend to plains; reaches borders of Algerian Sahara south of Monts des Ksours and Biskra (once Taggourt). Vagrant NW Libya (once, Jefren escarpment, Feb 1978) and N Sudan (Wadi Halfa, twice, Mar 1972). (A bird caught at Abe´che´, Chad (Salvan 1963) and claimed as this species can be discounted from its wing measurement.) Breeding density in maquis, Morocco, 5–34 pairs per km2 (av. 98) (The´venot 1982). Description. E. c. cia Linnaeus: only race in Africa; includes ‘africana’: Europe and N Africa east to W Caucasus. ADULT Y: centre of crown to hindneck grey, finely streaked black; broad stripe at side of crown and nape black, flecked with grey; mantle, back and scapulars cinnamon, streaked black, broadly and boldly on lower mantle and scapulars; rump bright rufous-cinnamon; uppertail-coverts cinnamon with black shaft streaks. Tail blackish brown; both webs of T1 fringed cinnamon-buff; inner web of T5–T6 with white distal wedge, extending along shaft to twothirds of length of T6, half length of T5; T6 also with narrow white outer edge. Broad superciliary stripe greyish white. Lores black; stripe behind eye black, extending to rear of ear-coverts; cheeks and ear-coverts grey, paler near bill base, enclosed by broad black malar stripe linked to black bar behind ear-coverts; sides of neck grey. Chin to upper breast grey, usually paler near bill base; rest of underparts cinnamon-rufous, grading to pale cinnamon-buff on centre of belly and thighs. Upperwing feathers dark grey-brown; primaries, primary coverts and small alula feather finely edged cinnamon; secondaries edged cinnamon and tertials fringed more broadly so; greater coverts broadly edged cinnamon and tipped whitish on outer webs; median coverts tipped whitish; lesser coverts grey. Axillaries and most underwingcoverts white; marginal underwing-coverts mottled dark grey. Upper mandible dark horn-grey or blue-black; lower mandible light grey or blue-grey, tinged yellow or pink at base; eyes dark brown to red-brown; legs pale flesh-brown or red-brown. ADULT X: similar to ad. Y, but grey of crown, supercilium and

Emberiza cia

ear-coverts tinged brown; black head stripes mixed with brown; central crown streaks broader; grey of chin to upper breast tinged buff, breast feathers with a few dark brown triangular spots when fresh. Complete moult July–Oct. SIZE (10 YY, 10 XX): wing, Y 79–87 (828), X 74–79 (766); tail, Y 73–85 (789), X 68–75 (715); bill, Y 135–145 (140), X 135–145 (140); tarsus, Y 195–21 (202), X 185–205 (196). WEIGHT: (Germany, Apr–June) Y (n ¼ 51) 21–29 (242), X (n ¼ 38) 20–27 (227); (Italy, Aug–Nov) unsexed (n ¼ 97) 19–28 (222). IMMATURE: juv. buffy brown above, heavily streaked dark brown, including rump; top of head paler, sandier, well streaked. Buff supercilium poorly marked; ear-coverts brown with pale buff central area; lower cheeks and malar area buffy white, flecked dark brown. Chin and throat off white, throat flecked dark brown; rest of underparts pale buff, with band of short dark brown streaks across breast and narrower streaks on flanks. Tail and wing as ad. but greater, median and lesser coverts sharply fringed pale buff. NESTLING: down dark grey, long, plentiful, restricted to upperparts, upperwing, vent and thigh; mouth uniform yellowpink, gape-flanges whitish.

Field Characters. Length 16 cm. Y easily recognized by silver-grey head with black stripes; body rich rufous. When flushed from side of road, rusty red rump and whiteedged tail characteristic (Jonsson 1992). X duller, but head

Emberiza cia Karoo, Sept–Oct and Mar–May; SW Kalahari, Feb–Mar; Transvaal, (nest-building, July); Zimbabwe, Mar–May. INCUBATION: by X. Period: 13 days. DEVELOPMENT AND CARE OF YOUNG: nestling period c. 12–13 days.

587

BREEDING SUCCESS/SURVIVAL: particularly prone to desert nest before completion (Vincent 1949).

Key References Byers, C. et al. (1995), Clancey, P.A. (1989), Harrison, J.A. et al. (1997), Maclean, G.L. (1993), Skead, C.J. (1960), Tyler, J. and Tyler, L. (2001), Vincent, A.W. (1949).

Emberiza cia Linnaeus. Rock Bunting. Bruant fou.

Plate 35

Emberiza cia Linnaeus, 1766. Syst. Nat., ed. 12, p. 310; Lower Austria.

(Opp. p. 538)

Range and Status. Resident, central and S Europe, NW Africa, Turkey to Caucasus and Iran, and central Asia south to Himalayas. Breeds in hills and mountains in N and W Morocco, N Algeria and N Tunisia; sedentary or dispersive; partly leaves high ground in winter. Morocco, Djebala, Rif and NE highlands to Moyen and Haut Atlas; and W slopes of Anti-Atlas, toward Anja and near Abeino; locally common; Algeria, coast to Atlas Tellien, Aure`s Mts, Atlas Saharien; locally common, but scarce in Oranie, absent from Mitidja and apparently from Hauts Plateaux. Tunisia, near massifs from Tunis south to Gafsa, uncommon. Breeds mainly at 600–1500 m, but up to 2000 m in Algeria, and at 2200– 2300 m (exceptionally above 2700 m) in Moroccan Atlas. In winter, some occur in non-breeding areas and descend to plains; reaches borders of Algerian Sahara south of Monts des Ksours and Biskra (once Taggourt). Vagrant NW Libya (once, Jefren escarpment, Feb 1978) and N Sudan (Wadi Halfa, twice, Mar 1972). (A bird caught at Abe´che´, Chad (Salvan 1963) and claimed as this species can be discounted from its wing measurement.) Breeding density in maquis, Morocco, 5–34 pairs per km2 (av. 98) (The´venot 1982). Description. E. c. cia Linnaeus: only race in Africa; includes ‘africana’: Europe and N Africa east to W Caucasus. ADULT Y: centre of crown to hindneck grey, finely streaked black; broad stripe at side of crown and nape black, flecked with grey; mantle, back and scapulars cinnamon, streaked black, broadly and boldly on lower mantle and scapulars; rump bright rufous-cinnamon; uppertail-coverts cinnamon with black shaft streaks. Tail blackish brown; both webs of T1 fringed cinnamon-buff; inner web of T5–T6 with white distal wedge, extending along shaft to twothirds of length of T6, half length of T5; T6 also with narrow white outer edge. Broad superciliary stripe greyish white. Lores black; stripe behind eye black, extending to rear of ear-coverts; cheeks and ear-coverts grey, paler near bill base, enclosed by broad black malar stripe linked to black bar behind ear-coverts; sides of neck grey. Chin to upper breast grey, usually paler near bill base; rest of underparts cinnamon-rufous, grading to pale cinnamon-buff on centre of belly and thighs. Upperwing feathers dark grey-brown; primaries, primary coverts and small alula feather finely edged cinnamon; secondaries edged cinnamon and tertials fringed more broadly so; greater coverts broadly edged cinnamon and tipped whitish on outer webs; median coverts tipped whitish; lesser coverts grey. Axillaries and most underwingcoverts white; marginal underwing-coverts mottled dark grey. Upper mandible dark horn-grey or blue-black; lower mandible light grey or blue-grey, tinged yellow or pink at base; eyes dark brown to red-brown; legs pale flesh-brown or red-brown. ADULT X: similar to ad. Y, but grey of crown, supercilium and

Emberiza cia

ear-coverts tinged brown; black head stripes mixed with brown; central crown streaks broader; grey of chin to upper breast tinged buff, breast feathers with a few dark brown triangular spots when fresh. Complete moult July–Oct. SIZE (10 YY, 10 XX): wing, Y 79–87 (828), X 74–79 (766); tail, Y 73–85 (789), X 68–75 (715); bill, Y 135–145 (140), X 135–145 (140); tarsus, Y 195–21 (202), X 185–205 (196). WEIGHT: (Germany, Apr–June) Y (n ¼ 51) 21–29 (242), X (n ¼ 38) 20–27 (227); (Italy, Aug–Nov) unsexed (n ¼ 97) 19–28 (222). IMMATURE: juv. buffy brown above, heavily streaked dark brown, including rump; top of head paler, sandier, well streaked. Buff supercilium poorly marked; ear-coverts brown with pale buff central area; lower cheeks and malar area buffy white, flecked dark brown. Chin and throat off white, throat flecked dark brown; rest of underparts pale buff, with band of short dark brown streaks across breast and narrower streaks on flanks. Tail and wing as ad. but greater, median and lesser coverts sharply fringed pale buff. NESTLING: down dark grey, long, plentiful, restricted to upperparts, upperwing, vent and thigh; mouth uniform yellowpink, gape-flanges whitish.

Field Characters. Length 16 cm. Y easily recognized by silver-grey head with black stripes; body rich rufous. When flushed from side of road, rusty red rump and whiteedged tail characteristic (Jonsson 1992). X duller, but head

588

EMBERIZIDAE

stripes still distinctive. In N Africa can only be confused with smaller House Bunting E. striolata, which has bright rufous wings, finely streaked head and breast, no white in tail. Juv. heavily streaked on head and breast, unlike juv. House Bunting, and has white in tail, striped wings with little rufous. Voice. Tape-recorded (62–73, 92, 105, 110–113, B). Extremely variable song thin and high-pitched but not unmusical, a medley of chips, short trills and slurred notes, with jerky quality reminiscent of Dunnock Prunella modularis, ‘trrr, see-so, chichi siwirr chit’, or ‘chyo titi tsiker ziwo sisisisi chewy’; rather fast and long for a bunting (up to 3–4 s). Common contact call a sharp ‘tsee’ or ‘tzit’; other calls include thin, descending ‘seeee’ or ‘seea’; short ‘tucc’, repeated to become twitter in flight; on rising, ‘chelut’, ‘chit vit’ or slightly vibrant ‘chee dee dee’ (Jonsson 1992). For sonagrams and further details see Cramp and Perrins (1994). General Habits. Inhabits high, open, semi-arid areas with sparse shrub vegetation and few trees; dry stony or rocky hillsides up to 2300 m, and cultivation on poor soil. Sometimes among xerophytes. Markedly terrestrial; unobtrusive, but not shy and may allow close approach. Occurs singly or in groups. Forms small parties in breeding area after 2nd broods fledge, and in winter, when it may associate with other seedeaters. May mix with House Buntings in winter on low, dry plains. Forages on ground among rocks and scrubby vegetation, and in short grass in fields and at woodland edges; feeds in bushes or tall herbs. Gait a hop; also creeps when feeding. Stance typically half upright. Flicks and spreads tail, exposing white outer feathers. When flushed, usually flies up to boulders or tree canopy (Cramp and Perrins 1994). Regular autumn passage migrant at Gibraltar, peaking in late Oct. Partly dispersive in NW Africa, wintering in areas where it does not breed, such as hills in NW Morocco (Pineau and Geraud-Audine 1979), the Mitidja in N Algeria (Ledant et al. 1981) and the Saharan borders. Food. Seeds, mainly of grasses, and other plant items; invertebrates in breeding season (A). Animal prey in Europe comprises mainly beetles, Hemiptera, Orthoptera, ad. and larval Lepidoptera, ants, spiders and snails.

Breeding Habits. Monogamous, territorial. Territory size (Germany, n ¼ 35) 11–43 (av. 255) ha. Adults range up to 150–200 m from nest to find food for nestlings. Y defends territory by singing, most frequently before nest-building, declining during incubation, infrequently when young being fed. Unpaired YY sing into summer. Often 2 broods in Europe. NEST: cup of dry grass, stalks and roots, sometimes leaves and bark, lined with fine grasses, rootlets and some hair; ext. diam. 100–160 (av. 129), int. diam. 55–65 (59), height 55–91 (72), cup depth 29–52 (40); on or close to ground, under a rock or bush, usually hidden by vegetation; sometimes low in dense tree or bush; built by X in 3–7 days with material collected within 10–30 m. Nest for 2nd brood built 8–65 m (av. 26 m) from 1st nest (Cramp and Perrins 1994). EGGS: 3–6 (Algeria, n ¼ 42, av. 414). Pale greyish to purplish white, heavily and intricately scrawled with dark violet to black hair streaks, sometimes forming ring at broad end; a few underlying grey or lilac-grey smears. SIZE: (n ¼ 7, N Africa) 20–22  16 (208–160). LAYING DATES: Algeria, early Apr–mid June. INCUBATION: by X only, starting with last egg. Period: 12–14 days. DEVELOPMENT AND CARE OF YOUNG: fed by both sexes, at first mainly by X; brooded only by X. Leave nest at 10–13 days, then remain close together for several more days in thick ground vegetation, until they fly well and move into bushes. Fledged young fed by parents for 2–3 weeks. BREEDING SUCCESS/SURVIVAL: in Germany, 35 fledged young per successful nest (n ¼ 21); 18 overall (n ¼ 41) (Cramp and Perrins 1994). Key References C.M. (1994).

Byers, C. et al. (1995), Cramp, S. and Perrins,

Plate 35

Emberiza leucocephalos Gmelin. Pine Bunting. Bruant a` calotte blanche.

(Opp. p. 538)

Emberiza leucocephala Gmelin, 1771. Nov. Comm. Acad. Sci. Petrop., 15, p. 480; Astrakhan. E Palearctic, breeding west to 55
E; winters south to Iran, Pakistan, N India and central China; vagrant to Arabia and N, W and S Europe; regular in Israel, almost so in NE Italy (100 records). 1 Y, presumably of nominate race, Oukaı¨meden, Morocco, Jan 1995, in a flock of Chaffinches Fringilla coelebs (Edgeller 1995). Forms a superspecies with Yellowhammer E. citrinella; the 2 overlap extensively in W Siberia (where YY of both species have been seen singing in same tree) but often hybridize, with gene flow distantly to both W and E (Cramp and Perrins 1994, Byers et al. 1995). Formerly, often treated as conspecific with E. citrinella. Adult Y (breeding): crown white, bordered on forehead, sides and hindneck with broad black band; malar stripe, cheeks and ear-coverts white, the last bordered with black; rest of head chestnut (supercilium, lores, behind eyes, chin, throat and

588

EMBERIZIDAE

stripes still distinctive. In N Africa can only be confused with smaller House Bunting E. striolata, which has bright rufous wings, finely streaked head and breast, no white in tail. Juv. heavily streaked on head and breast, unlike juv. House Bunting, and has white in tail, striped wings with little rufous. Voice. Tape-recorded (62–73, 92, 105, 110–113, B). Extremely variable song thin and high-pitched but not unmusical, a medley of chips, short trills and slurred notes, with jerky quality reminiscent of Dunnock Prunella modularis, ‘trrr, see-so, chichi siwirr chit’, or ‘chyo titi tsiker ziwo sisisisi chewy’; rather fast and long for a bunting (up to 3–4 s). Common contact call a sharp ‘tsee’ or ‘tzit’; other calls include thin, descending ‘seeee’ or ‘seea’; short ‘tucc’, repeated to become twitter in flight; on rising, ‘chelut’, ‘chit vit’ or slightly vibrant ‘chee dee dee’ (Jonsson 1992). For sonagrams and further details see Cramp and Perrins (1994). General Habits. Inhabits high, open, semi-arid areas with sparse shrub vegetation and few trees; dry stony or rocky hillsides up to 2300 m, and cultivation on poor soil. Sometimes among xerophytes. Markedly terrestrial; unobtrusive, but not shy and may allow close approach. Occurs singly or in groups. Forms small parties in breeding area after 2nd broods fledge, and in winter, when it may associate with other seedeaters. May mix with House Buntings in winter on low, dry plains. Forages on ground among rocks and scrubby vegetation, and in short grass in fields and at woodland edges; feeds in bushes or tall herbs. Gait a hop; also creeps when feeding. Stance typically half upright. Flicks and spreads tail, exposing white outer feathers. When flushed, usually flies up to boulders or tree canopy (Cramp and Perrins 1994). Regular autumn passage migrant at Gibraltar, peaking in late Oct. Partly dispersive in NW Africa, wintering in areas where it does not breed, such as hills in NW Morocco (Pineau and Geraud-Audine 1979), the Mitidja in N Algeria (Ledant et al. 1981) and the Saharan borders. Food. Seeds, mainly of grasses, and other plant items; invertebrates in breeding season (A). Animal prey in Europe comprises mainly beetles, Hemiptera, Orthoptera, ad. and larval Lepidoptera, ants, spiders and snails.

Breeding Habits. Monogamous, territorial. Territory size (Germany, n ¼ 35) 11–43 (av. 255) ha. Adults range up to 150–200 m from nest to find food for nestlings. Y defends territory by singing, most frequently before nest-building, declining during incubation, infrequently when young being fed. Unpaired YY sing into summer. Often 2 broods in Europe. NEST: cup of dry grass, stalks and roots, sometimes leaves and bark, lined with fine grasses, rootlets and some hair; ext. diam. 100–160 (av. 129), int. diam. 55–65 (59), height 55–91 (72), cup depth 29–52 (40); on or close to ground, under a rock or bush, usually hidden by vegetation; sometimes low in dense tree or bush; built by X in 3–7 days with material collected within 10–30 m. Nest for 2nd brood built 8–65 m (av. 26 m) from 1st nest (Cramp and Perrins 1994). EGGS: 3–6 (Algeria, n ¼ 42, av. 414). Pale greyish to purplish white, heavily and intricately scrawled with dark violet to black hair streaks, sometimes forming ring at broad end; a few underlying grey or lilac-grey smears. SIZE: (n ¼ 7, N Africa) 20–22  16 (208–160). LAYING DATES: Algeria, early Apr–mid June. INCUBATION: by X only, starting with last egg. Period: 12–14 days. DEVELOPMENT AND CARE OF YOUNG: fed by both sexes, at first mainly by X; brooded only by X. Leave nest at 10–13 days, then remain close together for several more days in thick ground vegetation, until they fly well and move into bushes. Fledged young fed by parents for 2–3 weeks. BREEDING SUCCESS/SURVIVAL: in Germany, 35 fledged young per successful nest (n ¼ 21); 18 overall (n ¼ 41) (Cramp and Perrins 1994). Key References C.M. (1994).

Byers, C. et al. (1995), Cramp, S. and Perrins,

Plate 35

Emberiza leucocephalos Gmelin. Pine Bunting. Bruant a` calotte blanche.

(Opp. p. 538)

Emberiza leucocephala Gmelin, 1771. Nov. Comm. Acad. Sci. Petrop., 15, p. 480; Astrakhan. E Palearctic, breeding west to 55
E; winters south to Iran, Pakistan, N India and central China; vagrant to Arabia and N, W and S Europe; regular in Israel, almost so in NE Italy (100 records). 1 Y, presumably of nominate race, Oukaı¨meden, Morocco, Jan 1995, in a flock of Chaffinches Fringilla coelebs (Edgeller 1995). Forms a superspecies with Yellowhammer E. citrinella; the 2 overlap extensively in W Siberia (where YY of both species have been seen singing in same tree) but often hybridize, with gene flow distantly to both W and E (Cramp and Perrins 1994, Byers et al. 1995). Formerly, often treated as conspecific with E. citrinella. Adult Y (breeding): crown white, bordered on forehead, sides and hindneck with broad black band; malar stripe, cheeks and ear-coverts white, the last bordered with black; rest of head chestnut (supercilium, lores, behind eyes, chin, throat and

Emberiza cirlus

589

sides of neck); broad white band across upper breast; rest of plumage, and bare parts, like ad. Y E. citrinella, q.v., except that breast, belly and flanks white where E. citrinella yellow. Other plumages (non-breeding Y; X, imm.) very like those of E. citrinella but whitish where E. citrinella yellowish; hybrids may be yellowish, however (see Byers et al. 1995, plate 3). Length 16–165 cm; wing, Y (n ¼ 39) 88–100 (940), X (n ¼ 17) 84–95 (889); tail, Y (n ¼ 27) 68–83 (750), X (n ¼ 10) 64–75 (691); weight (Belgium and France, winter) Y (n ¼ 4) 198–307 (251), X (n ¼ 3) 235–314 (280).

Emberiza citrinella Linnaeus. Yellowhammer. Bruant jaune.

Plate 35

Emberiza citrinella Linnaeus, 1758. Syst. Nat., ed. 10, p 177; Sweden.

(Opp. p. 538)

Europe and N Asia east to c. 110
E in Siberia. Sedentary or partial migrant; winter visitor to Spain, Italy, Balkans, W Turkey and N Israel; rare, Gibraltar, Malta (13 records Oct–Dec), Sicily (6) and Jordon. Rare vagrant, presumably European E. c. citrinella, to N Morocco (1, unconfirmed, Korifla, Jan 1980, 1, unconfirmed, Ras el Ma (Ifrane), Apr 1981, 1 near Kenitra, Nov 1991), N Algeria and Tunisia (Byers et al. 1995). A rather large bunting with yellow or yellow-tinged head and underparts bordered on sides by dark olive streaks; yellow side of head with dark stripes behind eye and around lower border of ear-coverts; upperparts warm brown with blackish streaking; underparts lemon-yellow with olive-grey and rufous tinged breast band, and long rufous-chestnut flank streaks. Autumn Y has broader head streaks and blackish streaks on breast and flanks. X duller, buffier above and paler yellow below with streaked crown centre, breast and flanks streaked dark brown rather than chestnut. Longer-tailed, less compact looking than Cirl Bunting E. cirlus, which lacks rufous rump. Length 16–17 cm; wing, Y (n ¼ 10) 86–93 (904), X (n ¼ 10) 79–88 (826); weight, SE France, winter, Y (n ¼ 27) 25–38 (289), X (n ¼ 26) 23–30 (274). Commonest call a clipped metallic ‘zit’ or ‘tzit’; in flight, single or multiple ‘tit’ notes. Frequents open country and cultivation with hedges and scattered trees. Forages on ground for grass seeds, usually with other species.

Emberiza cirlus Linnaeus. Cirl Bunting. Bruant zizi.

Plate 35

Emberiza cirlus Linnaeus 1766, Syst. Nat., ed. 12, p. 311; S Europe.

(Opp. p. 538)

Range and Status. Resident, W and S Europe and NW Africa; breeds north to S England, east to Balkans and Turkey. Resident N and W Morocco, N Algeria and extreme NW Tunisia; largely sedentary, but some local winter movement. Morocco, from Mediterranean coastal areas, Djebala and Rif to Moyen-Atlas, Azrou and Ke´nitra areas and N foothills of Haut-Atlas; also at W extremity of Anti-Atlas at Djebel Kest, Abeino, Ifni and Djebel Guir, but absent from Sous valley and Atlantic coast; frequent to common, mainly on lower valley slopes. Algeria, frequent from coast to N slopes of Atlas Tellien, common in Aure`s Mts; reaches edge of Sahara (El Kantara, Oumache, Mts des Ksours, Baniane) in winter. Tunisia, frequent in Kroumerie, south to Le Ke´f; scarce Medjerga valley, Zaghouan and St-Marie du Zit; rare south to Sousse, Kairouan and Gabe`s. Vagrant Egypt (once, Damietta, 1892). Breeding density, Morocco, 38–255 pairs per km2, av. 160 (The´venot 1982). Winter density, near Gibraltar, av. 61 birds per km2 in shrubland and 157 in cork oak (Arroyo and Tellerı´a 1984). Description. ADULT Y: top of head to hindneck greenish grey, forehead to nape narrowly streaked black; mantle and inner scapulars chestnut, broadly streaked black, feathers tipped greybuff when fresh; outer scapulars chestnut with little if any dark streaking; back, rump and uppertail-coverts greyish olive-green, with some chestnut on back and usually some dark grey shaft streaks. Tail dark grey-brown; T1 fringed greenish olive on outer

Emberiza cirlus

web, T2–T5 more narrowly so; inner web of T5–T6 with white distal wedge, extending along shaft to two-thirds of length of T6,

Emberiza cirlus

589

sides of neck); broad white band across upper breast; rest of plumage, and bare parts, like ad. Y E. citrinella, q.v., except that breast, belly and flanks white where E. citrinella yellow. Other plumages (non-breeding Y; X, imm.) very like those of E. citrinella but whitish where E. citrinella yellowish; hybrids may be yellowish, however (see Byers et al. 1995, plate 3). Length 16–165 cm; wing, Y (n ¼ 39) 88–100 (940), X (n ¼ 17) 84–95 (889); tail, Y (n ¼ 27) 68–83 (750), X (n ¼ 10) 64–75 (691); weight (Belgium and France, winter) Y (n ¼ 4) 198–307 (251), X (n ¼ 3) 235–314 (280).

Emberiza citrinella Linnaeus. Yellowhammer. Bruant jaune.

Plate 35

Emberiza citrinella Linnaeus, 1758. Syst. Nat., ed. 10, p 177; Sweden.

(Opp. p. 538)

Europe and N Asia east to c. 110
E in Siberia. Sedentary or partial migrant; winter visitor to Spain, Italy, Balkans, W Turkey and N Israel; rare, Gibraltar, Malta (13 records Oct–Dec), Sicily (6) and Jordon. Rare vagrant, presumably European E. c. citrinella, to N Morocco (1, unconfirmed, Korifla, Jan 1980, 1, unconfirmed, Ras el Ma (Ifrane), Apr 1981, 1 near Kenitra, Nov 1991), N Algeria and Tunisia (Byers et al. 1995). A rather large bunting with yellow or yellow-tinged head and underparts bordered on sides by dark olive streaks; yellow side of head with dark stripes behind eye and around lower border of ear-coverts; upperparts warm brown with blackish streaking; underparts lemon-yellow with olive-grey and rufous tinged breast band, and long rufous-chestnut flank streaks. Autumn Y has broader head streaks and blackish streaks on breast and flanks. X duller, buffier above and paler yellow below with streaked crown centre, breast and flanks streaked dark brown rather than chestnut. Longer-tailed, less compact looking than Cirl Bunting E. cirlus, which lacks rufous rump. Length 16–17 cm; wing, Y (n ¼ 10) 86–93 (904), X (n ¼ 10) 79–88 (826); weight, SE France, winter, Y (n ¼ 27) 25–38 (289), X (n ¼ 26) 23–30 (274). Commonest call a clipped metallic ‘zit’ or ‘tzit’; in flight, single or multiple ‘tit’ notes. Frequents open country and cultivation with hedges and scattered trees. Forages on ground for grass seeds, usually with other species.

Emberiza cirlus Linnaeus. Cirl Bunting. Bruant zizi.

Plate 35

Emberiza cirlus Linnaeus 1766, Syst. Nat., ed. 12, p. 311; S Europe.

(Opp. p. 538)

Range and Status. Resident, W and S Europe and NW Africa; breeds north to S England, east to Balkans and Turkey. Resident N and W Morocco, N Algeria and extreme NW Tunisia; largely sedentary, but some local winter movement. Morocco, from Mediterranean coastal areas, Djebala and Rif to Moyen-Atlas, Azrou and Ke´nitra areas and N foothills of Haut-Atlas; also at W extremity of Anti-Atlas at Djebel Kest, Abeino, Ifni and Djebel Guir, but absent from Sous valley and Atlantic coast; frequent to common, mainly on lower valley slopes. Algeria, frequent from coast to N slopes of Atlas Tellien, common in Aure`s Mts; reaches edge of Sahara (El Kantara, Oumache, Mts des Ksours, Baniane) in winter. Tunisia, frequent in Kroumerie, south to Le Ke´f; scarce Medjerga valley, Zaghouan and St-Marie du Zit; rare south to Sousse, Kairouan and Gabe`s. Vagrant Egypt (once, Damietta, 1892). Breeding density, Morocco, 38–255 pairs per km2, av. 160 (The´venot 1982). Winter density, near Gibraltar, av. 61 birds per km2 in shrubland and 157 in cork oak (Arroyo and Tellerı´a 1984). Description. ADULT Y: top of head to hindneck greenish grey, forehead to nape narrowly streaked black; mantle and inner scapulars chestnut, broadly streaked black, feathers tipped greybuff when fresh; outer scapulars chestnut with little if any dark streaking; back, rump and uppertail-coverts greyish olive-green, with some chestnut on back and usually some dark grey shaft streaks. Tail dark grey-brown; T1 fringed greenish olive on outer

Emberiza cirlus

web, T2–T5 more narrowly so; inner web of T5–T6 with white distal wedge, extending along shaft to two-thirds of length of T6,

Emberiza cirlus

589

sides of neck); broad white band across upper breast; rest of plumage, and bare parts, like ad. Y E. citrinella, q.v., except that breast, belly and flanks white where E. citrinella yellow. Other plumages (non-breeding Y; X, imm.) very like those of E. citrinella but whitish where E. citrinella yellowish; hybrids may be yellowish, however (see Byers et al. 1995, plate 3). Length 16–165 cm; wing, Y (n ¼ 39) 88–100 (940), X (n ¼ 17) 84–95 (889); tail, Y (n ¼ 27) 68–83 (750), X (n ¼ 10) 64–75 (691); weight (Belgium and France, winter) Y (n ¼ 4) 198–307 (251), X (n ¼ 3) 235–314 (280).

Emberiza citrinella Linnaeus. Yellowhammer. Bruant jaune.

Plate 35

Emberiza citrinella Linnaeus, 1758. Syst. Nat., ed. 10, p 177; Sweden.

(Opp. p. 538)

Europe and N Asia east to c. 110
E in Siberia. Sedentary or partial migrant; winter visitor to Spain, Italy, Balkans, W Turkey and N Israel; rare, Gibraltar, Malta (13 records Oct–Dec), Sicily (6) and Jordon. Rare vagrant, presumably European E. c. citrinella, to N Morocco (1, unconfirmed, Korifla, Jan 1980, 1, unconfirmed, Ras el Ma (Ifrane), Apr 1981, 1 near Kenitra, Nov 1991), N Algeria and Tunisia (Byers et al. 1995). A rather large bunting with yellow or yellow-tinged head and underparts bordered on sides by dark olive streaks; yellow side of head with dark stripes behind eye and around lower border of ear-coverts; upperparts warm brown with blackish streaking; underparts lemon-yellow with olive-grey and rufous tinged breast band, and long rufous-chestnut flank streaks. Autumn Y has broader head streaks and blackish streaks on breast and flanks. X duller, buffier above and paler yellow below with streaked crown centre, breast and flanks streaked dark brown rather than chestnut. Longer-tailed, less compact looking than Cirl Bunting E. cirlus, which lacks rufous rump. Length 16–17 cm; wing, Y (n ¼ 10) 86–93 (904), X (n ¼ 10) 79–88 (826); weight, SE France, winter, Y (n ¼ 27) 25–38 (289), X (n ¼ 26) 23–30 (274). Commonest call a clipped metallic ‘zit’ or ‘tzit’; in flight, single or multiple ‘tit’ notes. Frequents open country and cultivation with hedges and scattered trees. Forages on ground for grass seeds, usually with other species.

Emberiza cirlus Linnaeus. Cirl Bunting. Bruant zizi.

Plate 35

Emberiza cirlus Linnaeus 1766, Syst. Nat., ed. 12, p. 311; S Europe.

(Opp. p. 538)

Range and Status. Resident, W and S Europe and NW Africa; breeds north to S England, east to Balkans and Turkey. Resident N and W Morocco, N Algeria and extreme NW Tunisia; largely sedentary, but some local winter movement. Morocco, from Mediterranean coastal areas, Djebala and Rif to Moyen-Atlas, Azrou and Ke´nitra areas and N foothills of Haut-Atlas; also at W extremity of Anti-Atlas at Djebel Kest, Abeino, Ifni and Djebel Guir, but absent from Sous valley and Atlantic coast; frequent to common, mainly on lower valley slopes. Algeria, frequent from coast to N slopes of Atlas Tellien, common in Aure`s Mts; reaches edge of Sahara (El Kantara, Oumache, Mts des Ksours, Baniane) in winter. Tunisia, frequent in Kroumerie, south to Le Ke´f; scarce Medjerga valley, Zaghouan and St-Marie du Zit; rare south to Sousse, Kairouan and Gabe`s. Vagrant Egypt (once, Damietta, 1892). Breeding density, Morocco, 38–255 pairs per km2, av. 160 (The´venot 1982). Winter density, near Gibraltar, av. 61 birds per km2 in shrubland and 157 in cork oak (Arroyo and Tellerı´a 1984). Description. ADULT Y: top of head to hindneck greenish grey, forehead to nape narrowly streaked black; mantle and inner scapulars chestnut, broadly streaked black, feathers tipped greybuff when fresh; outer scapulars chestnut with little if any dark streaking; back, rump and uppertail-coverts greyish olive-green, with some chestnut on back and usually some dark grey shaft streaks. Tail dark grey-brown; T1 fringed greenish olive on outer

Emberiza cirlus

web, T2–T5 more narrowly so; inner web of T5–T6 with white distal wedge, extending along shaft to two-thirds of length of T6,

590

EMBERIZIDAE

one-third of T5; tip of inner web of T4 sometimes also white; T6 with middle and distal border of outer web white, shaft dark grey, widening on tip. Superciliary stripe pale yellow, broadest behind eye, extending back to side of nape; below this a black band through lores and upper ear-coverts, and another broad pale yellow stripe across upper cheek and lower ear-coverts. Lower cheek to chin and upper throat black (greyish tipped when fresh), extending back below yellow cheek stripe; below this a narrow yellow collar from lower throat to front lower side of neck and a broad greygreen band across upper breast, continuous with grey-green sides of neck. Sides of lower breast chestnut; centre of lower breast and belly pale yellow; flanks pale tawny buff with long dark brown streaks; undertail-coverts yellowish white. Large upperwing feathers dark grey-brown; primaries narrowly edged pale greenish olive, secondaries edged olive, primary coverts and small alula feather narrowly fringed pale green; outer webs and tips of tertials broadly bordered chestnut-brown; greater coverts broadly edged and tipped olive (more chestnut on inner feathers); median coverts narrowly fringed pale yellow or whitish. Lesser coverts grey, fringed greenish. Underwing-coverts and axillaries pale olive-yellow. Bill dark horn above, light bluish to grey below; eyes brown; feet brownish flesh. ADULT X: top of head greyish olive, streaked blackish; mantle to inner scapulars greyish olive, usually with some chestnut, feathers with black shaft streaks broadening at tip; outer scapulars more chestnut and with narrower streaking; back to uppertail-coverts greyish olive-green, with dusky streaking usually faint. Tail as in ad. Y. Superciliary stripe pale buffish, more conspicuous behind eye; post-ocular stripe dark brown, flecked blackish; lores, upper cheek and lower ear-coverts buffy yellow, mottled dusky, bordered by brown moustachial stripe below and narrow brown line behind, and with pale buff spot at rear of ear-coverts. Broad sub-moustachial stripe buffy yellow, flecked dusky, and below this a narrow blackish malar stripe. Underparts pale buffy yellow or oliveyellow, breast slightly browner, tinged tawny at sides; throat and breast with short blackish streaks, flanks with long streaks; undertail-coverts buffy white with fine dusky streaks. Wing as in ad. Y, but outer borders of tertials and inner greater coverts more olive-brown or tawny (less chestnut). Bare parts as in ad. Y. Complete moult July–Oct. SIZE (10 YY, 10 XX, NW Africa and Spain): wing, Y 75–83 (803), X 74–80 (769); tail, Y 68–73 (704), X 65–71 (678); bill, Y 13–145 (138), X 135–145 (139); tarsus, Y 18–20 (191), X 18–20 (191). WEIGHT: (S France) Y (n ¼ 39) 23–28 (257), X (n ¼ 24) 23–28 (255). IMMATURE: juv. similar to ad. X, but paler, buffier brown above, streaked blackish brown from forehead to uppertail-coverts; underparts pale buffish, less yellow, breast more strongly streaked; fringes of tertials, greater coverts and median coverts sharper, less broad, buffier. First-winter like ad. NESTLING: down long and plentiful; grey, grey-brown, greycinnamon. Inside of mouth pink or red, gape-flanges lemon; legs pale yellow.

Field Characters. Length 16 cm. Distinguished in all plumages from vagrant Yellowhammer E. citrinella by olive-brown (not rufous) rump. Y distinctive all year, with black and yellow face and throat, olive breast-band, chestnut breast patch and scapulars; X paler yellow below than X Yellowhammer, with finer streaking, more strongly patterned face; chestnut scapulars and tertials contrast with greyish back and rump. Juv. paler and buffier than X, with finely streaked upperparts, more heavily streaked breast; face marks buff rather than yellow. Voice. Tape-recorded (62–73, 105, 110–113, B). Song a short (c. 125 s) rattle or trill on 1 pitch at varying speeds, typically dry, sometimes liquid, or with buzzy quality

suggesting Yellowhammer (but without the terminal ‘cheese’); 1 bird may have several song types. Quieter trill of 4–5 notes given in winter. Commonest call a short ‘sit’, ‘zit’ or ‘tzip’; warning or alarm, a loud descending ‘seeee’ or ‘tzeeep’, recalling Song Thrush Turdus philomelos. For threat and other calls, see Cramp and Perrins (1994). General Habits. Inhabits slopes with scrub and myrtle Myrtus on edges of cork oak forest at medium altitudes; up to 1600 m in Haut-Atlas and 1800 m in Djurdjura; down to coastal plains and cultivation. Clearings in date palm groves, large gardens. In winter, favours areas with patches of dense thicket, scattered trees, rough pasture and cultivation. Outside breeding season, some birds remain paired on territories, but otherwise forms parties and small flocks. Forages almost wholly on ground, among sparse vegetation and low growth, often on trampled or grazed grass, tracks or field edges close to cover, rarely on bare soil. Not shy when feeding. Gait a hop, often slow and shuffling. Stance somewhat hunched, and may droop wingtips, exposing rump. Sits on 1 or on both tarsi. When anxious, spreads tail with flicking action, exposing white outer feathers. Often raises crest feathers. May ‘freeze’ when disturbed; usually then flies to low cover. Rests and preens in taller trees. Roosts in thick vegetation 1 m or more above ground (Cramp and Perrins 1994). Mainly sedentary, but some southward movement in autumn and a few birds reach N edge of Sahara in winter, south to Biskra and Mts des Ksours. Irregular winter visitor to Gibraltar, but not certainly an immigrant to Africa from Europe. Food. Seeds; and invertebrates in breeding season, including grasshoppers, earwigs, bugs, ad. and larval Lepidoptera, Hymenoptera, beetles, spiders, earthworms and snails. Young given mostly invertebrates, but also seeds; ripening grain apparently important for later broods. Breeding Habits. Monogamous; pair may stay together for several years. Strongly territorial; territories can be clumped, but Y excludes others from within 150 m of nest. Territory size (n ¼ 22, SW Germany) 048–183 ha (094) (Groh 1975). Territory boundaries established by singing, also by threat and fighting. Nests are sometimes close to those of Rock Bunting. Y sings throughout much of day, usually from tall tree, bushtop or wire, sometimes from within thorny bush, head often thrown back (emphasizing pattern), bill wide open, wings and tail lowered and body vibrating. Threatens intruding Y by crouching low, wings held out to side, head and tail slightly raised, bill agape; may pursue intruder and engage in breast-to-breast fight in the air. No noticeable courtship display. Pair on territory stays close together. Unpaired YY, often in poorer territories, perch and sing conspicuously. 2 broods (sometimes 3 in Europe). NEST: rather bulky, untidy cup of rough stalks, roots, grass, leaves and moss, lined with finer grass and hair. Ext. diam. 100–175 (120–125), int. diam. 55–82 (66), ext. height 58–102 (67), cup depth 32–61 (43); low down, well-hidden in dense tree, shrub, hedge or creeper; sometimes in rock

Emberiza cineracea crevice; usually below 2 m, rarely on ground. Built by X, often accompanied by Y, in 4–6 days. Used only once, but 2nd brood may be in nest built only a few m away. EGGS: 3–5 (50 clutches, Algeria) av. 416 (Heim de Balzac and Mayaud 1962); laid daily in early morning, starting 1–3 days after nest completed. Greyish white, often tinged pinkish, blue or green, finely speckled brown, with a few bold blackish scrawls and dots concentrated near broad end, and pale grey undermarkings. SIZE (n ¼ 343, Europe) 19–24  145–18 (214  161). LAYING DATES: NW Africa, early Apr–June. 2nd nest started c. 7 days after 1st brood becomes independent.

INCUBATION:

591

entirely by X, fed on nest by Y. Period

12–14 days. CARE AND DEVELOPMENT OF YOUNG: fed and cared for by both parents; brooded by X for up to 9 days. Fledge after 11–13 days; independent after another 8–16 days. BREEDING SUCCESS/SURVIVAL: in Valencia (Spain), of 86 eggs, 65% hatched, 31% produced fledged young; in Germany, 37% of 73 nests failed (Cramp and Perrins 1994).

Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Groh, G. (1975).

Emberiza cineracea C.L. Brehm. Cinereous Bunting. Bruant cendre ´.

Plate 36

Emberiza cineracea C.L. Brehm, 1845. Der vollsta¨ndige Vogelfang, p. 114; Smyrna.

(Opp. p. 539)

Range and Status. Greece (Lesbos and Chios), W and SE Turkey, SW Iran; winters NE Africa and SW Arabia. Winters NE Sudan (Red Sea Prov.) and Eritrea, inland to central plateau at 2200 m (Zinner 2001); uncommon. Rare passage migrant Egypt (Gebel Elba, Apr, El Gorah, May, near Au ¨ n Sukna, May); vagrant Tunisia (once Apr, Gabe`s). Description. E. c. semenowi Zarudnyi: SW Iran; winters SW Arabia, NE Sudan and Eritrea. ADULT Y: forehead to nape yellowish olive-green with faint dusky mottling; hindneck, sides of neck and upper mantle greyish olive; rest of upperparts dull brown or olive-brown, feathers of lower mantle and scapulars with browner centres and blackish shaft streaks, rump plain. Tail blackish brown; T1 fringed buff; inner webs of T5–T6 with square-cut white distal patch, occupying about one-third of T5 and half of T6; middle part of outer web of T6 also white. Lores and supercilium tinged yellower than crown; prominent narrow yellow eye-ring; upper cheeks and ear-coverts yellowish olivegrey; lower cheeks tinged yellower, bordered by faint olive-grey malar stripe. Chin and throat yellow, merging into olive-green breast band, sides and flanks; centre of lower breast and belly yellow; undertail-coverts yellowish buff; thighs buffish brown. Wing feathers dark grey-brown; primaries narrowly edged buff, secondaries edged cinnamon-buff, primary coverts and small alula feather narrowly fringed buff; outer webs and tips of tertials and outer webs of greater coverts fringed cinnamon-buff; greater coverts and median coverts tipped paler buff or grey-buff to form 2 wing-bars. Lesser coverts greyish olive-green. Underwingcoverts and axillaries greyish white, tinged olive-yellow. Bill pale blue-grey or horn-grey, darker on culmen; eyes brown; legs pale brown or flesh. ADULT X: upperparts dull brown or olive-brown, top of head sometimes tinged greenish, upper mantle and neck greyer; forehead to nape, lower mantle and scapulars narrowly streaked blackish brown. Ear-coverts and upper cheeks olivebrown, flecked yellowish buff, lores and supercilium slightly paler, more yellowish; lower cheeks pale olive-yellow, with faint broken dusky moustachial and malar stripes. Underparts pale olive-yellow, upper breast, sides and flanks tinged grey; chin and throat with dusky flecking, upper breast feathers with narrow dusky shaft streaks; undertail-coverts pale buff. Tail and wings as ad. Y. Bare parts as ad. Y. Complete moult July–Sept. SIZE (7 YY, 7 XX): wing, Y 89–98 (929), X 84–88 (863); tail, Y 72–79 (753), X 65–74 (699); bill, Y 145–155 (150), X 145–15 (149); tarsus, Y 195–205 (202), X 19–205 (195). WEIGHT: (NE Sudan, Sept, n ¼ 9) 185–25 (211).

Emberiza cineracea

IMMATURE: 1st-winter like ad., but finely streaked on breast, flanks and rump, X with throat pale buffy yellow; in retained juv. tail, white distal patches on T5–T6 often more V-shaped than in ad. E. c. cineracea C.L. Brehm: W and S Turkey; winters E Sudan and Eritrea (rare, Ghadi Saati, Mareb R.). Y differs from semenowi in having lower part of breast band, sides and flanks brownish grey (not greenish), merging into greyish white lower breast centre and belly; undertail-coverts off-white; upperparts duller brown, less olive, rump grey-brown. X has breast band grey-brown, underparts buffish white, sometimes little or no yellow on throat.

Field Characters. Length 16–17 cm. A rather large, plain bunting, streaked mainly on back, with noticeable pale yellow eye-ring and white outer tail-feathers. Y of nominate race ashy grey with yellow throat, olive breast-band, white belly; Y of eastern semenowi has underparts mainly yellow.

Emberiza cineracea crevice; usually below 2 m, rarely on ground. Built by X, often accompanied by Y, in 4–6 days. Used only once, but 2nd brood may be in nest built only a few m away. EGGS: 3–5 (50 clutches, Algeria) av. 416 (Heim de Balzac and Mayaud 1962); laid daily in early morning, starting 1–3 days after nest completed. Greyish white, often tinged pinkish, blue or green, finely speckled brown, with a few bold blackish scrawls and dots concentrated near broad end, and pale grey undermarkings. SIZE (n ¼ 343, Europe) 19–24  145–18 (214  161). LAYING DATES: NW Africa, early Apr–June. 2nd nest started c. 7 days after 1st brood becomes independent.

INCUBATION:

591

entirely by X, fed on nest by Y. Period

12–14 days. CARE AND DEVELOPMENT OF YOUNG: fed and cared for by both parents; brooded by X for up to 9 days. Fledge after 11–13 days; independent after another 8–16 days. BREEDING SUCCESS/SURVIVAL: in Valencia (Spain), of 86 eggs, 65% hatched, 31% produced fledged young; in Germany, 37% of 73 nests failed (Cramp and Perrins 1994).

Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Groh, G. (1975).

Emberiza cineracea C.L. Brehm. Cinereous Bunting. Bruant cendre ´.

Plate 36

Emberiza cineracea C.L. Brehm, 1845. Der vollsta¨ndige Vogelfang, p. 114; Smyrna.

(Opp. p. 539)

Range and Status. Greece (Lesbos and Chios), W and SE Turkey, SW Iran; winters NE Africa and SW Arabia. Winters NE Sudan (Red Sea Prov.) and Eritrea, inland to central plateau at 2200 m (Zinner 2001); uncommon. Rare passage migrant Egypt (Gebel Elba, Apr, El Gorah, May, near Au ¨ n Sukna, May); vagrant Tunisia (once Apr, Gabe`s). Description. E. c. semenowi Zarudnyi: SW Iran; winters SW Arabia, NE Sudan and Eritrea. ADULT Y: forehead to nape yellowish olive-green with faint dusky mottling; hindneck, sides of neck and upper mantle greyish olive; rest of upperparts dull brown or olive-brown, feathers of lower mantle and scapulars with browner centres and blackish shaft streaks, rump plain. Tail blackish brown; T1 fringed buff; inner webs of T5–T6 with square-cut white distal patch, occupying about one-third of T5 and half of T6; middle part of outer web of T6 also white. Lores and supercilium tinged yellower than crown; prominent narrow yellow eye-ring; upper cheeks and ear-coverts yellowish olivegrey; lower cheeks tinged yellower, bordered by faint olive-grey malar stripe. Chin and throat yellow, merging into olive-green breast band, sides and flanks; centre of lower breast and belly yellow; undertail-coverts yellowish buff; thighs buffish brown. Wing feathers dark grey-brown; primaries narrowly edged buff, secondaries edged cinnamon-buff, primary coverts and small alula feather narrowly fringed buff; outer webs and tips of tertials and outer webs of greater coverts fringed cinnamon-buff; greater coverts and median coverts tipped paler buff or grey-buff to form 2 wing-bars. Lesser coverts greyish olive-green. Underwingcoverts and axillaries greyish white, tinged olive-yellow. Bill pale blue-grey or horn-grey, darker on culmen; eyes brown; legs pale brown or flesh. ADULT X: upperparts dull brown or olive-brown, top of head sometimes tinged greenish, upper mantle and neck greyer; forehead to nape, lower mantle and scapulars narrowly streaked blackish brown. Ear-coverts and upper cheeks olivebrown, flecked yellowish buff, lores and supercilium slightly paler, more yellowish; lower cheeks pale olive-yellow, with faint broken dusky moustachial and malar stripes. Underparts pale olive-yellow, upper breast, sides and flanks tinged grey; chin and throat with dusky flecking, upper breast feathers with narrow dusky shaft streaks; undertail-coverts pale buff. Tail and wings as ad. Y. Bare parts as ad. Y. Complete moult July–Sept. SIZE (7 YY, 7 XX): wing, Y 89–98 (929), X 84–88 (863); tail, Y 72–79 (753), X 65–74 (699); bill, Y 145–155 (150), X 145–15 (149); tarsus, Y 195–205 (202), X 19–205 (195). WEIGHT: (NE Sudan, Sept, n ¼ 9) 185–25 (211).

Emberiza cineracea

IMMATURE: 1st-winter like ad., but finely streaked on breast, flanks and rump, X with throat pale buffy yellow; in retained juv. tail, white distal patches on T5–T6 often more V-shaped than in ad. E. c. cineracea C.L. Brehm: W and S Turkey; winters E Sudan and Eritrea (rare, Ghadi Saati, Mareb R.). Y differs from semenowi in having lower part of breast band, sides and flanks brownish grey (not greenish), merging into greyish white lower breast centre and belly; undertail-coverts off-white; upperparts duller brown, less olive, rump grey-brown. X has breast band grey-brown, underparts buffish white, sometimes little or no yellow on throat.

Field Characters. Length 16–17 cm. A rather large, plain bunting, streaked mainly on back, with noticeable pale yellow eye-ring and white outer tail-feathers. Y of nominate race ashy grey with yellow throat, olive breast-band, white belly; Y of eastern semenowi has underparts mainly yellow.

592

EMBERIZIDAE

X darker and browner above, with some streaks on breast, throat buffy yellow. Plain head, pale eye-ring and throat pattern suggest Ortolan or Cretzschmar’s Bunting E. hortulana and E. caesia, but distinguished by grey (not pink) bill, grey rump and lack of rufous on underparts. X told from X Black-headed Bunting E. melanocephala (also with plain head, grey bill, yellow-tinged underparts) by white in tail, more prominent eye-ring, yellower throat, lack of rufous on rump. Voice. Tape-recorded (62–73, 92, B, VIEL, MIL). Song, 3–4 ringing notes on one pitch, dropping off at end with 1–2 lower tuneless ones, ‘twee-twee-twee-twee-drit’, ‘dir-dir-dir-dir-dli-dlu’; lasts c. 1 s, given at intervals of c. 5 s. Calls include ‘tsik’ or ‘kyip’, hard ‘jeet’ and softer, clear descending ‘dyew’. General Habits. On passage and in winter, frequents dry country with scanty vegetation; short grassland, semidesert and cultivated areas; low rocky or scrub-covered hills; coastal plains. Unobtrusive but easy to approach.

Forages on ground, singly or in small groups. Feeds on autumn migration in dry stubble fields. In Sudan, associates with parties of Ortolan and Cretzschmar’s Buntings. 2 migration routes: south through Levant and Israel (mainly nominate cineracea) to E Sudan and Eritrea; and through Arabian Gulf areas (semenowi only) to winter quarters on either side of S Red Sea (Knijff 1991). Migrates at night, sometimes apparently with Ortolan and Cretzschmer’s Buntings (G. Nikolaus, pers. comm.). In autumn, passes through Israel coast and Red Sea Hills (mainly semenowi) from early Sept, mostly in mid–late Sept. In spring, returns through Israel (mainly cineracea) from late Aug to late Sept; arrives Sudan from late Mar to early May, commoner than in autumn. Food. On migration and in winter, mainly seeds; also small invertebrates. Passage birds at Elat (Israel) fed on beetles and spiders (Knijff 1991). Key References Collar, N.J. et al. (1994), Cramp, S. and Perrins, C.M. (1994), Knijff, P. de (1991), Svensson, L. (1992).

Plate 36

Emberiza hortulana Linnaeus. Ortolan Bunting. Bruant ortolan.

(Opp. p. 539)

Emberiza hortulana Linnaeus, 1758. Syst. Nat., ed. 10, p. 177; Sweden. Range and Status. Europe, S Russia and N Kazakhstan to NW Mongolia; Turkey and Caucasus to N Iran and Turkmenistan; and NW Africa. Winters in N tropics of Africa. Vagrant south to Seychelles. Rare breeder N Algeria: in Djurdjura (dozens of pairs since at least 1977), Petite-Kabylie, and probably in Aure`s Mts (Burnier 1977, Ledant et al. 1981); perhaps southeast of Setif (Cramp and Perrins 1994). Passage migrant in N Africa and Saharan oases, mainly in spring: uncommon Morocco; rare to uncommon Algeria, Tunisia and Libya; in Egypt, frequent in autumn (confined mainly to Suez Canal area and Eastern Desert), more common and widespread in spring (including Nile Valley and Western Desert). On S edge of Sahara, passage in SW Mauritania (common in autumn), Senegal (mainly in N), Gambia (rare), Mali (uncommon), Nigeria and Chad (rare) and N and central Sudan. Common to abundant autumn migrant in coastal Sudan and coastal Eritrea; scarce autumn records in N Somalia, presumably of passage birds. Winters between 7
and 15
N, mainly in upland areas, range apparently discontinuous: in NE Sierra Leone (Loma and Tingi Mts, locally common); S Guinea (Mt Nimba, common); N Nigeria (around Zaria and Jos, uncommon); W Sudan (Darfur, common); Eritrea and Ethiopia, frequent to common, in Eritrea at 50–2500 m (Zinner 2001), in Ethiopia mainly above 1200 m. Rare Liberia, Kenya (5 records), Uganda (once, Kibale, Oct; T. Stones, pers. comm.) and Zaı¨re (once, Goma; B.W. Finch, pers. comm.).

Emberiza hortulana ??

Description. ADULT Y (breeding): forehead to hindneck, sides of neck, upper cheeks and ear-coverts grey, tinged olive-green, sometimes with paler, yellow-tinged supraloral stripe; eye-ring pale yellow. Mantle and scapulars olive-brown or fulvous brown, boldly streaked blackish brown; back to uppertail-coverts tawny

592

EMBERIZIDAE

X darker and browner above, with some streaks on breast, throat buffy yellow. Plain head, pale eye-ring and throat pattern suggest Ortolan or Cretzschmar’s Bunting E. hortulana and E. caesia, but distinguished by grey (not pink) bill, grey rump and lack of rufous on underparts. X told from X Black-headed Bunting E. melanocephala (also with plain head, grey bill, yellow-tinged underparts) by white in tail, more prominent eye-ring, yellower throat, lack of rufous on rump. Voice. Tape-recorded (62–73, 92, B, VIEL, MIL). Song, 3–4 ringing notes on one pitch, dropping off at end with 1–2 lower tuneless ones, ‘twee-twee-twee-twee-drit’, ‘dir-dir-dir-dir-dli-dlu’; lasts c. 1 s, given at intervals of c. 5 s. Calls include ‘tsik’ or ‘kyip’, hard ‘jeet’ and softer, clear descending ‘dyew’. General Habits. On passage and in winter, frequents dry country with scanty vegetation; short grassland, semidesert and cultivated areas; low rocky or scrub-covered hills; coastal plains. Unobtrusive but easy to approach.

Forages on ground, singly or in small groups. Feeds on autumn migration in dry stubble fields. In Sudan, associates with parties of Ortolan and Cretzschmar’s Buntings. 2 migration routes: south through Levant and Israel (mainly nominate cineracea) to E Sudan and Eritrea; and through Arabian Gulf areas (semenowi only) to winter quarters on either side of S Red Sea (Knijff 1991). Migrates at night, sometimes apparently with Ortolan and Cretzschmer’s Buntings (G. Nikolaus, pers. comm.). In autumn, passes through Israel coast and Red Sea Hills (mainly semenowi) from early Sept, mostly in mid–late Sept. In spring, returns through Israel (mainly cineracea) from late Aug to late Sept; arrives Sudan from late Mar to early May, commoner than in autumn. Food. On migration and in winter, mainly seeds; also small invertebrates. Passage birds at Elat (Israel) fed on beetles and spiders (Knijff 1991). Key References Collar, N.J. et al. (1994), Cramp, S. and Perrins, C.M. (1994), Knijff, P. de (1991), Svensson, L. (1992).

Plate 36

Emberiza hortulana Linnaeus. Ortolan Bunting. Bruant ortolan.

(Opp. p. 539)

Emberiza hortulana Linnaeus, 1758. Syst. Nat., ed. 10, p. 177; Sweden. Range and Status. Europe, S Russia and N Kazakhstan to NW Mongolia; Turkey and Caucasus to N Iran and Turkmenistan; and NW Africa. Winters in N tropics of Africa. Vagrant south to Seychelles. Rare breeder N Algeria: in Djurdjura (dozens of pairs since at least 1977), Petite-Kabylie, and probably in Aure`s Mts (Burnier 1977, Ledant et al. 1981); perhaps southeast of Setif (Cramp and Perrins 1994). Passage migrant in N Africa and Saharan oases, mainly in spring: uncommon Morocco; rare to uncommon Algeria, Tunisia and Libya; in Egypt, frequent in autumn (confined mainly to Suez Canal area and Eastern Desert), more common and widespread in spring (including Nile Valley and Western Desert). On S edge of Sahara, passage in SW Mauritania (common in autumn), Senegal (mainly in N), Gambia (rare), Mali (uncommon), Nigeria and Chad (rare) and N and central Sudan. Common to abundant autumn migrant in coastal Sudan and coastal Eritrea; scarce autumn records in N Somalia, presumably of passage birds. Winters between 7
and 15
N, mainly in upland areas, range apparently discontinuous: in NE Sierra Leone (Loma and Tingi Mts, locally common); S Guinea (Mt Nimba, common); N Nigeria (around Zaria and Jos, uncommon); W Sudan (Darfur, common); Eritrea and Ethiopia, frequent to common, in Eritrea at 50–2500 m (Zinner 2001), in Ethiopia mainly above 1200 m. Rare Liberia, Kenya (5 records), Uganda (once, Kibale, Oct; T. Stones, pers. comm.) and Zaı¨re (once, Goma; B.W. Finch, pers. comm.).

Emberiza hortulana ??

Description. ADULT Y (breeding): forehead to hindneck, sides of neck, upper cheeks and ear-coverts grey, tinged olive-green, sometimes with paler, yellow-tinged supraloral stripe; eye-ring pale yellow. Mantle and scapulars olive-brown or fulvous brown, boldly streaked blackish brown; back to uppertail-coverts tawny

Emberiza hortulana brown, feather centres sometimes with faint dusky marking. Tail blackish brown; both webs of T1 fringed tawny brown; inner web of T6 with white distal wedge 25–40 mm long, inner web of T5 with wedge 22–30 mm long, inner web of T4 sometimes with short white edge or subterminal spot; T6 also with base and middle of outer web white, shaft black. Broad pale yellow submoustachial stripe, extending around lower rear edge of earcoverts, bordered below by olive-grey malar stripe. Chin and throat pale yellow; broad band across upper breast olive-grey; rest of underparts rufous-cinnamon, brighter on breast, paler, more cinnamon-buff on flanks, thighs and undertail-coverts. Flight feathers, primary coverts and alula blackish brown; primaries narrowly edged buff, secondaries more broadly edged cinnamonbuff, primary coverts and small alula feather narrowly fringed buff. Tertials, greater and median coverts black; tertials fringed cinnamon-buff, broadly on outer edge; edges and tips of greater coverts and tips of median coverts cinnamon-buff or grey-buff. Lesser coverts dark brown, tipped olive-grey. Underwing-coverts and axillaries white, tinged olive-yellow. Bill pinkish or reddish flesh; eyes dark brown; legs light brown or pinkish brown. ADULT Y (non-breeding): like breeding Y, but top and side of head more greenish, with fine dusky streaks; upper breast band yellowish olive-grey, with a few dark olive-grey streaks. ADULT X: differs from ad. Y in having distinct fine blackish shaft streaks from forehead to hindneck; upper cheeks and ear-coverts brown; submoustachial stripe, chin and throat pale yellowish buff or creamy buff, chin and throat flecked blackish brown; malar stripe more prominent, with small blackish streaks; sides of neck more brownish (less grey); upper breast band grey-buff with short blackish brown streaks; underparts paler, cinnamon or cinnamonbuff. Moults July–early Sept, complete except for some secondaries; partial moult of head and body in Africa, Dec–Mar. SIZE (10 YY, 10XX): wing, Y 84–93 (890), X 81–88 (836); tail, Y 63–71 (678), X 61–65 (635); bill, 13–145 (139), X 125–14 (135); tarsus, Y 185–205 (197), X 185–20 (193). WEIGHT: coastal Sudan, Sept, Y (n ¼ 95) 155–25 (202), X (n ¼ 117) 14–24 (189); central Nigeria, Mar–Apr, unsexed (n ¼ 5) 22–36; Morocco, spring, Y (n ¼ 8) 17–26 (201), X (n ¼ 3) 17–205. IMMATURE: juv. has entire upperparts brown, top of head to mantle and scapulars broadly streaked blackish; sides of head brown; sub-moustachial stripe, chin and throat buffy white. Underparts pale buff, malar stripe and upper breast band with broad triangular blackish feather centres, lower breast and flanks streaked blackish. Tail as ad. Wings as ad. but buff fringes on tertials extend more prominently around inner web; fringes of greater and median coverts paler buff, narrower, more sharply demarcated. 1st-winter Y like ad. X, but top of head browner, more boldly streaked; breast band with short heavy streaks, and some fine streaks also on lower breast and flanks; cinnamon-buff underparts less sharply demarcated, merging with yellow-buff of upper breast; sub-moustachial stripe and throat usually yellower. New cinnamon-buff outer edges of tertials contrast with narrow buffy white outer edges of retained juv. outer greater coverts. 1stwinter X like 1st-winter Y but buff of throat less yellow-tinged. NESTLING: down fairly long and plentiful, pearl-grey with buff tinge; on top of head, shoulder and back, traces on belly; mouth pink or flesh red, gape flanges pale cream-yellow (Cramp and Perrins 1994).

Field Characters. Length 16–17 cm. Plain head, pale eyering, pink bill and rufous-brown underparts shared with Cretzschmar’s Bunting E. caesia, but Y has yellow throat and sub-moustachial stripe, grey-green head, breast and malar stripe. X colours more subdued, with streaks on head and breast; no clear-cut division between grey breast and rufous underparts. 1st-winter birds often heavily streaked but still retain pale stripe curving under earcoverts, pink bill and whitish eye-ring, and underparts

tinged rufous-buff. When flushed shows light brown rump (grey in Cinereous Bunting E. cineracea, rufous in Cretzschmar’s and Yellowhammer E. citrinella). Voice. Tape-recorded (62–73, 76–89, 92, 105, 110–113, B, C). Trilled song of 6–8 notes in 2 parts, lasting 15–20 s; 1st part ringing but with buzzy overtone, 2nd part lowerpitched, sometimes a single note, ‘tsitsitsitsitsitsi-woy’, ‘pwee-pwee-pwee-pwee-woy’, sometimes a faster trill, ‘tseetsee-tsee-wawawawawa’, ‘weesu-weesu-weesu-chachacha’; African variations not described, but in S Europe second part usually a single hoarse note, ‘weez-weez-weez-zhaa’. Calls include alternating ‘tsyeep . . . tyep . . . tsyeep . . . tyep’ given by migrants; down-slurred ‘seeoo’ from night-flying migrants and in other contexts; plaintive ‘tyup’ or ‘tyoop’; short ‘tyip’ in flight or on take-off. For full treatment see Cramp and Perrins (1994). General Habits. Breeds in Algeria in open mountain forest at 1000–1600 m. On migration, frequents dry open country, with or without trees; short grassland, semidesert, cereal fields, steppe and open bush, and woodland edges. Winters in savannas and cultivated country, mainly in upland areas; in Guinea, on broken ground on Mt Nimba at 1250–1400 m, in Nigeria on open grassy plateaux with bare rock patches, in Eritrea and Ethiopia above 900 m, in grassland, acacia savanna and cleared forest. Rather quiet and secretive. Occurs often singly, but in parties and small flocks on passage and in winter. Hundreds of newly arrived migrants visit drinking places in the Sudan Red Sea Hills in Sept, associating with Cretzschmar’s Buntings (G. Nikolaus, pers. comm., Prendergast 1985). In Sierra Leone, occurs in flocks together with Cinnamon-breasted Rock Bunting E. tahapisi. Feeds mainly on ground, with hopping gait. In breeding season forages in trees, sometimes on high branches; picks off caterpillars while hovering and may catch insects in flight. Occasionally sings on spring passage in N Africa and Middle East. Autumn migration inconspicuous in NW Africa: late Sept to mid-Oct on W Moroccan coast, Sept–Oct in Algeria, Tunisia and Libya, late Sept–Nov or Dec in Mauritania, Senegal, Gambia and Mali. In east of range, small numbers are regular in E Egypt in mid-Aug to midOct; strong passage Sudan and Eritrea, late Aug–Oct. Migrates at night, when ‘tsjo’ flight calls frequently heard on Sudan coast. Arrives in wintering areas in Darfur (W Sudan) and central Ethiopia from early Oct, Nigeria from Nov. Departure from Ethiopia takes place in Mar, complete by end of month. Migration through N Africa and Saharan oases occurs in late Mar–early May; more pronounced in spring than autumn. Strong return passage through Egypt and Middle East, mainly mid-Mar to early May. A bird ringed Finland recovered N Morocco (Mar), 1 ringed Belgium recovered N Morocco (Oct), 1 ringed Switzerland recovered Tunisia (Apr) (Zink 1985). Food. Invertebrates (more insectivorous than most buntings); also seeds, especially in non-breeding season. In Europe, mainly ants, beetles, grasshoppers and caterpillars;

593

594

EMBERIZIDAE

young fed especially with caterpillars (Cramp and Perrins 1994). Breeding Habits. Mainly monogamous. Pairs only for 1 breeding attempt. Pairs widely separated or in small, loose ‘colonies’. Territories not strictly defended; song-posts often only 25–50 m apart. Breeding birds collect food up to 200 m from nests, but usually closer (Cramp and Perrins 1994). Y arrives and begins singing; joined by X after a few days. During pair-bonding, Y may pursue X in fast twisting flight, giving repeated dry ‘tip tip tip’ call. Later, pair often feeds and stays close together when X not on nest. Y sings from treetop, wire, low bush or boulder; usually with head slightly above horizontal, neck somewhat extended, wings drooping and rump feathers raised, but may sing in resting attitude. Y also sings in displayflight, rising steeply from song-post with undulating flight and descending abruptly. Display-flight given when X incubating, presumably in territory defence (Cramp and Perins 1994). NEST: cup of stalks, stems, roots and leaves lined with fine grasses, rootlets and hair. On ground, often in rather deep depression; ext. diam. 90–140 (110–120), int. diam.

60–70 (65); cup depth 30–40 (34); ext. height c. 75. Built by X only, sometimes accompanied by Y; in 2–3 days, taking material from within 50 m; may excavate depression with feet. EGGS: 3–6 (usually 4–5); glossy; bluish, greyish or pinkish, marked sparsely with blackish speckles, blotches and scrawls, sometimes forming ring at broad end, and with greyish undermarkings. SIZE: (n ¼ 637 Europe) 17–22  14–17 (199  154). LAYING DATES: Algeria, (newly fledged young mid June). INCUBATION: by X only; period 11–12 (13) days, starting with last or penultimate egg. DEVELOPMENT AND CARE OF YOUNG: fed and cared for by both parents. Brooded by X for 3–4 days. Fledging period 12–13 days; young may leave nest before able to fly. Fed by parents for another 4–5 days; independent 8–12 days after leaving nest. BREEDING SUCCESS/SURVIVAL: in Russia, of 155 eggs in 33 clutches, 65% hatched, 51% produced fledged young (Cramp and Perrins 1994). Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Small, B. (1992).

Plate 36

Emberiza caesia Cretzschmar. Cretzschmar’s Bunting. Bruant cendrillard.

(Opp. p. 539)

Emberiza caesia Cretzschmar, 1828. In Ru ¨ ppell, Atlas Reise no ¨rdlichen Afrika, Vo ¨gel, p. 17, pl. 10, fig. b; Kurgos Island, Nile River, Berber District, Sudan. Range and Status. Greece, Cyprus, and W and S Turkey to N Israel and NW Jordon; migrates to NE Africa. Winters Sudan (west to Darfur, south to Sennar, mainly Nile system and Red Sea Prov.) and lowlands of Eritrea, including offshore islands; locally frequent to common. Frequent on both passages in Egypt, mainly in Nile Delta and Valley and Eastern Desert; no recent confirmation of wintering as claimed by Meinertzhagen (1930). Vagrant E Libya (El Bardia, Apr), S Algeria (once), Chad (once, Abe´che´, Jan; Salvan 1963) and Djibouti (once, Tadjoura, Mar). Description. ADULT Y (breeding): forehead to hindneck, sides of neck, upper cheeks and ear-coverts bluish grey; lores and fore supercilium cinnamon-rufous; eye-ring creamy white. Mantle and scapulars tawny brown or rufous-cinnamon, lower mantle and scapulars broadly streaked blackish; back to uppertail-coverts brighter rufous-cinnamon. Tail blackish brown; T1 fringed cinnamon; tip of inner web of T4 fringed buffy white; inner web of T5–T6 with white distal wedge, occupying about a quarter of length of T5, one-third of T6. Broad cinnamon-rufous stripe across lower cheek to just behind lower ear-coverts, bordered by narrow bluish grey malar stripe connecting with grey lower side of neck. Chin and throat rufous-cinnamon, sharply demarcated from malar stripe and from broad bluish grey band across upper breast. Rest of underparts deep rufous-chestnut, slightly paler on sides, flanks, thighs and undertail-coverts. Flight feathers, primary coverts and alula blackish brown; primaries edged buff, secondaries more broadly with cinnamon-buff, primary coverts and small alula feather narrowly fringed buff. Tertials and greater

Emberiza caesia

and median coverts black; tips and outer webs of tertials, edges and tips of greater coverts and tips of median coverts broadly fringed

594

EMBERIZIDAE

young fed especially with caterpillars (Cramp and Perrins 1994). Breeding Habits. Mainly monogamous. Pairs only for 1 breeding attempt. Pairs widely separated or in small, loose ‘colonies’. Territories not strictly defended; song-posts often only 25–50 m apart. Breeding birds collect food up to 200 m from nests, but usually closer (Cramp and Perrins 1994). Y arrives and begins singing; joined by X after a few days. During pair-bonding, Y may pursue X in fast twisting flight, giving repeated dry ‘tip tip tip’ call. Later, pair often feeds and stays close together when X not on nest. Y sings from treetop, wire, low bush or boulder; usually with head slightly above horizontal, neck somewhat extended, wings drooping and rump feathers raised, but may sing in resting attitude. Y also sings in displayflight, rising steeply from song-post with undulating flight and descending abruptly. Display-flight given when X incubating, presumably in territory defence (Cramp and Perins 1994). NEST: cup of stalks, stems, roots and leaves lined with fine grasses, rootlets and hair. On ground, often in rather deep depression; ext. diam. 90–140 (110–120), int. diam.

60–70 (65); cup depth 30–40 (34); ext. height c. 75. Built by X only, sometimes accompanied by Y; in 2–3 days, taking material from within 50 m; may excavate depression with feet. EGGS: 3–6 (usually 4–5); glossy; bluish, greyish or pinkish, marked sparsely with blackish speckles, blotches and scrawls, sometimes forming ring at broad end, and with greyish undermarkings. SIZE: (n ¼ 637 Europe) 17–22  14–17 (199  154). LAYING DATES: Algeria, (newly fledged young mid June). INCUBATION: by X only; period 11–12 (13) days, starting with last or penultimate egg. DEVELOPMENT AND CARE OF YOUNG: fed and cared for by both parents. Brooded by X for 3–4 days. Fledging period 12–13 days; young may leave nest before able to fly. Fed by parents for another 4–5 days; independent 8–12 days after leaving nest. BREEDING SUCCESS/SURVIVAL: in Russia, of 155 eggs in 33 clutches, 65% hatched, 51% produced fledged young (Cramp and Perrins 1994). Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Small, B. (1992).

Plate 36

Emberiza caesia Cretzschmar. Cretzschmar’s Bunting. Bruant cendrillard.

(Opp. p. 539)

Emberiza caesia Cretzschmar, 1828. In Ru ¨ ppell, Atlas Reise no ¨rdlichen Afrika, Vo ¨gel, p. 17, pl. 10, fig. b; Kurgos Island, Nile River, Berber District, Sudan. Range and Status. Greece, Cyprus, and W and S Turkey to N Israel and NW Jordon; migrates to NE Africa. Winters Sudan (west to Darfur, south to Sennar, mainly Nile system and Red Sea Prov.) and lowlands of Eritrea, including offshore islands; locally frequent to common. Frequent on both passages in Egypt, mainly in Nile Delta and Valley and Eastern Desert; no recent confirmation of wintering as claimed by Meinertzhagen (1930). Vagrant E Libya (El Bardia, Apr), S Algeria (once), Chad (once, Abe´che´, Jan; Salvan 1963) and Djibouti (once, Tadjoura, Mar). Description. ADULT Y (breeding): forehead to hindneck, sides of neck, upper cheeks and ear-coverts bluish grey; lores and fore supercilium cinnamon-rufous; eye-ring creamy white. Mantle and scapulars tawny brown or rufous-cinnamon, lower mantle and scapulars broadly streaked blackish; back to uppertail-coverts brighter rufous-cinnamon. Tail blackish brown; T1 fringed cinnamon; tip of inner web of T4 fringed buffy white; inner web of T5–T6 with white distal wedge, occupying about a quarter of length of T5, one-third of T6. Broad cinnamon-rufous stripe across lower cheek to just behind lower ear-coverts, bordered by narrow bluish grey malar stripe connecting with grey lower side of neck. Chin and throat rufous-cinnamon, sharply demarcated from malar stripe and from broad bluish grey band across upper breast. Rest of underparts deep rufous-chestnut, slightly paler on sides, flanks, thighs and undertail-coverts. Flight feathers, primary coverts and alula blackish brown; primaries edged buff, secondaries more broadly with cinnamon-buff, primary coverts and small alula feather narrowly fringed buff. Tertials and greater

Emberiza caesia

and median coverts black; tips and outer webs of tertials, edges and tips of greater coverts and tips of median coverts broadly fringed

Emberiza rustica cinnamon or rufous brown. Lesser coverts greyish buff. Underwing-coverts and axillaries greyish white. Bill pinkish horn, culmen horn brown to dark slate grey; eyes brown or blackish; legs pinkish orange or pale flesh brown. ADULT Y (nonbreeding): like breeding Y but grey of head and neck tinged brown (less blue); crown feathers often with fine dusky shaft streaks. ADULT X: differs from ad. Y in having grey of head and upper breast band tinged brown, especially on ear-coverts; forehead to hindneck narrowly streaked blackish; malar stripe more prominent and flecked blackish; breast band with short narrow blackish streaks, less sharply demarcated; lower cheek stripe, throat patch and underparts paler rufous-cinnamon. Complete moult July–Aug; partial moult of head and body in Africa. SIZE (10 YY, 10 XX): wing, Y 81–89 (847), X 77–83 (801); tail, Y 62–70 (665), X 60–67 (626); bill, Y 125–135 (128), X 12–13 (125); tarsus, Y 18–20 (189), X 175–19 (185). WEIGHT: Sudan coast, Sept–Oct, Y (n ¼ 56) 145–225 (189), X (n ¼ 59) 15–21 (186); Greece, Turkey and Israel, Mar–July, Y (n ¼ 8) 198–23 (212), X (n ¼ 2) 198, 222. IMMATURE: 1st winter similar to ad. X, but top of head brown or buffy brown, more strongly streaked; lower cheek stripe, chin and throat buff or cinnamon-buff; sides of neck and breast band with only a tinge of grey; breast more boldly and extensively streaked, and a few streaks on flanks; underparts paler cinnamonbuff; retained juv. outer greater coverts fringed pale buff or whitish.

Field Characters. Length 15–16 cm. Shares plain head, whitish eye-ring and pink bill with Ortolan Bunting E. hortulana, but head and breast bluish grey without olive tones, throat and sub-moustachial stripe orange-chestnut. X duller, with streaked head, like X Ortolan, but throat and sub-moustachial stripe rufous-buff, not yellow, head greyer; in all plumages more rufous above, including rump, and deeper rufous below. Juv. very similar to juv. Ortolan but overall more rufous, head greyer, throat tinged cinnamon, not yellowish. In all plumages lower wing-bar more cinnamon (i.e. less prominent) than Ortolan. Voice. Tape-recorded (62–73, 92, 110–113, B, MIL). Song of 3–4 notes, not unlike Ortolan but with single longer

595

note at end; some songs drier and buzzier than Ortolan, ‘dzee-dzee-dzee-dzreee’, others ringing and sweet, ‘tooytooy-tooy-toooo’, ‘wee-hee-hee-heeee’ or more languid ‘wooey, wooey, wheeee’; lasts 1–15 s, given c. 8 times per min. 1 Y may have several song types. Contact call a hard, sharp ‘jit’ or ‘jititit’; other calls more liquid, ‘tyeep’ or ‘tyup’; dry ‘plet’ given in flight.

General Habits. On passage and in winter occupies low, dry habitats: steppe and desert borders; bare rock desert; Acacia savanna; short grass coastal plains; dry hillsides, bare or with sparse vegetation; rocky wadis with scrub; cultivated areas and gardens within arid regions. Winters mainly at lower altitude than Ortolan Bunting; in Eritrea below 600 m (Smith 1957). Often in small parties. May congregate in larger flocks, e.g. at drinking spots (Sudan coast, Sept) or on ripening cereal fields (Eritrea, Feb). Tame and approachable. Highly terrestrial; perches on bushes only when alarmed; feeds almost exclusively on ground, often among semi-arid herbs and scrub. On passage, forages with Ortolans on rocky or scrubby slopes and in stubble fields and desert edge cultivation. Migrates on broad front across and around E Mediterranean in both autumn and spring. Moves mainly at night. Fairly common in Egypt, where main autumn passage is mid Aug to late Sept (latest early Nov; 21, Zaranik, on one day). Arrives Sudan from early Sept (mainly mid Sept–Oct), departs in Mar. Spring passage through Egypt is mainly mid-Mar to late Apr (occasional from mid Feb); through Israel mainly in Mar.

Food. Seeds and small invertebrates.

Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Small, B. (1992), Smith, K.D. (1957).

Emberiza rustica Pallas. Rustic Bunting. Bruant rustique.

Plate 36

Emberiza rustica Pallas, 1776. Reise d. versch. Prov. Russ. Reichs, 3, p. 698; Dauria.

(Opp. p. 539)

N Eurasia, from Scandinavia to E Siberia; winters E China to Japan, and a few in central Asia. Vagrant (E. r. rustica) to W and S Europe, Middle East and Egypt (1 collected Bahig, Oct 1971). Slightly smaller and shorter-tailed than Reed Bunting E. schoeniclus. Rufous brown above, with dark streaked mantle and mottled rufous rump; white below, with rufous streaked sides and flanks. Shows broad pale supercilium behind eye, 2 narrow white wing-bars and conspicuous white tail edges. Spring Y has black crown and black face, and contrasting white supercilium, white moustache and white throat; plain rufous hindneck and rufous chest band. X and autumn/winter Y have complex but more subdued head pattern: dark brown crown sides and narrow central stripe, pale brown face bordered behind by blackish stripes, pale buff supercilium, moustachial stripe and throat; rufous breast mottling and flank streaks paler than in spring Y. Length 14–15 cm; wing, Y (n ¼ 10) 76–81 (779), X (n ¼ 10) 73–78 (745); weight, N Russia, Y (n ¼ 6) 181–208 (192), X (n ¼ 8) 171–199 (183). Commonest calls a high-pitched ‘zit’ or ‘tsip’ and a disyllabic ‘tit-tit’. Inhabits moist, wooded lowlands. In non-breeding season in E Asia, cool woodlands, clearings and cultivation, often in marshy places or along river banks. Forms flocks, often with other buntings. Forages on ground for seeds among weeds and grass and stubble, and feeds on berries in low bushes.

Emberiza rustica cinnamon or rufous brown. Lesser coverts greyish buff. Underwing-coverts and axillaries greyish white. Bill pinkish horn, culmen horn brown to dark slate grey; eyes brown or blackish; legs pinkish orange or pale flesh brown. ADULT Y (nonbreeding): like breeding Y but grey of head and neck tinged brown (less blue); crown feathers often with fine dusky shaft streaks. ADULT X: differs from ad. Y in having grey of head and upper breast band tinged brown, especially on ear-coverts; forehead to hindneck narrowly streaked blackish; malar stripe more prominent and flecked blackish; breast band with short narrow blackish streaks, less sharply demarcated; lower cheek stripe, throat patch and underparts paler rufous-cinnamon. Complete moult July–Aug; partial moult of head and body in Africa. SIZE (10 YY, 10 XX): wing, Y 81–89 (847), X 77–83 (801); tail, Y 62–70 (665), X 60–67 (626); bill, Y 125–135 (128), X 12–13 (125); tarsus, Y 18–20 (189), X 175–19 (185). WEIGHT: Sudan coast, Sept–Oct, Y (n ¼ 56) 145–225 (189), X (n ¼ 59) 15–21 (186); Greece, Turkey and Israel, Mar–July, Y (n ¼ 8) 198–23 (212), X (n ¼ 2) 198, 222. IMMATURE: 1st winter similar to ad. X, but top of head brown or buffy brown, more strongly streaked; lower cheek stripe, chin and throat buff or cinnamon-buff; sides of neck and breast band with only a tinge of grey; breast more boldly and extensively streaked, and a few streaks on flanks; underparts paler cinnamonbuff; retained juv. outer greater coverts fringed pale buff or whitish.

Field Characters. Length 15–16 cm. Shares plain head, whitish eye-ring and pink bill with Ortolan Bunting E. hortulana, but head and breast bluish grey without olive tones, throat and sub-moustachial stripe orange-chestnut. X duller, with streaked head, like X Ortolan, but throat and sub-moustachial stripe rufous-buff, not yellow, head greyer; in all plumages more rufous above, including rump, and deeper rufous below. Juv. very similar to juv. Ortolan but overall more rufous, head greyer, throat tinged cinnamon, not yellowish. In all plumages lower wing-bar more cinnamon (i.e. less prominent) than Ortolan. Voice. Tape-recorded (62–73, 92, 110–113, B, MIL). Song of 3–4 notes, not unlike Ortolan but with single longer

595

note at end; some songs drier and buzzier than Ortolan, ‘dzee-dzee-dzee-dzreee’, others ringing and sweet, ‘tooytooy-tooy-toooo’, ‘wee-hee-hee-heeee’ or more languid ‘wooey, wooey, wheeee’; lasts 1–15 s, given c. 8 times per min. 1 Y may have several song types. Contact call a hard, sharp ‘jit’ or ‘jititit’; other calls more liquid, ‘tyeep’ or ‘tyup’; dry ‘plet’ given in flight.

General Habits. On passage and in winter occupies low, dry habitats: steppe and desert borders; bare rock desert; Acacia savanna; short grass coastal plains; dry hillsides, bare or with sparse vegetation; rocky wadis with scrub; cultivated areas and gardens within arid regions. Winters mainly at lower altitude than Ortolan Bunting; in Eritrea below 600 m (Smith 1957). Often in small parties. May congregate in larger flocks, e.g. at drinking spots (Sudan coast, Sept) or on ripening cereal fields (Eritrea, Feb). Tame and approachable. Highly terrestrial; perches on bushes only when alarmed; feeds almost exclusively on ground, often among semi-arid herbs and scrub. On passage, forages with Ortolans on rocky or scrubby slopes and in stubble fields and desert edge cultivation. Migrates on broad front across and around E Mediterranean in both autumn and spring. Moves mainly at night. Fairly common in Egypt, where main autumn passage is mid Aug to late Sept (latest early Nov; 21, Zaranik, on one day). Arrives Sudan from early Sept (mainly mid Sept–Oct), departs in Mar. Spring passage through Egypt is mainly mid-Mar to late Apr (occasional from mid Feb); through Israel mainly in Mar.

Food. Seeds and small invertebrates.

Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Small, B. (1992), Smith, K.D. (1957).

Emberiza rustica Pallas. Rustic Bunting. Bruant rustique.

Plate 36

Emberiza rustica Pallas, 1776. Reise d. versch. Prov. Russ. Reichs, 3, p. 698; Dauria.

(Opp. p. 539)

N Eurasia, from Scandinavia to E Siberia; winters E China to Japan, and a few in central Asia. Vagrant (E. r. rustica) to W and S Europe, Middle East and Egypt (1 collected Bahig, Oct 1971). Slightly smaller and shorter-tailed than Reed Bunting E. schoeniclus. Rufous brown above, with dark streaked mantle and mottled rufous rump; white below, with rufous streaked sides and flanks. Shows broad pale supercilium behind eye, 2 narrow white wing-bars and conspicuous white tail edges. Spring Y has black crown and black face, and contrasting white supercilium, white moustache and white throat; plain rufous hindneck and rufous chest band. X and autumn/winter Y have complex but more subdued head pattern: dark brown crown sides and narrow central stripe, pale brown face bordered behind by blackish stripes, pale buff supercilium, moustachial stripe and throat; rufous breast mottling and flank streaks paler than in spring Y. Length 14–15 cm; wing, Y (n ¼ 10) 76–81 (779), X (n ¼ 10) 73–78 (745); weight, N Russia, Y (n ¼ 6) 181–208 (192), X (n ¼ 8) 171–199 (183). Commonest calls a high-pitched ‘zit’ or ‘tsip’ and a disyllabic ‘tit-tit’. Inhabits moist, wooded lowlands. In non-breeding season in E Asia, cool woodlands, clearings and cultivation, often in marshy places or along river banks. Forms flocks, often with other buntings. Forages on ground for seeds among weeds and grass and stubble, and feeds on berries in low bushes.

596

EMBERIZIDAE

Plate 36

Emberiza pusilla Pallas. Little Bunting. Bruant nain.

(Opp. p. 539)

Emberiza pusilla Pallas, 1776. Reise d. versch. Prov. Russ. Reichs, 3, p. 647; Daurian Alps. N Eurasia, from N Norway to E Siberia; winters SE Asia. Vagrant to W and S Europe, SW Asia, Morocco (1, Charaf, Agadir, Oct 1994) and Egypt (1, Agamy, Nov 1980). A small bunting with a small, sharply pointed bill and shortish, narrowly white-edged tail. Well-patterned head shows rufous face, narrow white eye-ring, blackish or streaky brown crown with pale central stripe, broad rufous or creamy supercilium, dark line behind eye and around ear-coverts, and dark malar stripe. Buff to grey-brown above with dark streaks; white below with short dark streaks on breast and flanks. 2 inconspicuous narrow wing-bars, the front one whitish. Length 12–14 cm; wing, Y (n ¼ 10) 70–76 (726), X (n ¼ 10) 67–70 (688); weight, Netherlands, autumn, unsexed (n ¼ 7) 127–165 (141). Calls include 2 short flat monosyllables given by migrants, and a sharp clicking ‘pwick’ or ‘tik’. In Asia in non-breeding season, inhabits forest fringes, bushy hillsides, crops and stubble, marshes and river banks. Joins winter flocks feeding on ground on grass and cereal seeds; also eats invertebrates. Tame and approachable but can vanish among ground cover. Tends to skulk rather than fly and quickly dives back to ground when flushed.

Plate 36

Emberiza schoeniclus Linnaeus. Common Reed Bunting. Bruant des roseaux.

(Opp. p. 539)

Fringilla schoeniclus Linnaeus, 1758. Syst. Nat., ed. 10, p. 182; Sweden. Range and Status. Eurasia, north to Scandinavia and central Siberia, east to Japan and Kamchatka, south to NW China, Iran, Turkey, Iberia and Morocco. Partial migrant; winters in W and S part of range, also Mediterranean area and S and E Asia. Breeds or has bred at up to 4 sites in Morocco; a few winter in NW Africa. Morocco, occasional pairs breed some years in marshes at Bas-Loukkos; probably breeds also at Merja Zerga and Sidi-Bou-Rhaba; outside breeding season rare to uncommon in lowlands throughout, south to Haut-Atlas; several records in most years, of single birds or flocks of up to 12, mainly in Nov–Mar in mouths of Moulouya, Souss and Massa, also at Douyiet, Marja Barga, Oued Smir, Sidi Bettache, Merja Halloufa, palm groves at Oud Tensift and salt-pans at Lixus. N Algeria and Tunisia, irregular in non-breeding season south to Hauts-Plateaux, Atlas Saharien and Gulf of Gabe`s; has reached edge of Sahara at Loghouat and Beni Abbe`s. Scarce and local NW Libya, perhaps regular in coastal Tripolitania; vagrant N Egypt (4 records, Nov–Mar). Description. E. s. schoeniclus (Linnaeus): W and N Europe; some winter in Morocco to Tunisia. ADULT Y: forehead to nape, earcoverts and cheeks black; collar around hindneck and sides of neck white, connecting with broad white sub-moustachial stripe. Upper mantle below white collar grey-brown spotted with black; rest of mantle and scapulars ochre and cinnamon-rufous, broadly streaked black; back, rump and uppertail-coverts olive-brown or grey-brown, back with narrow blackish streaks. Tail blackish; T1 broadly edged cinnamon; inner web of T5 with white distal wedge, extending along shaft for about 60% of feather length; T6 with inner web (apart from base of inner border) and basal twothirds of outer web white. Chin and throat to bib on centre of upper breast black; rest of underparts greyish white with a few long brown and blackish streaks on flanks. Upperwing feathers blackish brown; primaries and secondaries edged cinnamonrufous; outer webs and tips of tertials and greater coverts broadly bordered cinnamon-rufous; primary coverts and alula narrowly fringed cinamon-rufous; lesser coverts, and broad tips to median coverts brighter rufous-chestnut. Underwing-coverts and axillaries

Emberiza schoeniclus

white. In winter, black and white head areas and bib partly obscured by brown feather tips, and breast mottled brownish. In fresh autumn plumage, collar may be fully hidden, top of head mottled buff-brown and head pattern more complex, the buffish superciliary stripe, cental ear-coverts, sub-moustachial stripe and lower side of neck contrasting with dark ear-covert borders and distinct moustachial stripe. Bill blackish when breeding; in winter, upper mandible dark grey, lower mandible paler horngrey. Eyes dark brown; legs pinkish brown, flesh-grey or dark brown. ADULT X: forehead to nape pale olive-brown, narrowly streaked blackish, sides of crown and nape brighter, more rufous; hindneck olive-brown and rufous-brown; upper mantle greyish olive, mottled with dusky; rest of upperparts as ad. Y. Tail as ad. Y. Supercilium pale buff, broad behind eye, narrower in front.

596

EMBERIZIDAE

Plate 36

Emberiza pusilla Pallas. Little Bunting. Bruant nain.

(Opp. p. 539)

Emberiza pusilla Pallas, 1776. Reise d. versch. Prov. Russ. Reichs, 3, p. 647; Daurian Alps. N Eurasia, from N Norway to E Siberia; winters SE Asia. Vagrant to W and S Europe, SW Asia, Morocco (1, Charaf, Agadir, Oct 1994) and Egypt (1, Agamy, Nov 1980). A small bunting with a small, sharply pointed bill and shortish, narrowly white-edged tail. Well-patterned head shows rufous face, narrow white eye-ring, blackish or streaky brown crown with pale central stripe, broad rufous or creamy supercilium, dark line behind eye and around ear-coverts, and dark malar stripe. Buff to grey-brown above with dark streaks; white below with short dark streaks on breast and flanks. 2 inconspicuous narrow wing-bars, the front one whitish. Length 12–14 cm; wing, Y (n ¼ 10) 70–76 (726), X (n ¼ 10) 67–70 (688); weight, Netherlands, autumn, unsexed (n ¼ 7) 127–165 (141). Calls include 2 short flat monosyllables given by migrants, and a sharp clicking ‘pwick’ or ‘tik’. In Asia in non-breeding season, inhabits forest fringes, bushy hillsides, crops and stubble, marshes and river banks. Joins winter flocks feeding on ground on grass and cereal seeds; also eats invertebrates. Tame and approachable but can vanish among ground cover. Tends to skulk rather than fly and quickly dives back to ground when flushed.

Plate 36

Emberiza schoeniclus Linnaeus. Common Reed Bunting. Bruant des roseaux.

(Opp. p. 539)

Fringilla schoeniclus Linnaeus, 1758. Syst. Nat., ed. 10, p. 182; Sweden. Range and Status. Eurasia, north to Scandinavia and central Siberia, east to Japan and Kamchatka, south to NW China, Iran, Turkey, Iberia and Morocco. Partial migrant; winters in W and S part of range, also Mediterranean area and S and E Asia. Breeds or has bred at up to 4 sites in Morocco; a few winter in NW Africa. Morocco, occasional pairs breed some years in marshes at Bas-Loukkos; probably breeds also at Merja Zerga and Sidi-Bou-Rhaba; outside breeding season rare to uncommon in lowlands throughout, south to Haut-Atlas; several records in most years, of single birds or flocks of up to 12, mainly in Nov–Mar in mouths of Moulouya, Souss and Massa, also at Douyiet, Marja Barga, Oued Smir, Sidi Bettache, Merja Halloufa, palm groves at Oud Tensift and salt-pans at Lixus. N Algeria and Tunisia, irregular in non-breeding season south to Hauts-Plateaux, Atlas Saharien and Gulf of Gabe`s; has reached edge of Sahara at Loghouat and Beni Abbe`s. Scarce and local NW Libya, perhaps regular in coastal Tripolitania; vagrant N Egypt (4 records, Nov–Mar). Description. E. s. schoeniclus (Linnaeus): W and N Europe; some winter in Morocco to Tunisia. ADULT Y: forehead to nape, earcoverts and cheeks black; collar around hindneck and sides of neck white, connecting with broad white sub-moustachial stripe. Upper mantle below white collar grey-brown spotted with black; rest of mantle and scapulars ochre and cinnamon-rufous, broadly streaked black; back, rump and uppertail-coverts olive-brown or grey-brown, back with narrow blackish streaks. Tail blackish; T1 broadly edged cinnamon; inner web of T5 with white distal wedge, extending along shaft for about 60% of feather length; T6 with inner web (apart from base of inner border) and basal twothirds of outer web white. Chin and throat to bib on centre of upper breast black; rest of underparts greyish white with a few long brown and blackish streaks on flanks. Upperwing feathers blackish brown; primaries and secondaries edged cinnamonrufous; outer webs and tips of tertials and greater coverts broadly bordered cinnamon-rufous; primary coverts and alula narrowly fringed cinamon-rufous; lesser coverts, and broad tips to median coverts brighter rufous-chestnut. Underwing-coverts and axillaries

Emberiza schoeniclus

white. In winter, black and white head areas and bib partly obscured by brown feather tips, and breast mottled brownish. In fresh autumn plumage, collar may be fully hidden, top of head mottled buff-brown and head pattern more complex, the buffish superciliary stripe, cental ear-coverts, sub-moustachial stripe and lower side of neck contrasting with dark ear-covert borders and distinct moustachial stripe. Bill blackish when breeding; in winter, upper mandible dark grey, lower mandible paler horngrey. Eyes dark brown; legs pinkish brown, flesh-grey or dark brown. ADULT X: forehead to nape pale olive-brown, narrowly streaked blackish, sides of crown and nape brighter, more rufous; hindneck olive-brown and rufous-brown; upper mantle greyish olive, mottled with dusky; rest of upperparts as ad. Y. Tail as ad. Y. Supercilium pale buff, broad behind eye, narrower in front.

Emberiza schoeniclus Narrow eye-ring pale buff. Lores finely mottled dark brown and buff; broad stripe behind eye dark brown, tinged rufous; below eye to centre of ear-coverts brown and buff, bordered below by dark brown moustachial stripe from base of bill, and behind by dark bar. Broad sub-moustachial stripe pale buff, connecting with buffy white bar on lower side of neck; below this a bold black and brown streaked malar stripe, extending back and broadening across base of throat. Chin and throat pale buff; upper breast pale buff-brown, with short dark brown streaks; rest of underparts whitish or creamy, often sullied with grey, side of breast and flanks with long dark brown streaks. Wing as ad. Y. Bare parts as ad. Y (non-breeding). Complete moult July–Sept. SIZE (10 YY, 10 XX): wing, Y 76–85 (805), X 72–76 (742); tail, Y 64–72 (672), X 60–67 (628); bill, Y 115–13 (122), X 11–12 (116); tarsus, Y 195–205 (202), X 19–205 (196). Bill depth, YX (n ¼ 48) 48–57 (Cramp and Perrins 1994). WEIGHT: SE France, Oct–Feb, Y (n ¼ 217) 180–250 (212), X (n ¼ 159) 155–225 (186). IMMATURE: 1st-winter birds like ads, but tail feathers more pointed. E. s. witherbyi von Jordons: Iberia, S France; probably this race breeding Morocco; a few winter in NW Africa. Similar in colour to schoeniclus or slightly darker; bill much heavier. E. s. intermedia Degland: Corsica, Italy and Adriatic coast of Croatia. A few winter in Algeria, Tunisia and perhaps Libya and Egypt. Like witherbyi, with heavy bill and curved culmen, but slightly paler, especially on rump, flanks streaked rufous-brown rather than blackish; wing longer, Y (n ¼ 14) 80–87 (837); bill depth, YX (n ¼ 23) 69–78 (Cramp and Perrins 1994). (E. s. tschusii Reiser and Almasy: Danube valley in Bulgaria and Rumania to S Ukraine; perhaps the form occurring in Egypt. Like intermedia, but upperparts paler, with narrower black streaking. Wing, Y (n ¼ 5) 83–86 (850).)

Field Characters. Length 15–16 cm. Breeding Y distinctive, with black head, white collar and moustachial stripe; black obscured in autumn by buff feather tips, but outline still apparent. Spring X has broad black malar stripe, dark brown face, grey sides of neck. Other plumages more nondescript, but typically has dark malar stripe and triangular brown face patch with blackish border, surrounded by buff supercilium and sides of neck and whitish moustachial stripe. Scapulars and wings have rufous tone in all plumages; rump grey. Rustic Bunting E. rustica has chestnut stripes on flanks and breast, rufous rump, distinct white wing-bars and pale spot at rear of dark ear-coverts; Little Bunting E. pusilla smaller, with chestnut face patch, pale eye-ring and grey (not rufous) lesser coverts. Characteristic metallic ‘seeeu’ call a clue to bird’s presence when hidden in vegetation. Voice. Tape-recorded (62–73, 76–89, 105, 110–113, B, C). Song high-pitched, ringing but not musical; at slower speeds, a series of single notes and short trills given in hesitant, disjointed manner, ‘tirrrup . . . tsit . . . tsweet . . . tsweet . . . tirrrup . . . tsit . . . tsweet’; faster songs have trills of varying speeds, ‘tsee-tsee-tway-tway-tsitsitsitsi’. Common call a penetrating descending ‘seeeu’; low buzzy ‘bzree’ or ‘jup’ given in flight and by migrants. For further songs and calls see Cramp and Perrins (1994). General Habits. In breeding season, inhabits tall herbage and small shrubs in marshy areas by fresh or brackish water, usually in valleys and lowlands; especially mixed reed and willow. On passage and in winter, occurs on lakeshores and marshes, but also well away from water, in

cultivated fields, weedy areas, woodland clearings and marram grass on sand dunes. Not shy; easy to observe. Regularly forms groups or small feeding flocks outside breeding season. Often associates with other seedeaters. Roosts communally in thick vegetation, such as reeds and growth fringing lake shores. Forages mainly on ground, among sedges, rushes and reeds, or in pasture and marshy grasslands; also from reeds and tall grass stems, and low down in waterside bushes and trees. Quite agile when foraging. Often takes flying insects. Commonly perches rather upright on prominent bush or stem. Frequently flicks and spreads tail. Hops on ground, and creeps when feeding. Populations in N and E Europe and most of Asia are migratory; those of Britain, S Europe and S central Asia mainly sedentary. Scandinavian and central European birds move W to SSW, mainly in late Sept–early Nov. Many winter south of breeding range in Mediterranean area, mid Nov–Feb. Regular on passage at Gibraltar, and small numbers widespread in winter in NW Africa, in Mediterranean coastal areas and south to Atlas range, occasionally reaching Saharan borders: mainly nominate schoeniclus but including some thick-billed witherbyi and intermedia. Spring return migration in Europe is mainly in mid Feb–Apr. Food. In breeding season, mainly invertebrates, especially Diptera, caterpillars, beetles, spiders, Odonata and Orthoptera. In non-breeding season, from late July, mainly seeds of grasses, knotgrass and goosegrass; also other plant material. Eats invertebrates opportunistically, including spiders, springtails, bugs, flies and beetles, and many larvae and pupae. Breeding Habits. Very well known in Europe. Monogamous, but extra-pair paternity common; occasionally polygynous. Solitary and territorial. Territory used for pair formation and nesting. Size variable; in Dutch study,

597

598

EMBERIZIDAE

96–3500 m2 (av. 1595 m2); neighbouring territories overlap in densely populated areas. Breeding density in Europe high (140–180 per km2) in suitable marshy areas, much lower (usually less than 15 per km2) in areas of mixed habitat (Cramp and Perrins 1994). Pairing takes place in Y’s territory after break-up of winter flocks. Y is then frequently in conflict with rivals; gives threat display, with body and head held horizontal, and aggressive calls; often engages in face to face fights. Y sings from bush or reed stem, in upright relaxed posture with head and rump feathers ruffled (A). Song most intense when Y unpaired or when X incubating (Cramp and Perrins 1994). 2, perhaps 3, broods. NEST: cup of stems and blades of sedges, grasses, waterside plants, a few twigs; lined with moss, rootlets, sometimes hair or feathers; ext. diam. c. 80–90, int. diam. c. 50, cup depth c. 45; usually well hidden on ground or sedge tussocks, by water; or up to 4 m high in e.g. Salix bushes; built by X in 1–5 days, with material collected close by. EGGS: 3–7 (usually 4–5); slightly glossy, pale purplish, lilac grey or olive-brown, with scrawls, spots and blotches

of purplish or brownish black, irregularly distributed or concentrated at each end. SIZE (E. s. schoeniclus, n ¼ 859) 17–23  14–16 (195  146). LAYING DATES: no African data. NE Spain, late Apr–Aug. INCUBATION: by X only. Period: 12–15 (13) days. DEVELOPMENT AND CARE OF YOUNG: fed and cared for by both parents; brooded by X for first few days. Fledging period 10–12 days; young leave nest 3–5 days before able to fly. BREEDING SUCCESS/SURVIVAL: in European studies (E. s. schoeniclus), many nests lost to predators. In a 6 year study in England, only 20% of nests produced fledged young (16 fledged young per breeding pair); of 1846 eggs laid, 67% hatched, and about 30% produced flying young (Cramp and Perrins 1994).

Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Prs-Jones, R. (1984).

Plate 36

Emberiza melanocephala Scopoli. Black-headed Bunting. Bruant melanocephale.

(Opp. p. 539)

Emberiza melanocephala Scopoli, 1769. Annus I Historica-Nat., p. 142; Carnicola. Italy, Balkans, Turkey, Levant, S Russia and W Caspian to Iran; winters India. Vagrant to Morocco (once Tangier), Algeria (once Djanet, Aug 1960), Tunisia (once Sousse, Aug 1975) and Egypt (twice, Port Said, Mar 1946 and Suez, May 1982). A large bunting with rather long grey bill and long legs, separated from other buntings in Africa by combination of pale unstreaked underparts and no white in outer tail. Breeding Y has black head, chestnut back, and bright yellow collar and underparts (nape to mantle masked with buff when freshly moulted in winter). X rather featureless, with faintly streaked dull brown head and upperparts and yellow tinged underparts with buffy breast; lacks supercilium but may show thin malar stripe, and sometimes dark greyish crown and ear-coverts. Autumn Y like X, but with blacker lores and ear-coverts and yellower underparts. 1st-winter like X, but with finely streaked breast and flanks, whiter fringes to wing-coverts and no yellow below. Similar Cinereous Bunting E. cineracea semenowi has white outer tail, pale eye-ring and greyer underparts. Length 16–18 cm; wing, Y (n ¼ 10) 94–100 (971), X (n ¼ 10) 86–92 (893); weight, breeding area, Y (n ¼ 29) 25–33 (298), X (n ¼ 18) 23–32 (266). Calls include ‘cheuh’ (like House Sparrow Passer domesticus), and in flight a metallic ‘pchlu’ or ‘plut’ like Ortolan E. hortulana. Frequents open areas with scattered bushes and trees, orchards and olive groves. Often perches on commanding features. Forms flocks in Asia in non-breeding season. Feeds on ground on grass seeds and cereals or on berries in shrubs and low trees.

Genus Miliaria Brehm

Single species of large, W Palearctic bunting, formerly placed in Emberiza; but Y 20% heavier than X; plumage nondescript, sexes alike; cutting edges of mandibles curved and of lower mandible angled (A); and has complete post-juv. moult. Palate knobbed (as in Emberiza).

598

EMBERIZIDAE

96–3500 m2 (av. 1595 m2); neighbouring territories overlap in densely populated areas. Breeding density in Europe high (140–180 per km2) in suitable marshy areas, much lower (usually less than 15 per km2) in areas of mixed habitat (Cramp and Perrins 1994). Pairing takes place in Y’s territory after break-up of winter flocks. Y is then frequently in conflict with rivals; gives threat display, with body and head held horizontal, and aggressive calls; often engages in face to face fights. Y sings from bush or reed stem, in upright relaxed posture with head and rump feathers ruffled (A). Song most intense when Y unpaired or when X incubating (Cramp and Perrins 1994). 2, perhaps 3, broods. NEST: cup of stems and blades of sedges, grasses, waterside plants, a few twigs; lined with moss, rootlets, sometimes hair or feathers; ext. diam. c. 80–90, int. diam. c. 50, cup depth c. 45; usually well hidden on ground or sedge tussocks, by water; or up to 4 m high in e.g. Salix bushes; built by X in 1–5 days, with material collected close by. EGGS: 3–7 (usually 4–5); slightly glossy, pale purplish, lilac grey or olive-brown, with scrawls, spots and blotches

of purplish or brownish black, irregularly distributed or concentrated at each end. SIZE (E. s. schoeniclus, n ¼ 859) 17–23  14–16 (195  146). LAYING DATES: no African data. NE Spain, late Apr–Aug. INCUBATION: by X only. Period: 12–15 (13) days. DEVELOPMENT AND CARE OF YOUNG: fed and cared for by both parents; brooded by X for first few days. Fledging period 10–12 days; young leave nest 3–5 days before able to fly. BREEDING SUCCESS/SURVIVAL: in European studies (E. s. schoeniclus), many nests lost to predators. In a 6 year study in England, only 20% of nests produced fledged young (16 fledged young per breeding pair); of 1846 eggs laid, 67% hatched, and about 30% produced flying young (Cramp and Perrins 1994).

Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Prs-Jones, R. (1984).

Plate 36

Emberiza melanocephala Scopoli. Black-headed Bunting. Bruant melanocephale.

(Opp. p. 539)

Emberiza melanocephala Scopoli, 1769. Annus I Historica-Nat., p. 142; Carnicola. Italy, Balkans, Turkey, Levant, S Russia and W Caspian to Iran; winters India. Vagrant to Morocco (once Tangier), Algeria (once Djanet, Aug 1960), Tunisia (once Sousse, Aug 1975) and Egypt (twice, Port Said, Mar 1946 and Suez, May 1982). A large bunting with rather long grey bill and long legs, separated from other buntings in Africa by combination of pale unstreaked underparts and no white in outer tail. Breeding Y has black head, chestnut back, and bright yellow collar and underparts (nape to mantle masked with buff when freshly moulted in winter). X rather featureless, with faintly streaked dull brown head and upperparts and yellow tinged underparts with buffy breast; lacks supercilium but may show thin malar stripe, and sometimes dark greyish crown and ear-coverts. Autumn Y like X, but with blacker lores and ear-coverts and yellower underparts. 1st-winter like X, but with finely streaked breast and flanks, whiter fringes to wing-coverts and no yellow below. Similar Cinereous Bunting E. cineracea semenowi has white outer tail, pale eye-ring and greyer underparts. Length 16–18 cm; wing, Y (n ¼ 10) 94–100 (971), X (n ¼ 10) 86–92 (893); weight, breeding area, Y (n ¼ 29) 25–33 (298), X (n ¼ 18) 23–32 (266). Calls include ‘cheuh’ (like House Sparrow Passer domesticus), and in flight a metallic ‘pchlu’ or ‘plut’ like Ortolan E. hortulana. Frequents open areas with scattered bushes and trees, orchards and olive groves. Often perches on commanding features. Forms flocks in Asia in non-breeding season. Feeds on ground on grass seeds and cereals or on berries in shrubs and low trees.

Genus Miliaria Brehm

Single species of large, W Palearctic bunting, formerly placed in Emberiza; but Y 20% heavier than X; plumage nondescript, sexes alike; cutting edges of mandibles curved and of lower mandible angled (A); and has complete post-juv. moult. Palate knobbed (as in Emberiza).

598

EMBERIZIDAE

96–3500 m2 (av. 1595 m2); neighbouring territories overlap in densely populated areas. Breeding density in Europe high (140–180 per km2) in suitable marshy areas, much lower (usually less than 15 per km2) in areas of mixed habitat (Cramp and Perrins 1994). Pairing takes place in Y’s territory after break-up of winter flocks. Y is then frequently in conflict with rivals; gives threat display, with body and head held horizontal, and aggressive calls; often engages in face to face fights. Y sings from bush or reed stem, in upright relaxed posture with head and rump feathers ruffled (A). Song most intense when Y unpaired or when X incubating (Cramp and Perrins 1994). 2, perhaps 3, broods. NEST: cup of stems and blades of sedges, grasses, waterside plants, a few twigs; lined with moss, rootlets, sometimes hair or feathers; ext. diam. c. 80–90, int. diam. c. 50, cup depth c. 45; usually well hidden on ground or sedge tussocks, by water; or up to 4 m high in e.g. Salix bushes; built by X in 1–5 days, with material collected close by. EGGS: 3–7 (usually 4–5); slightly glossy, pale purplish, lilac grey or olive-brown, with scrawls, spots and blotches

of purplish or brownish black, irregularly distributed or concentrated at each end. SIZE (E. s. schoeniclus, n ¼ 859) 17–23  14–16 (195  146). LAYING DATES: no African data. NE Spain, late Apr–Aug. INCUBATION: by X only. Period: 12–15 (13) days. DEVELOPMENT AND CARE OF YOUNG: fed and cared for by both parents; brooded by X for first few days. Fledging period 10–12 days; young leave nest 3–5 days before able to fly. BREEDING SUCCESS/SURVIVAL: in European studies (E. s. schoeniclus), many nests lost to predators. In a 6 year study in England, only 20% of nests produced fledged young (16 fledged young per breeding pair); of 1846 eggs laid, 67% hatched, and about 30% produced flying young (Cramp and Perrins 1994).

Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Prs-Jones, R. (1984).

Plate 36

Emberiza melanocephala Scopoli. Black-headed Bunting. Bruant melanocephale.

(Opp. p. 539)

Emberiza melanocephala Scopoli, 1769. Annus I Historica-Nat., p. 142; Carnicola. Italy, Balkans, Turkey, Levant, S Russia and W Caspian to Iran; winters India. Vagrant to Morocco (once Tangier), Algeria (once Djanet, Aug 1960), Tunisia (once Sousse, Aug 1975) and Egypt (twice, Port Said, Mar 1946 and Suez, May 1982). A large bunting with rather long grey bill and long legs, separated from other buntings in Africa by combination of pale unstreaked underparts and no white in outer tail. Breeding Y has black head, chestnut back, and bright yellow collar and underparts (nape to mantle masked with buff when freshly moulted in winter). X rather featureless, with faintly streaked dull brown head and upperparts and yellow tinged underparts with buffy breast; lacks supercilium but may show thin malar stripe, and sometimes dark greyish crown and ear-coverts. Autumn Y like X, but with blacker lores and ear-coverts and yellower underparts. 1st-winter like X, but with finely streaked breast and flanks, whiter fringes to wing-coverts and no yellow below. Similar Cinereous Bunting E. cineracea semenowi has white outer tail, pale eye-ring and greyer underparts. Length 16–18 cm; wing, Y (n ¼ 10) 94–100 (971), X (n ¼ 10) 86–92 (893); weight, breeding area, Y (n ¼ 29) 25–33 (298), X (n ¼ 18) 23–32 (266). Calls include ‘cheuh’ (like House Sparrow Passer domesticus), and in flight a metallic ‘pchlu’ or ‘plut’ like Ortolan E. hortulana. Frequents open areas with scattered bushes and trees, orchards and olive groves. Often perches on commanding features. Forms flocks in Asia in non-breeding season. Feeds on ground on grass seeds and cereals or on berries in shrubs and low trees.

Genus Miliaria Brehm

Single species of large, W Palearctic bunting, formerly placed in Emberiza; but Y 20% heavier than X; plumage nondescript, sexes alike; cutting edges of mandibles curved and of lower mandible angled (A); and has complete post-juv. moult. Palate knobbed (as in Emberiza).

Miliaria calandra

599

Miliaria calandra (Linnaeus). Corn Bunting. Bruant proyer.

Plate 36

Emberiza calandra Linnaeus, 1758. Syst. Nat., ed. 10, p. 176; Europe [Sweden].

(Opp. p. 539)

Range and Status. Europe north to Scotland, Denmark and Oka R., Russia; Canary Is, N Africa and Mediterranean islands; east to Transcaspia, Afghanistan and W China. Resident; partial migrant in east of range; winter visitor south to Oman. Resident and winter visitor. Morocco, very common in plains and hills north of Haut Atlas; frequent in NE (Nador, Triffa, mouth of Moulaya, Beni-Snassen massif, south to Aı¨n Beni Mathar); up to 1000 m in Plateau Central, 1500– 2100 m in Moyen Atlas (Azrou-Aı¨n Leuh, Aguelmane Sidi Ali) and 2000–2500 m in Haut Atlas (Sour, Oukaı¨meden); south of Atlas Mts, breeds in Er Rachidia and Oued Draa, occurs in Souss, Agdz, Mhamid and Aoulouz-Oulad Berehil and in winter at Taliouine, Taroudant, Tiznit, Ouarzazate region (Tifoultout, Amerzgane) and Tafilalet region; vagrant south to Bouanane and Figuig. Migrants from Europe cross Strait of Gibraltar in winter, when they (or local birds) extend somewhat south of Moroccan breeding range; in E, large numbers wintering in some years are thought to include many European birds (A. Brosset in Cramp and Perrins 1994). Algeria, common and widespread from coast to northern oases (south of Biskra and Laghouat. Abundant in corn-growing districts south of Biskra: Jourdain 1915); up to 1800 m on Jebel Mahmel and 1900 m in Djurdjura; makes altitudinal and other movements in winter, when vagrant south to Beni Abbe`s and Hassi Touiel (El Gole´a) (Isenmann and Moali 2000). Tunisia, common to very common resident in N, south to line from Feriana to Gabe`s, abundant near coast, uncommon to frequent south to Sidi Mansour; in winter scarce but regular south to Douz and Libyan border (Thomsen and Jacobsen 1979). Libya, locally common in N, Sept–Apr, in flocks of up to 50 on coastal plain and occasionally south to Jebel Nafusa; singing YY in barley fields on Jefara and Azizia Plain; singing YY at Al Beyda, and c. 130 pairs and 2 nests near At Tamimi (Massa 1999); locally common in winter on plains south of Barce, Cyrenaica, in flocks of up to 300; rare at Serir – 1, 1, 2, 3, 10 and 30 birds in Sept 1969 to May 1970 (Bundy 1976). Egypt, regular in winter in Nile Delta, in Wadi el Natrun and along Nile south to Mallawi (27
440 N), also in Suez area and Faiyum; irregular on Mediterranean coast west to El Dab’a and Salum, south to Kharga (25
260 N), and on Red Sea coast to Hurghada, Gezira Umm el Showra and Ras Banas; flock of 1200 birds at Heliopolis (Cairo), Feb 1980 (Goodman and Meininger 1989). Mauritania, vagrant: Nouadhibou (Cansado), 1, 1, 3 and 1–4 in Dec 1979 to Feb 1980; Cap Timirist, 1, Oct 1987; Nouakchott, 1, Nov 1984 (Farnsworth 1994). Senegal, vagrant: 3 in harvested ricefield, Senegal R. delta, Feb 1974. Breeding density of 128 pairs (or trios) per km2 in maquis, Morocco (The´venot 1982) (and in area north of Strait of Gibraltar, 38–24 birds per 10 ha). Description. M. c. calandra (Linnaeus) (only race in Africa): N Africa and Europe east to Caucasus, Turkey and coastal Levant.

Miliaria calandra

ADULT Y: forehead to hindneck grey-brown, boldly streaked black, feathers of forehead and forecrown edged buffish; upper mantle grey-brown, narrowly and sparsely streaked black; lower mantle and scapulars grey-brown to tawny brown, boldly streaked black; back to uppertail-coverts grey-brown with fine blackish shaft streaks, rump slightly buffier. Tail dark brown; outer web of T1 fringed buff, and of T2–T5 narrowly so; tips of T4–T6 also fringed buff. Lores and indistinct superciliary stripe buff, streaked dark brown; upper cheeks and ear-coverts tawny brown, flecked buff. Broad moustachial stripe pale buff, spotted dark brown, linked to pale buff bar behind lower ear-coverts; below this a narrow broken blackish brown malar stripe. Side of neck grey-brown, streaked buff and blackish. Underparts buffish white, chin and throat spotted dark brown; sides of breast and flanks pale tawny brown, streaked blackish brown. Flight feathers, primary coverts and alula dark grey-bown; primaries edged pale buff, secondaries edged tawny buff, primary coverts and small alula feather fringed pale buff. Tertials, greater coverts and median coverts blackish, tertials and greater coverts broadly edged and tipped deep tawny-buff, median coverts tipped pale buff. Lesser coverts grey-brown, tipped buff. Underwing-coverts and axillaries buffish white. Bill pinkish, yellowish or greyish horn, grey to blackish on culmen; eyes dark brown; legs flesh brown, yellowish brown or light horn. Sexes alike. SIZE (10 YY, 10 XX, N Africa): wing, Y 98–105 (102), X 90–97 (931); tail, Y 74–80 (767), X 66–74 (685); bill, Y 16–175 (167), X 15–17 (159); tarsus, Y 25–265 (259), X 24–255 (246). WEIGHT (Morocco) 1 (447); (Malta, Mar–early June) Y (n ¼ 11) 44–56 (478), X (n ¼ 21) 35–43 (387); late June–Feb, Y (n ¼ 13) 43–52 (473), X (n ¼ 34) 34–49 (386). IMMATURE: juv. has upperparts buffier than ad., streaking browner and extending to rump and uppertail-coverts; scapulars

600

EMBERIZIDAE

and lower mantle feathers fringed pale buff; sides of head plainer, buffier, with distinct brown moustachial and malar stripes; underparts buffier (less white), with chin and throat unspotted, a few short streaks on upper breast, and narrow streaks on flanks. Tail feathers browner, more pointed; wing feathers browner, tertials, greater coverts and median coverts more sharply fringed pale buff. NESTLING: down long, yellow-buff, confined to upperparts, especially head, shoulder and back; mouth flesh-coloured; gape flanges yellow.

Field Characters. Length 18 cm. A large, plump bunting commonly seen singing its jangling song from telephone wires among grainfields. Nondescript streaked brown plumage suggests Sky Lark Alauda arvensis, but readily distinguished by short thick bill, big round head without crest, lack of white trailing edge to wing, no white in tail, very different voice. Flight heavy, legs often dangling (especially singing Y). Voice. Tape-recorded (62–73, 92, 105, 110–113, B, C). Song begins with short ‘ti’ notes which accelerate into long, buzzy, nasal, jangling trill, which may change timbre halfway through: ‘ti-ti-ti-ti-titititititi zweeeeee-zweeerrrz’; sometimes only first ‘zweeeeee’ given; stated similarity to splintering glass or bunch of keys being shaken rather farfetched. Flight call a double ‘titup’, second syllable lower; other (contact?) calls include ‘chup’, buzzy ‘dzeet’, ‘dzweeep’ or ‘bzzzz’, and hard tikking notes from birds in feeding flock. For sonagrams and further details see Cramp and Perrins (1994). General Habits. Inhabits maquis, undulating lowland, coastal plains and cultivation; less common in mountains, up to 2500 m; avoids forest, broken terrain and built-up areas, and keeps to open arable land with wheat, barley, rice and pea fields (Morocco) and salt marshes (Libya), fences, scattered shrubs, short rough grass, wasteland, roadsides, and in autumn fields where root crops have been harvested or dung spread. Lives almost solitarily in breeding season but markedly gregarious out of it. Flocks of >100 roost in marshy vegetation (Morocco), and once a flock of several thousand at roost in reeds in wadi at Titaf (Er Rachidia), Jan. Forages almost entirely on ground, in flocks of up to 300 (Morocco), but readily uses low perches: fence posts and fence wires, telegraph wires, walls, bushes and small trees when not feeding and (YY) for singing. Gait on ground a hop. Stance on perch upright, tail of Y on song post often hanging straight down. Breeding birds quite approachable; Y perches and sings openly; but foraging flocks in winter generally secretive, remaining still on arable land or in stubble; when person 40 m away approaches, whole flock flies off in rippling flight low down over ground and drops back into cover 200 m away. Complete moult in July–Oct. Migrates by day. Passage at Strait of Gibraltar Oct–Nov and late Feb–May; passage at Bahig, Egypt, mid Feb to mid Mar (K.O. Horner in Goodman and Meininger 1989). Recovery from Cap Bon (Tunisia) to SW Spain.

Food. Fallen seeds, especially of cereals. Young fed with seeds, insects and insect larvae. Breeding Habits. Few data from Africa; well known in N Europe, especially NE Scotland and Slovakia, where biology may differ from that in south of range; following summary from Cramp and Perrins (1994). Solitary nester; monogamous, successively polygynous, occasionally polyandrous; pair bond not strong, and paired Y and X are seldom together. Territorial; Y attacks rival Y entering territory; threatens in Head-forward posture (like that of Emberiza buntings); sometimes fights in air breast-to-breast with rival Y; occasionally attacks other bird species including buntings. Y sings at song post and, when approaching or leaving it, often sings in flight with Dangling-legs display, with head lowered, legs and feet dangled conspicuously and pointing somewhat outward, accompanied by shallow, rapid wingbeats. Y sings sporadically all year, regularly when breeding; X does not sing. NEST: quite large cup, loosely made of grass stems, herb stalks and roots; lined with fine grass, rootlets and some hair. SIZE: ext. diam. av. 126, int. diam. av. 76, ext. height av. 74, int. depth av. 61. Sited usually in small depression on ground, in thick, tangled grass and thistles, or (Morocco) often in low branches of shrub. Built in 1–3 days by X, accompanied on collecting trips by Y; X collects material from ground within 50 m of site, sometimes pulling up green grass. EGGS: 4–7 (NW Africa, n ¼ 131 clutches), av. 492 (Heim de Balsac and Mayaud 1962). Shape and colour variable; generally sub-elliptical and whitish, often tinged blue, buff or purple, with sparse but bold blackish blotches, scrawls and hairstreaks. SIZE: (NW Africa, n ¼ 94) 230–265  170–185, (Europe, n ¼ 844) 208–286  158– 192 (241  176). WEIGHT: av. 39. LAYING DATES: Morocco and Algeria, Apr–mid June, Libya, Apr. INCUBATION: by X only. Period: 12–14 days. DEVELOPMENT AND CARE OF YOUNG: young fed and cared for mainly by X; both parents remove faecal sacs; Y helps with feeding later on. Nestling period 9–13 days (9–10 days probably when birds disturbed). On leaving nest young can fly only weakly; they move up to 20 m on 1st day and are fed by parents for c. 2 weeks. BREEDING SUCCESS/SURVIVAL: estimated annual mortality 42% (Germany). Oldest ringed bird 105 years.

Key References Byers, C. et al. (1995), Cramp, S. and Perrins, C.M. (1994), Goodman, S.M. and Meininger, P.L. (1989), Heim de Balsac, H. and Mayaud, N. (1962), Isenmann, P. and Moali, A. (2000), The´venot, M. (1982).

Plectrophenax nivalis

601

Genus Plectrophenax Stejneger

2 species of bunting, boldly pied, sexes dissimilar, with distinctive voices and habitats: breed in Arctic, nesting amongst rocks, often snowy and icy (P. nivalis), and in hollow driftwood on shorelines (P. hyperboreus, of Bering Sea islands and coasts). P. nivalis is Holarctic, breeding between July isotherms of 2
C and 15
C and wintering in mountains and on coasts; gregarious in winter; 5 subspecies.

Plectrophenax nivalis (Linnaeus). Snow Bunting. Bruant des neiges.

Plate 36

Emberiza nivalis Linnaeus, 1758. Syst. Nat., ed. 10, p. 176; Lapland.

(Opp. p. 539)

Breeding range circumpolar, mainly north of 65
N, south to S Norway, Kamchatka, Aleutian Is, Hudson Strait and S Greenland. Migrates to winter between 40
and 60
N, south in Europe to E Croatia, but since late 1970s small numbers (up to 50) have become annual in Azores. Vagrant south to Canary Is, Madeira, Azores, Gibraltar, NW Africa, Malta and Bulgaria. Morocco, a few records Feb–Mar along Atlantic coast from Tangier to Oualidia; once Algeria, Dec 1961–Jan 1962, Tigzirt (on coast, 3
540 E). Vagrants in Africa not identified subspecifically: probably N European P. n. nivalis. Ad. Y mainly snow-white but mantle, back, scapulars and tertials black, alula and outer 3rd of wing black, rump and uppertail-coverts mottled black and white and middle of tail mainly black. Bill and eyes black, legs slaty. Ad. X, imm. and juv. variegated rufous and buff where ad. Y black; underparts mainly white; black streaks on mantle, black scallops on scapulars and long black chevrons on rufous tertials; mid-line of forehead dark brown, crown and hindneck warm brown, cheeks and breast rufous; above eyes, sides of neck, chin and throat white, flanks buffy; wings and tail boldly pied, like ad. Y; bill horn-yellow, eyes black, legs slaty. Length 16–17 cm; wing, Y 104–118 (110), X 100–107 (104); weight (n ¼ 41, Netherlands, winter) 28–44 (329), exhausted birds (n ¼ 11, Netherlands, winter) 18–23. Easily told by boldly pied plumage (Y) or sparrowy rufous upperparts and pied wings (X). Vocal, and even a solitary bird often utters distinctive short, rippling ‘tzwee-tzwee-chu-we-tu-wee’ and variants, at once melodious but buzzy and slightly twangy. Inhabits snowy tundra and mountains; in winter commonly coastal; all African vagrants have occurred on sandy coasts. Forages on saltings and along tidelines; eats seeds, some insects. X very like Snow Finch Montifringilla nivalis, a Pyrenean-Balkan bird accidental south to Malta and Canary Is but not yet seen in Africa (both species: see Cramp and Perrins 1994).

Plectrophenax nivalis

601

Genus Plectrophenax Stejneger

2 species of bunting, boldly pied, sexes dissimilar, with distinctive voices and habitats: breed in Arctic, nesting amongst rocks, often snowy and icy (P. nivalis), and in hollow driftwood on shorelines (P. hyperboreus, of Bering Sea islands and coasts). P. nivalis is Holarctic, breeding between July isotherms of 2
C and 15
C and wintering in mountains and on coasts; gregarious in winter; 5 subspecies.

Plectrophenax nivalis (Linnaeus). Snow Bunting. Bruant des neiges.

Plate 36

Emberiza nivalis Linnaeus, 1758. Syst. Nat., ed. 10, p. 176; Lapland.

(Opp. p. 539)

Breeding range circumpolar, mainly north of 65
N, south to S Norway, Kamchatka, Aleutian Is, Hudson Strait and S Greenland. Migrates to winter between 40
and 60
N, south in Europe to E Croatia, but since late 1970s small numbers (up to 50) have become annual in Azores. Vagrant south to Canary Is, Madeira, Azores, Gibraltar, NW Africa, Malta and Bulgaria. Morocco, a few records Feb–Mar along Atlantic coast from Tangier to Oualidia; once Algeria, Dec 1961–Jan 1962, Tigzirt (on coast, 3
540 E). Vagrants in Africa not identified subspecifically: probably N European P. n. nivalis. Ad. Y mainly snow-white but mantle, back, scapulars and tertials black, alula and outer 3rd of wing black, rump and uppertail-coverts mottled black and white and middle of tail mainly black. Bill and eyes black, legs slaty. Ad. X, imm. and juv. variegated rufous and buff where ad. Y black; underparts mainly white; black streaks on mantle, black scallops on scapulars and long black chevrons on rufous tertials; mid-line of forehead dark brown, crown and hindneck warm brown, cheeks and breast rufous; above eyes, sides of neck, chin and throat white, flanks buffy; wings and tail boldly pied, like ad. Y; bill horn-yellow, eyes black, legs slaty. Length 16–17 cm; wing, Y 104–118 (110), X 100–107 (104); weight (n ¼ 41, Netherlands, winter) 28–44 (329), exhausted birds (n ¼ 11, Netherlands, winter) 18–23. Easily told by boldly pied plumage (Y) or sparrowy rufous upperparts and pied wings (X). Vocal, and even a solitary bird often utters distinctive short, rippling ‘tzwee-tzwee-chu-we-tu-wee’ and variants, at once melodious but buzzy and slightly twangy. Inhabits snowy tundra and mountains; in winter commonly coastal; all African vagrants have occurred on sandy coasts. Forages on saltings and along tidelines; eats seeds, some insects. X very like Snow Finch Montifringilla nivalis, a Pyrenean-Balkan bird accidental south to Malta and Canary Is but not yet seen in Africa (both species: see Cramp and Perrins 1994).

BIBLIOGRAPHY The bibliography is in three parts: (1) general and regional references, (2) references by family or group of genera (journal titles abbreviated) and (3) acoustic references. Together, lists (1) and (2) comprise the publications cited in the text and all other significant works consulted; titles in (1) are generally not repeated in (2), except for some family monographs. If a reference cited in the text does not appear in the appropriate systematic lists in (2), it will be found in (1). A few of the most recent publications listed have been consulted for some but not all bird species.

General and Regional References Backhurst, G. C., Britton, P. L. and Mann, C. F. (1973). The less common Palaearctic migrant birds of Kenya and Tanzania. Journal of the East Africa Natural History Society and National Museum 140, 1–38. Balanc¸a, G. and Visscher, M.-N. de (1997). Composition et ´evolution saisonnie`re d’un peuplement d’oiseaux au nord du Burkina Faso (nord-Yatenga). Malimbus 19, 68–94. Balchin, C. S. (1988). Recent observations of birds from the Ivory Coast. Malimbus 10, 201–206. Balchin, C. S. (1990). Further observations of birds from the Ivory Coast. Malimbus 12, 52–53. Bannerman, D. A. (1932). Account of the birds collected (i.) by Mr. G. L. Bates on behalf of the British Museum in Sierra Leone and French Guinea; (ii.) by Lt.-Col. G. H. Houghton, R. A. M. C., in Sierra Leone, recently acquired by the British Museum. Part III. Ibis 1932, 217–261. Bannerman, D. A. (1948, 1949, 1951). ‘The Birds of Tropical West Africa’, Vols 6, 7 and 8. The Crown Agents for the Colonies, London. Barlow, C., Wacher, T. and Disley, T. (1997). ‘A Field Guide to Birds of Gambia and Senegal’. Pica Press, Robertsbridge. Barreau, D., Bergier, P. and Lesne, L. (1987). ‘L’Avifaune de l’Oukaimeden, 2200–3600 m (Haut Atlas, Maroc)’. L’Oiseau et la Revue Franc¸aise d’Ornithologie 57(4), 307–367. Basilio, A. (1963). ‘Aves de la Isla de Fernando Poo’. Editorial Coculsa, Madrid. Bates, G. L. (1909). Field-notes on the birds of southern Kamerun, West Africa. Ibis 1909, 25–74. Bates, G. L. (1911). Further notes on birds of southern Cameroon. Ibis 1911, 479–545, 581–631. Bates, G. L. (1924). On the birds collected in northwestern and northern Cameroon and parts of northern Nigeria. Ibis 1924, 1–45. Bates, G. L. (1927). Notes on some birds of Cameroon and the Lake Chad region: their status and breeding-times. Ibis 1927, 1–64. Bates, G. L. (1930). ‘Handbook of the Birds of West Africa’. John Bale, Sons and Danielsson, London. Bates, G. L. (1934). Birds of the southern Sahara and adjoining countries in French West Africa. Ibis 1934, 439–466, 685– 717. Beals, E. W. (1966). Sight additions to the avifaunal list of Ethiopia. Journal of the East Africa Natural History Society, National Museum Kenya 25, 227–228. Beals, E. W. (1970). Birds of an Euphorbia-Acacia woodland in Ethiopia: habitat and seasonal changes. Journal of Animal Ecology 39, 277–297.

van den Akker, M. (2003) Birds of Niaouli forest, southern Benin. Bulletin of the African Bird Club 10, 16–22. Ali, S. and Ripley, S. D. (1972–1974). ‘Handbook of the Birds of India and Pakistan’, Vols 5, 7, 9, and 10. Oxford University Press, Bombay. Allan, D. G. and Davies, G. B. (1999). The birds (Aves) of the middle Komati River, Swaziland. Durban Museum Novitates 24, 22–42. Allport, G., Ausden, M., Hayman, P. V., Robertson, P. and Wood, P. (1989). ‘The Conservation of the Birds of Gola Forest, Sierra Leone’. ICBP Study Report 38, 1–104. International Council for Bird Preservation, Cambridge. Amadon, D. (1953). Avian systematics and evolution in the Gulf of Guinea. Bulletin of the American Museum of Natural History 100, 393–452. Anon. (1996). ‘Important Bird Areas of Ethiopia’. Ethiopian Wildlife and Natural History Society, Addis Ababa. Archer, G. and Godman, E. M. (1961). ‘The Birds of British Somaliland and the Gulf of Aden’, Vol. 4. Oliver and Boyd, Edinburgh. Ash, J. S. (1969). Spring weights of trans-Saharan migrants in Morocco. Ibis 111, 1–10. Ash, J. S. (1983). Over fifty additions to the Somali list including two hybrids, together with notes from Ethiopia and Kenya. Scopus 7, 54–79. Ash, J. S. (1990). Additions to the avifauna of Nigeria, with notes on distributional changes and breeding. Malimbus 11, 104– 116. Ash, J. S., Dowsett, R. J. and Dowsett-Lemaire, F. (1989). New ornithological distribution records from eastern Nigeria. Tauraco Research Report 1, 13–27. Ash, J. S. and Miskell, J. E. (1993). Birds of Somalia: their habitat, status and distribution. Scopus Suppl. 1, 1–96. Ash, J. S. and Miskell, J. E. (1998). ‘Birds of Somalia’. Pica Press, Robertsbridge. Ash, J. S. and Sharland, R. E. (1986). ‘Nigeria: Assessment of Bird Conservation Priorities’. ICBP/Nigerian Conservation Foundation, Cambridge. Aspinwall, D. R. (1979). Bird notes from the Zambezi district, North-Western Province. Zambian Ornithological Society Occasional Paper 2, 1–60. Aspinwall, D. R. and Beel, C. (1998). ‘A Field Guide to Zambian Birds not found in Southern Africa’. Zambian Ornithological Society, Lusaka. Atkinson, P., Peet, N. and Alexander, J. (1991). The status and conservation of the endemic bird species of Sa˜o Tome´ and Prı´ncipe, West Africa. Bird Conservation International 1, 255–282.

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Bock, W. J. and Morony, J. J., Jr. (1978). Relationships of the Passerine Finches. Bonn. zool. Beitr. 29, 122–147. Bortoli, L. (1969). Contribution `a l’e´tude du proble`me des oiseaux granivores en Tunisie. Bull. Fac. Agron. 22–23, 32–153. Bortoli, L. (1973). Sparrows in Tunisia. In ‘Productivity, Population Dynamics and Systematics of Granivorous Birds’ (Ed. S. C. Kendeigh and J. Pinowski), 249–252. PWN-Polish Scientific Publishers, Warsaw. Bowden, C. G. R. (1987). The Arabian Golden Sparrow in North Yemen. Sandgrouse 9, 94–97. Brooker, A. (1986). Breeding the Chestnut Sparrow Sorella (Passer) eminibey. Avicult. Mag. 92, 184–185. Brosset, A. (1956). Le re´gime alimentaire de l’Effraye Tyto alba au Maroc oriental. Alauda 24, 303–305. Browne, P. W. P. (1981). Breeding of six Palaearctic birds in southwest Mauritania. Bull. Br. Orn. Club 101, 306–310. Bruggers, R. L. and Bortoli, L. (1976). Dry season nesting of the Golden Sparrow near Richard-Toll, Senegal. Terre et Vie 30, 521–527. Bundy, G. and Morgan, J. H. (1969). Notes on Tripolitanian birds. Bull. Br. Orn. Club 89, 151–159. Catry, P. and Monteiro, H. (2003). House sparrow Passer domesticus colonises Guinea-Bissau. Malimbus 25, 58–59. Chapin, J. P. (1917). The classification of the weaver-birds. Bull. Am. Mus. Nat. Hist. 37, 243–280. Clancey, P. A. (1974). Subspeciation studies in some Rhodesian birds. Arnoldia (Rhod.) 6(28), 1–43. Clouet, M., Goar, J.-L. and Barrau, C. (1998). Contribution `a l’e´tude ornithologique de l’Iˆle de Socotra. Alauda 66, 235– 246. Collias, N. E. and Collias, E. C. (1984). ‘Nest Building and Bird Behavior’. Princeton Univ. Press, Princeton. Cook, C. and Cook, J. (1998). Is this a nest building/nesting record? Bee-eater 49, 17. Craig, A., Every, B. and Summers-Smith, D. (1987). The spread of the Southern Greyheaded Sparrow in the Cape Province. Ann. Cape Prov. Mus. (Nat. Hist.) 16, 191–200.

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Deakin, R. (2001). Unusual sparrow behaviour. Afr. Birds & Birding 6(3), 11. Dean, W. R. J. (1977). The moult of the Cape Sparrow. Ostrich Suppl. 12, 108–116. Dean, W. R. J. (1978). Life expectancy of the Cape Sparrow. Ostrich 49, 16–20. De Bont, A. F. (1970). Some results of bird banding in the Congo (Kinshasa). Ostrich 41, 195–199. Demey, R., Herroelen, P. and Pedersen, T. (2000). Additions and annotations to the avifauna of Congo-Kinshasa (ex-Zaı¨re). Bull. Br. Orn. Club 120, 154–172. Densley, M. (1990). Desert Sparrow in Morocco. British Birds 83, 195–201. Doumandji, S., Metref, S. and Baziz, B. (1994). Note sur la pre´sence exceptionnelle du Moineau blanc Passer simplex sur le littoral alge´rois. Alauda 62, 246. Dupuy, A. (1969). Catalogue ornithologique du Sahara alge´rien. Oiseau R.F.O. 39, 140–160. Dymond, J. N. (1996). Biometric data of birds in southern Yemen and Socotra, spring 1993. Sandgrouse 17, 158–164. Earle´, R. A. (1988). Reproductive isolation between urban and rural populations of Cape Sparrows and House Sparrows. Acta XIX Congr. Int. Orn., II, 1778–1786. Flowers, S. S. (1925). Contribution to our knowledge of the duration of life in vertebrate animals. Proc. Zool. Soc. London, Ser. A, 95, 1365–1421. Franchimont, J., El Ghazi, A., The´venot, M. and Bergier, P. (1997). Liste GOMAC re´vise´e et statuts des espe`ces re´gulie`rement observables au Maroc. Porphyrio, 9, 28–44. Fraser, M., McMahon, L. and Underhill, G. (1992). Hybrid Cape Sparrow  House Sparrow. Promerops 206, 13. Gavrilov, E. I. (1962). A contribution to the biology of the Spanish Sparrow (Passer hispaniolensis transcaspicus). Ibis 104, 416–417. Gavrilov, E. I. (1963). The biology of the Eastern Spanish Sparrow, Passer hispaniolensis transcaspicus Tschusi, in Kazakhstan. J. Bombay Nat. Hist. Soc. 60, 301–317. Ghabbour, S. I. (1976). The ecology and pest status of sparrows (Passer) in Egypt. Int. Stud. Sparrows 9, 17–29. Guichard, K. M. (1955). The birds of Fezzan and Tibesti. Ibis 97, 393–421. Harrison, J. A. (2000). Changing suburban birds – terrestrial species. Bird Numbers 9, 46–51. Hartert, E. (1921). Capt. A. Buchanan’s Aı¨r expedition IV. The birds collected by Capt. A. Buchanan during a journey from Kano to Aı¨r or Aslen. Novit. Zool. 28, 78–141. Harwin, R. M. and Irwin, M. P. S. (1966). The spread of the House Sparrow, Passer domesticus, in south-central Africa. Arnoldia (Rhod.) 2, 1–17. Hayes, J. (1982). Notes on bird behaviour. E. Afr. Nat. Hist. Soc. Bull. 1982, 40–41. Heim de Balzac, H. (1929). Remarques sur l’ethologie de Passer simplex. Alauda 1, 68–77. Hobbs, J. (1997). More Cape Sparrow me´nages `a trois. Promerops 230, 12. Hobbs, J. (1999). Have House Sparrows left CBD? Promerops 240, 15. Holyoak, D. T. and Seddon, M. B. (1991). Notes sur la re´partition des oiseaux du Niger (2e`me partie). Alauda 59, 116–120. Hustler, K., Irwin, M. P. S. and Tree, A. J. (1991). Third report of the OAZ Rarities Committee. Honeyguide 37, 165–170. Hustler, K. and Pollard, C. (2001). The Northern Grey-headed Sparrow in Zimbabwe. Honeyguide 47(1), 64–65. Immelmann, K. (1970). Brutbiologische Beobachtungen am Kapsperling (Passer melanurus) im su ¨ dlichen Afrika. Beitr. Vogelkd. 16, 195–204.

Irwin, M. P. S. (2001). A chronology of the spread and ecological requirements of the Southern Grey-headed Sparrow in Zimbabwe. Honeyguide 47, 66–72. Irwin, M. P. S. (2002). The origins and spread of the Northern and Southern Grey-headed Sparrows in Zambia. Honeyguide 48, 41–46. Irwin, M. P. S. (2002). What determines the status of the House Sparrow in Harare? Honeyguide 48, 90–91. Jacob, J.-P. and de Schaetzen, R. (1984). De´couverte du moineau dore´ Passer luteus dans l’extreˆme sud de l’Alge´rie en relation avec le peuplement du Nord du Niger. Malimbus 6, 73–74. Johnston, R. F. (1969). Taxonomy of House Sparrows and their allies in the Mediterranean basin. Condor 71, 120–139. Jones, J. M. B. (2002). More chirpings on grey-headed sparrows. Honeyguide 48, 92. Jourdain, F. C. R. (1915). Notes on the Bird-Life of eastern Algeria. Ibis 1915, 133–139. Kirwan, G. M., Martins, R. P., Morton, K. M. and Showler, D. A. (1996). The status of birds in Socotra and ’Abd Al-Kuri and the records of the OSME survey in spring 1993. Sandgrouse 17, 83–101. Klein, R. (1989a). Erfassung und Bewertung von Einflu ¨ ssen einer Tierart auf Sahelische Kulturlandschaften am Beispiel des Goldsperlings (Passer luteus) in der Republik Niger. Geomethodica 14, 141–165. Klein, R. (1989b). Moralita¨t im Nest des Goldsperlings (Passer luteus). J. Orn. 130, 361–365. Klein, R. (1992). The pest status of the Golden Sparrow in the Republic of Niger. Proc. VII Pan-Afr. Orn. Congr., 277–283. Klein, R. and Riss, B. (1990). Die Nestlingsnahrung von Weidensperling (Passer hispaniolensis Temm.) und Gold¨ kol. Vo¨gel sperling (Passer luteus Licht.) – ein Vergleich. O 12, 53–61. Kopij, G. (2000). Bird communities of Acacia karroo dominated riparian habitats in the Free State. Mirafra 17, 33–42. Kopij, G. (2001). ‘Atlas of Birds of Bloemfontein’. National University of Lesotho, Roma, and Free State Bird Club, Bloemfontein. Kunkel, R. (1961). Allgemeines und soziales Verhalten des Braunruckengoldsperlings (Passer [Auripasser] luteus Licht.). Z. Tierpsychol. 18, 471–489. Lambert, K. (1978). Feldsperling, Passer montanus, und Bergfink, Fringilla montifringilla, in den Tropen. Beitr. Vogelkd. 24, 103. Lamm, D. W. (1959). Feeding rates for the Grey-headed Sparrow Passer g. griseus at Accra, Ghana. Ostrich 30, 161–163. Lewis, A. D. (1981). The Somali Sparrow Passer castanopterus: a breeding record for East Africa. Scopus 5, 83. Little, J. de V. (1961). Anting behaviour in the East African Grey-headed Sparrow, Passer griseus ugandae Reichw. Ostrich 32, 182. Lynes, H. (1924). On the birds of North and Central Darfur, with notes on the West-Central Kordofan and North Nuba Provinces of British Sudan. Ibis 1924, 648–719. Lynes, H. (1926). On the birds of North and Central Darfur. Taxonomic Appendix, Part I. Ibis 1926, 346–405. Maciver, M. (1998). Observing Cape Sparrows can be rewarding. Promerops 235, 18. Madsen, J. J. (1990). Desert Sparrow in Tunisia in January 1989. Dutch Birding 12, 77. McFarlane, J. (1997a). Cape Sparrow me´nage `a trois. Promerops 229, 10. McFarlane, J. (1997b). Hole-nesting Cape Sparrows. Promerops 229, 12–13. Meinertzhagen, R. (1930). ‘Nicoll’s Birds of Egypt’. Hugh Rees, London.

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Meinertzhagen, R. (1940). Autumn in Central Morocco. Ibis 1940, 14. Metzmacher, M. (1986a). La distribution des moineaux, Passer, en Alge´rie: observations comple´mentaires. Gerfaut 76, 132–139. Metzmacher, M. (1986b). Moineaux domestiques, Passer domesticus, et espagnols, Passer hispaniolensis, dans une re´gion de l’ouest alge´rien: analyse comparative de leur morphologie externe. Gerfaut 76, 317–334. Metzmacher, M. (1986c). Moineaux domestiques, Passer domesticus, et espagnols, Passer hispaniolensis, dans l’ouest alge´rien: aperc¸u des variations regionales de la morphologie externe. Gerfaut 76, 335–342. Metzmacher, M. (1986d). L’organisation spatio-temporelle de la reproduction chez le Moineau espagnol Passer hispaniolensis Temm. en zone semi-aride alge´rienne. Oiseau R.F.O. 56, 229–262. Metzmacher, M. (1990). Climate factors, activity budgets and breeding success of the Spanish Sparrow (Passer hispaniolensis (Temm.)). In ‘Granivorous Birds in Agricultural Landscape’ (Ed. J. Pinowski and J. D. Summers-Smith), 151–168. Polish Scientific Publishers, Warsaw. Milton, S. and Dean, R. (1999). Nesting thyme: the use of aromatic plants in Cape Sparrow nests. Afr. Birds and Birding 4, 37–39. Mirza, Z. B. (1974). A preliminary study of the breeding, food, sexual dimorphism and distribution of the Spanish Sparrow, Passer hispaniolensis Temm. in Libya. Int. Stud. Sparrows 7, 76–87. Mirza, Z. B., Kora, A., Sadik, L. S. and Dahnous, K. (1975). A study of breeding populations and food of Spanish Sparrow Passer hispaniolensis Temm., in Tripoli and Azizia, Libyan Arab Republic. Int. Stud. Sparrows 8, 117–123. Mlikovsky, J. and Mlikovska, R. (1985). Une contribution `a l’avifaune de Hammamet, Tunisie. Vest. cs. Spolec. Zool. 49, 208–210. Morel, G. J. and Morel, M.-Y. (1976). Nouvelles observations sur la reproduction du Moineau dore´ Passer luteus en zone semi-aride de l’ouest africain. Terre Vie 30, 493–520. Morel, G. J. and Morel, M.-Y. (1978). Ele´ments de comparaison entre Quelea qu. quelea (L.) et Passer luteus (Lichtenstein) dans les savanes tropicales de l’ouest africain. Cah. ORSTOM, se´r. biol. 13, 347–358. Morel, G. J. and Morel, M.-Y. (1980). Has the Golden Sparrow replaced the Black-faced Dioch in West Africa? Acta XVII Congr. Int. Orn., Berlin, 1150–1154. Morel, M.-Y. (1988). Successful establishment of the House Sparrow Passer domesticus in Senegambia. Proc. VI Pan-Afr. Orn. Congr. 159–162. Morel, M.-Y. and Morel, G. (1973a). Premie`res observations sur la reproduction du Moineau dore´, Passer luteus (Licht.), en zone semi-aride de l’ouest africain. Oiseau R.F.O. 43, 97–118. Morel, M.-Y. and Morel, G. (1973b). Ele´ments de comparaison du comportement reproducteur colonial de trois espe`ces de Ploceide´s: Passer luteus, Ploceus cucullatus et Quelea quelea, en zone semi-aride de l’ouest africain. Oiseau R.F.O. 43, 314–329. Moyer, D. C. and Sion, E. (1993). The spread of the House Sparrow Passer domesticus indicus in Africa with new records from Tanzania. Scopus 16, 101–105. Msimanga, A. and Slotow, R. (2000). The House Sparrow Passer domesticus (Passeriformes: Ploceidae) invasion of southern Africa: dispersal rates and the Allee effect. Durban Mus. Novit. 25, 5–11. Ndao, B. (1980). Le Moineau domestique (Passer domesticus) espe`ce nouvelle pour le Se´ne´gal. Bull. Inst. Fond. Afr. Noire 42, ser. A, 2, 422–425.

Nhlane, M. E. D. (2000). The breeding biology of the House ˆ i. Ostrich 71, Sparrow Passer domesticus at Blantyre, Malaw 80–82. Niethammer, G. (1971). Some problems connected with the House Sparrow’s colonisation of the world. Ostrich Suppl. 8, 445–448. Oatley, T. B. and Skead, D. M. (1972). Nectar feeding by South African birds. Lammergeyer 15, 65–74. Ogilvie-Grant, W. R. (1912). On the birds of Ngamiland. Ibis 1912, 355–404. Oschadleus, D. (1997). Fiscal Shrike kills Cape Sparrow. Laniarius 64, 11. Payne, R. B. (1969). Nest parasitism and display of Chestnut Sparrows in a colony of Grey-capped Social Weavers. Ibis 111, 300–307. Peacock, F. (1997). Observations on the vocalisation patterns of Cape Sparrows, Passer melanurus. Laniarius 67, 7–8. Pierce, A. J. and Rayment, M. D. (1998). Disturbance of Plain Martin Riparia paludicola nests by Grey-headed Sparrow Passer griseus swainsonii. Scopus 20, 51–52. Pollard, C. J. W. (2002). Field identification of the grey-headed sparrows. Honeyguide 48, 91–92. Rainey, H. and Lachenaud, O. (2002). Recent observations from Ivory Coast. Malimbus 24, 23–37. Riss, B. (1989). Die Verbreitung des Weidensperlings (Passer hispaniolensis) in Marokko. J. Orn. 130, 367–375. Rossouw, J. D. (2001). New records of uncommon and poorly known species for Ugandan National Parks and Forest Reserves. Scopus 21, 23–24. Rowan, M. K. (1966). Some observations of reproduction and mortality in the Cape Sparrow Passer melanurus. Ostrich Suppl. 6, 424–434. Rudnai, J. (1991). Sparrows a wrestling match. E. Afr. Nat. Hist. Soc. Bull. 21, 13–14. Ruelle, P. J. (1982). Le Moineau dore´, Passer luteus (Lichtenstein), comme de´pre´dateur des ce´re´ales en Afrique de l’Ouest. Bull. Inst. Fond. Afr. Noire 44, se´r. A, 384–405. Ruelle, P. J. and Semaille, R. (1982). Note sur l’envahissement du nord du Se´ne´gal par le Moineau dore´ Passer luteus (Lichtenstein) en pe´riode de reproduction. Malimbus 4, 27–32. Schmidt, O. (1999). Whither the House Sparrow? Promerops 238, 12–13. Siegfried, W. R. (1973). Breeding success and reproductive potential in the Cape Sparrow Passer melanurus (Mu ¨ ller). In ‘Productivity, Population Dynamics and Systematics of Granivorous Birds’ (Ed. S. C. Kendeigh and J. Pinowski), 167–179. PWN-Polish Scientific Publishers, Warsaw. Slotow, R. and Goodfriend, W. (1996). Ecogeographic variation in body size and shape of Cape sparrows (Passer melanurus) in southern Africa. J. Zool. London 238, 279–286. Slotow, R., Goodfriend, W. and Maclean, G. L. (1993). The social structure of captive Cape Sparrow flocks. Ostrich 64, 63–66. Snow, D. W. and Perrins, C. M. (1998). ‘The Birds of the Western Palearctic, Concise Edition’. Vol. 2, Passerines. Oxford Univ. Press, Oxford. Stephan, B. (1999). Zur Taxonomie mediterraner Sperlinge der Gattung Passer—Probleme weiterhin aktuell: Hybridisation, italiae, tingitanus. Mitt. Mus. Nat. Berl. Zool. Reihe 75, 3–9. Steyn, P. (2000). Grey-headed Sparrow nest site. Promerops 243, 11. Summers-Smith, J. D. (1983a). The Great Sparrow Passer motitensis motitensis (A. Smith) 1836. Babbler 6, 9–15. Summers-Smith, J. D. (1983b). The Great Sparrow. Bokmakierie 35, 51–55.

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Petronia Balchin, C. S. (1988). Recent observations of birds from Ivory Coast. Malimbus 10, 201–206. Clancey, P. A. (1995). On the generic attribution of the Yellowthroated Sparrow Petronia superciliaris of the southern Afrotropics. Honeyguide 41, 268–269. Louchart, A. (1999). Invalidite´ de la note 3321 (Alauda 67 (1): 72– 73): ‘‘Observation du Moineau bride´, Petronia superciliaris `a N’Djamena (Tchad)’’. Alauda 67, 356. Medland, R. D. (1989). Observations on breeding Red-headed Weavers and Yellow-throated Sparrows. Nyala 14, 128–130. Salewski, V. (1997). Notes on some bird species from Comoe´ National Park, Ivory Coast. Malimbus 19, 61–67. Tree, A. J. (1991). Recent reports. Honeyguide 37, 187–195. Trollope, J. (1977). The breeding and behaviour of the Bush Petronia Petronia dentata. Avicult. Mag. 83, 69–74. Trollope, J. (1979). The breeding and behaviour of the Yellowthroated Sparrow Petronia xanthocollis. Avicult. Mag. 85, 135–138.

Family PLOCEIDAE Bentz, G. D. (1979). The appendicular myology and phylogenetic relationships of the Ploceidae and Estrildidae (Aves: Passeriformes). Bull. Carnergie Mus. Nat. Hist. 15, 1–25. Boetticher, H. von (1952). ‘Die Widahvo ¨gel und Witwen’. Geest and Portig, Leipzig. Borrett, R. P. and Brooke, R. K. (1970). A field guide to the weaver finches of the genera Quelea, Euplectes and Vidua in Rhodesia. Honeyguide 63, 17–24. Chapin, J. P. (1917). The classification of the weaverbirds. Bull. Am. Mus. Nat. Hist. 37, 243–280. Collias, N. E. and Collias, E. C. (1964). Evolution of nest-building in the weaverbirds (Ploceidae). Univ. California Publ. Zool. 73, 1–162. Craig, A. (1982d). Mortality rates of some South African Ploceidae. Ostrich 53, 54–57. Craig, A. J. F. K. (1995). Adaptation and evolution in ploceid weaver nests. Ostrich 66, 100–102. Craig, A. J. F. K. (1999). Weaving a story: the relationships of the endemic Ploceidae of Madagascar. Proc. XXII Intl. Orn. Congr., Durban, 3063–3070. BirdLife South Africa, Johannesburg. Crook, J. H. (1959). Behaviour study and the classification of West African weaver birds. Proc. Linnean Soc. 170, 147–153. Crook, J. H. (1960). Nest form and construction in certain West African weaver-birds. Ibis 102, 1–25. Crook, J. H. (1962). The adaptive significance of pair formation types in weaver birds. Symp. Zool. Soc. London 8, 57–70. Crook, J. H. (1963). A comparative analysis of nest structure in the weaver birds (Ploceinae). Ibis 105, 238–262. Crook, J. H. (1964). The evolution of social organisation and visual communication in the weaverbirds (Ploceinae). Behaviour Suppl. 10, 1–178. Crook, J. H. (1969). Functional and ecological aspects of vocalization in weaver birds. In ‘Bird Vocalizations’ (Ed. R. A. Hinde). Cambridge Univ. Press, Cambridge, 265–289.

Friedmann, H. (1950). The breeding habits of the weaverbirds: a study in the biology of behavior patterns. Ann. Rep. Smithsonian Inst. 1949, 293–316. Johnson, D. F. and Horner, R. F. (1986). Identifying widows, bishops and queleas in female plumage. Bokmakierie 38, 13–17. Leisler, B. (1995). Zirkeln bei Webervo ¨geln (Ploceidae): Beziehungen zu Lebensweisen und Schnabelbau. J. Orn. 136, 267–272. ¨ komorLeisler, B., Winkler, W. and Siebenrock, K.-H. (1997). O phologische Untersuchungen am Beispiel der Webervo ¨gel (Ploceidae) und Eisvo ¨gel (Alcedinidae). Mitt. Zoo1. Mus. Berlin 73 Suppl. Ann. Orn. 21, 17–43. Mayr, E. and Greenway, J. C. (Eds) (1962). ‘Peters Check-list of Birds of the World’. Vol. XV. Museum of Comparative Zoology, Cambridge, Mass. Moreau, R. E. (1959). Notes on Ploceinae. Part one. Bull. Br. Orn. Club 79, 117–124. Moreau, R. E. (1959). Notes on the Ploceinae. Part two. Bull. Br. Orn. Club 79, 159–163. Moreau, R. E. (1959). Weaver-bird problems. Ibis 101, 334–335. Moreau, R. E. (1960). Conspectus and classification of the ploceine weaver-birds. Ibis 102, 298–321, 443–471. Morlion, M. (1964). Pterylography of the wing of the Ploceidae. Gerfaut 54, 111–124. Morlion, M. L. (1980). Pterylosis as a secondary criterion in the taxonomy of the African ploceidae and estrildidae. Proc. IV Pan-Afr. Orn. Congr., 27–41. Oschadleus, H. D. (2000). Leaf stripping in African weaverbirds. Bird Numbers 9, 28–30. Reichenow, A. (1914). ‘Die Vo ¨gel. Handbuch der Systematischen Ornithologie’. II. Ferdinand Enke, Stuttgart. Schnell, G. D. (1973). A reanalysis of nest structure in the weavers (Ploceinae) using numerical taxonomy techniques. Ibis 115, 93–106.

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Reid, J. C. (1980). The Ortolan and Cretzschmar’s Bunting: an ornithological enigma (Aves, Emberizidae). Bonn. Zool. Beitr. 30, 357–366. Roux, P., Chakir, N. and Lesne, L. (1990). Etude compare´e de la reproduction du Bruant striole´ (Emberiza striolata sahari Levaillant) dans deux types d’environnement urbain `a Marrakech (Maroc). Bie`vre 11, 13–20. Salewski, V. and Go ¨ken, F. (1999). A southern record of Cinnamon-breasted Rock Bunting Emberiza tahapisi in Lamto, Ivory Coast. Malimbus 21, 121–122. Salvan, J. (1963). Quelques observations de migrateurs du Pale´arctique dans l’est du Tchad. L’Oiseau et R.F.O. 33, 161–162. Skead, C. J. (1960). ‘The Canaries, Seedeaters and Buntings of Southern Africa’. South African Bird Book Fund, Cape Town. Skead, D. M. (1966). Some observations on the Cape Bunting Emberiza capensis (Linnaeus). Ostrich 37, 59–60. Small, B. (1992). Ageing and sexing of Ortolan and Cretzschmar’s Buntings in the field. Birding World 5, 223–228. Taylor, I. R. and Macdonald, M. A. (1978). The status of some northern guinea savanna birds in Mole National Park, Ghana. Bull. Niger. Orn. Soc. 14(45), 5–8. Tree, A. J. (1994). Recent reports. Honeyguide 40, 187–196. Tyler, S. and Tyler, L. (2001). Biometrics and moult data for nomadic Lark-like Buntings, Emberiza impetuani, in southeastern Botswana. Ostrich 72, 118–120. Vieillard, J. (1971–1972). Donne´es bioge´ographiques sur l’avifaune d’Afrique centrale. Alauda 39, 227–248, 40, 63–92.

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Acoustic References Discs and Cassettes 2. Queeny, E. M. (1951). Songs of East African Birds. American Museum of Natural History, New York. Three 12-inch, 78 r.p.m. discs; approximately 48 species. Some errors on the labels and misidentifications were noted by Boswall and North (1967). 3. Queeny, E. M. (1951). Birds of Lake Nyibor. American Museum of Natural History, New York. One 12-inch, 78 r.p.m. disc; 14 species. 4. Cowles, R. B. (1956). Sounds of a South African Homestead. Folkways Records and Service Corp., 117 W 46 St., New York 10036. One 12-inch, 3313 r.p.m. disc, No. FPX 151. 30 species. 5. North, M. E. W. (1958). Voices of African Birds. Cornell University Press, 159 Sapsucker Woods Road, Ithaca, N. Y. 14850. One 12-inch, 3313 r.p.m. disc. 42 species. The first African record concerned mainly with identification. Species are presented in systematic order, grouped on separate bands, and details given of circumstances, place and date of recording. 7. Haagner, C. H. (1961). Birds of the Kruger National Park. International Library of African Music, P.O. Box 138, Roodeport, near Johannesburg, South Africa. Two 7-inch, 45 r.p.m. discs, Nos. XTR 17044 and XTR 27045. 31 species in systematic order following Roberts (1957, ‘Birds of South Africa’, Trustees of the John Voelcker Bird Book Fund, Cape Town) and with the Roberts number; each on a separate band. 9. Haagner, C. H. (1964). Birds of Zululand. Same publisher as No. 7. Two 7-inch, 45 r.p.m. discs., Nos. XTR 4 7094 and XTR 5 7095. 27 species. 10. North, M. E. W. and McChesney, D. S. (1964). More Voices of African Birds. Houghton Mifflin Co., Boston, U.S.A. One 12-inch, 3313 r.p.m. disc. 90 species. Details of recordings are given in an accompanying booklet. Discs Nos 5 and 10 together contain the voices of 132 species, and are the first major reference work for African bird voices. 14. Stannard, J. and Niven, P. (1967). Bird Song of the Forest. Percy FitzPatrick Institute. One 12-inch 3313 r.p.m. disc. GALP 1559. 32 species. 20. Henley, A. and Pooley, A. C. (1970). Birds of the Drakensberg. Published by the authors and obtainable from Wildlife Society of South Africa. One 12-inch 3313 r.p.m. stereo disc, BD 100. 41 species. 22. Walker, A. (1970). Garden Birds of Southern Africa. Gallo (Africa) Ltd., Johannesburg; obtainable from Wildlife Society of South Africa. One 12-inch 3313 r.p.m. stereo disc, SGALP 1598. 40 species. 30. Roche´, J.-C. (1971). Birds of Kenya. Birds and Wild Beasts of Africa, No. 1. L’Oiseau Musicien, France. One 12-inch, 3313 r.p.m. stereo disc, G.07. 32 species. 32. Keith, G. S. and Gunn, W. W. H. (1971). Birds of the African Rain Forests. Sounds of Nature No. 9. Federation of Ontario Naturalists, 1262 Don Mills Road, Don Mills, Ontario M3B 2WB, Canada, and American Museum of Natural History, New York. Two 12-inch, 3313 r.p.m. discs. 95 species. The most important reference work since the records of North (Nos. 5 and 10) and the first specializing in forest birds, many of which are here published for the first time. Most species are from East Africa, some from central Africa. Species are arranged in systematic order and

33.

36.

38.

39.

46.

54.

58.

59.

62.

66.

72.

73.

75.

76.

grouped in bands; a simple announcement of the name accompanies each species, and some information is provided on the jacket. Stannard, J. (1971). Bird Sounds and Songs. Fitzpatrick Institute. Issued in conjunction with Ostrich Supplement 9. One 7-inch, 45 r.p.m. disc, NV1. 20 species. Ker, A. (1972). Safari 99. Equator Sound Studios Ltd., P.O. Box 30068, Nairobi, Kenya. One 12-inch, 3313 r.p.m. disc, ESS 1001. 63 species. Keibel, W. D. (1972). Wildlife of South West Africa. Wildlife Society of South Africa, P.O. Box 3508, Windhock, Namibia. 1 cassette, 48 species. Roche´, J.-C. (1973). Birds of South Africa. Birds and Wild Beasts of Africa, No. 2. L’Oiseau Musicien, France. One 12-inch, 3313 r.p.m. stereo disc, G. 08. About 65 species. 7 environments are presented without commentary, created by 3 or 4 birds singing simultaneously. Anon. (1966). A Night at Treetops. Sapra Studios, York Street, Nairobi, Kenya. Heartbeat of Africa, Series 1, No. 3. One 17-cm, 45 r.p.m. disc. About 10 species. Petersen, H. (1975). Belauschte Welt der Tiere. Penny, Postfach 1, 6000 Frankfurt 1, Germany. One 30-cm, 3312 r.p.m. stereo disc, Penny S 1475/10. 9 species. Natal Bird Club (c. 1978). Bird Calls, Vols 1 and 2. Natal Bird Club, P.O. Box 10909, Marine Parade, Durban 4056, South Africa. 2 cassettes. 136 species presented in random order. Lengthy and numerous cuts provided for each species. Nicolai, J. (1965). Prachtfinken. 5 Kosmos-Lehrmittel, Postfach 640, 7000 Stuttgart 1, Germany. One 7-inch, 45 r.p.m. disc, 75-0932. 4 captive species. Palmer, S. and Boswall, J. (1969–1972). A Field Guide to the Bird Songs of Britain and Europe. SR Records, Swedish Broadcasting Corp., 105 10 Stockholm, Sweden. Twelve 12-inch, 3313 r.p.m. discs, RFLP 5001–5012. 530 species, nesting or accidental in Europe, mostly wintering in Africa. Presented in systematic order, on separate bands, announced by scientific name. The most important reference work for Palearctic birds wintering in Africa. Kabaya, T. (1978). Birds of the World. I: Africa. King Records Co., Japan. One 30-cm, 3313 r.p.m. stereo disc, King Records SKS (H) 2007. 20 species. Audio Three (1981). Bird Calls. See No. 58. 3 cassettes, of which the first 2 are the same as those of No. 58; the third contains additional species. Palmer, S. and Boswall, J. (1981). A Field Guide to the Bird Songs of Britain and Europe. 16 cassettes, RFLP 5021– 5036. An updated edition of No. 62. 612 species, in boxes of 4 cassettes with commentary and list of species in each box. A first class reference collection. Audio Three, Bird Calls: Bird Families, Vol IV. 2 cassettes, 171 species. Many of these species already appear on No. 72, but here all are in systematic order. Chappuis, C. (1984). Oiseaux de France: Migrateurs et Hivernants, parts I and II. Obtainable from the author, 10 Vallon du Fer `a Cheval, 76530 La Bouille, France. Two cassettes with booklets. Present mainly flight and contact calls, not full songs, of Palearctic birds; useful because these are the vocalizations typically made in Africa by migrants. 147 species.

651

652

ACOUSTIC REFERENCES

85. Ricci, S. (1979). Bruits et ambiance d’Afrique. One 17-cm, 45 r.p.m. mono disc, No. OCR 23. Office de coope´ration radiophonique, Paris. 3 species. 86. Stjernstedt, R. (1986–93). Bird Songs of Zambia. Distributor: Wildsounds, P.O. Box 9, Holt, Norfolk NR25 7AW, UK. 3 mono cassettes. Cassette No. 3: Sylviidae (Prinia) to Emberizidae, 140 species presented in systematic order, often with several types of vocalization per species. One of the major collections of African bird voices, specializing in birds of E-central Africa. 88. Gillard, L. (1987). Southern African Bird Calls. Gillard Bird Cassettes, P.O. Box 72059, Parkview 2122, Johannesburg, South Africa. Three 90-min cassettes. 540 species presented in systematic order, often with several types of song and call per species. Cassette No. 3: Achaetops to Emberiza. A very significant and comprehensive collection, at the time with the largest number of species yet published. 89. Chappuis, C. (1990). Sounds of Migrant and Wintering Birds, Western Europe. Obtainable from the author: 10, Vallon du Fer `a Cheval, 76530 La Bouille, France. Two mono cassettes with booklet; English (and revised) version of No. 76. 91. Gibbon, G. (1991). Southern African Bird Sounds. Southern African Birding, P.O. Box 24106, Hillary 4024, South Africa. Six 90-min. mono cassettes in box with booklet. 880 species in systematic order. A huge reference collection covering almost every species in southern Africa; sound reproduction is excellent. A magnificent achievement. 99. Gibbon, G. (1995). Southern African Bird Sounds. Six CDs, 900 species. Enlarged version of No. 91 (same distributor). 101. Stro ¨mberg, M. (1994). Moroccan Bird Songs and Calls. Waxne¨sq 44, 65341 Karlstad, Sweden. One cassette with booklet; 76 species. 102. Stjernstedt, R. (1996). Rare Birds of Zambia. Distributor: see No. 86. One cassette, 94 species. 103. Stjernstedt, R. (1996). Bird Songs of Zambia, addenda. Distributor: see No. 86. One cassette, 37 species. 104. Chappuis, C. (2000). African Bird Sounds 2: Birds of North, West and Central Africa. S. E. O. F., 55, Rue Buffon, 75005 Paris, France. 11 mono CDs with book. 1045 species breeding south of the Sahara.

105. Chappuis, C. (2000). African Bird Sounds 1: Birds of Sahara, Maghreb, Madeira, Canary and Cape Verde Islands. 4 mono CDs with book. 423 species; includes almost all Palearctic birds wintering south of the Sahara. Volumes 1 and 2 of this monumental work contain c. 3500 recordings of 1468 species, about 95% of those present in the region. It is by far the largest collection of bird sounds ever published anywhere, and is the most complete source for those of African rain forests, where knowledge of vocalizations is indispensable. Some problems of speciation are also touched on. This project began 32 years ago, and was made possible by the collaboration of 141 recordists. 106. Chappuis, C. (1976). Oiseaux de Corse et de Mediterrane´e. Alauda 44, 475–503. One 30-cm, 3313 r.p.m. mono disc. Distributor, see No. 104. 61 species from the NW perimeter of the Mediterranean. 108. Smith, S. (1994). Bird recordings from The Gambia. Available from the author: 1 Serrells Barn Cottages, Langton Matravers, Swanage, Dorset BH19 3HX, England. One cassette, 52 species. 109. Smith, S. (1996). Bird Recordings from Ethiopia. Available from the author (see No. 108). One cassette, 66 species. 110. Roche´, J. C. (1993). All the Bird Songs of Britain and Europe. Available from Sittelle, Rue des Jardins, 38710 Mens, France, or Wildsounds (address under No. 86). 4 CDs, 396 species in systematic order. Lengthy cuts of superb quality. See also No. 113. 111. Thomas, R. and Thomas, S. (1996). Birds of The Gambia. Available from Wildsounds (see No. 86). One mono cassette, 50 species. 112. Roche´, J. C. and Chevereau, J. (1998). Oiseaux du Maghreb: Birds of North-West Africa. Available from Sittelle (see No. 110). 1 CD with booklet, 52 species. 113. Roche´, J. C. and Chevereau, J. (2000). Guide sonore des oiseaux d’Europe. Available from Sittelle (see No. 110). 10 CDs with 5 illustrated booklets, 414 species. Revised and enlarged version of No. 110, but organized differently. The species are grouped into 5 separate habitats, and within each habitat are listed in alphabetical order. Unfortunately the time and place of each recording are not given.

Principal disc, cassettes and CDs by region East Africa: Nos 5, 10, 32, 36. West Africa: No. 104.

Southern Africa: Nos 72, 86, 88, 91, 99, 102. North Africa: Nos 73, 89, 105, 110, 112, 113.

Institutions with Sound Libraries A. BBC B. C.

E.

Audio Three, 6 Larch Road, Durban, South Africa. British Broadcasting Corporation, Natural History Recording Library: through NSA. BLOWS: British Library of Wildlife Sounds, now NSA. Cornell University, Library of Natural Sounds, Laboratory of Ornithology, 159 Sapsucker Woods Road, Ithaca NY 14850, USA. Fonoteca Zoologica, Museo de Zoologia, Parque de le Ciutadela, 08003 Barcelona, Spain.

F.

N. NSA S.

Fitzpatrick Bird Communication Library, Bird Department, Transvaal Museum, P.O. Box 413, Pretoria 001, South Africa. Natal Bird Club, P.O. Box 10909, Marine Parade, Durban 4056, South Africa. National Sound Archive, Wildlife Section, 96 Euston Road, London NW1 2DB, UK. South African Broadcasting Corporation, Library of Wildlife Sounds, P.O. Box 4559, Johannesburg 2000, South Africa.

ERRATA

Individual Recordists ALEX ASP BAR BRU BUT CART CHA CHR CRO DAV DEV ERA FISP GARD GBG GIB GIL GREG GRI GUT

Alexander-Marrack, P. D., NSA Aspinwall, D. R., FITZ Barlow, C. R. Brunel, J. Butynski, T. M. Carter, C., FITZ, NSA Chappuis, C., NSA Christy, P. Crook, J. Davidson, P. Devez, A. Erard, C. Fishpool, L. D. C., NSA Gardner, N. Greenberg, D. A. Gibbon, G., FITZ Gillard, L., FITZ Gregory, A. R. Grimes, L., NSA Guttinger, H. R.

HAM HAN HAZ HOR HRS HUG JOHN JO PJ KAE P KEI LEM LER LUT MAC McVIC MEES MIL MOR MOY NIC

Hammick, J. Hansen, P. S. Hazevoet, C. J., NSA Horne, J., Cornell Harris, T. Huguet, P. Johnson, E. D. H. Jones, P. J. Kaestner, P., Cornell Keith, S., Cornell Dowsett-Lemaire, F., FITZ, NSA Lernould, J. M. Lutgens, H., NSA Macaulay, L. R., Cornell McVicker, R., FITZ, NSA Mees, V., NSA Mild, K. Morel, G. Moyer, D., Cornell Nicolai, J.

NOR PAR PARK PAY PEA ROC RODE SINC SM STJ SUN TBS TUCK TYE VIEL WALK WAT WHI ZIM

North, M. E. W., Cornell, NSA Parker, V., FITZ Parker, T. A., Cornell Payne, R. B., NSA Pearson, D. J. Roche´, J. C., NSA Rodewald, P. Sinclair, J. C. Smith, H., NSA Stjernstedt, R., NSA Sundler, J. Smith, T. B. Tucker, N. Tye, A., NSA Vielliard, J., NSA Walker, A., FITZ, NSA Watts, D. E., NSA White, T. C., NSA Zimmerman, D. and M.

ERRATA Volume VI p. vii, Authorship. We apologize for the omission of Deleornis from the list of genera written by M. P. S. Irwin and of Lamprotornis pulcher from the list of species accounts prepared by R. Wilkinson. p. 176. Cyanomitra obscura obscura is indistinguishable from C. o. cephaelis (see Tye, A. and Macaulay, L. R., 1991, Malimbus 14, 65–66). The taxon cephaelis Bates 1930 must therefore be subsumed in obscura Jardine 1843. pp. 387, 393. The remark under Malaconotus monteiri, p. 387, ‘‘1–2 seen at 1250 m at L. Edib, Bakossi Mts, 1997 (Williams 1997)’’ refers to M. gladiator and should have appeared on p. 393. Plate 26, opp. p. 399. Eyes of Laniarius mufumbiri and L. erythrogaster are nearly white, rather than the straw colour shown.

p. 522. Dicrurus ludwigii is mapped slightly too far north in Ghana. p. 604. Weights given under Lamprotornis chalybaeus nordmanni refer instead to L. c. sycobius. pp. 646–648 and 706. Three papers were overlooked in the account of Zavattariornis stresemanni: Dellelegn, Y. (1993). Observations on the Ethiopian Bush Crow Zavattariornis stresemanni in Yabello, southern Ethiopia. Proc. VIII Pan-Afr. Orn. Congr., 469–474. Londei, T. (1996). Sur corvide di Zavattari, Zavattariornis stresemanni Moltoni, 1938, in memoria del suo descrittore. Riv. Ital. Orn., Milano 65(2) 93–100. Tesfaye Hundessa. (1991). Survival status review of the Ethiopian Bushcrow (Zavattariornis stresemanni 1938) in the Boreno area, Ethiopia. Walia 13, 9–13.

653

ERRATA

Individual Recordists ALEX ASP BAR BRU BUT CART CHA CHR CRO DAV DEV ERA FISP GARD GBG GIB GIL GREG GRI GUT

Alexander-Marrack, P. D., NSA Aspinwall, D. R., FITZ Barlow, C. R. Brunel, J. Butynski, T. M. Carter, C., FITZ, NSA Chappuis, C., NSA Christy, P. Crook, J. Davidson, P. Devez, A. Erard, C. Fishpool, L. D. C., NSA Gardner, N. Greenberg, D. A. Gibbon, G., FITZ Gillard, L., FITZ Gregory, A. R. Grimes, L., NSA Guttinger, H. R.

HAM HAN HAZ HOR HRS HUG JOHN JO PJ KAE P KEI LEM LER LUT MAC McVIC MEES MIL MOR MOY NIC

Hammick, J. Hansen, P. S. Hazevoet, C. J., NSA Horne, J., Cornell Harris, T. Huguet, P. Johnson, E. D. H. Jones, P. J. Kaestner, P., Cornell Keith, S., Cornell Dowsett-Lemaire, F., FITZ, NSA Lernould, J. M. Lutgens, H., NSA Macaulay, L. R., Cornell McVicker, R., FITZ, NSA Mees, V., NSA Mild, K. Morel, G. Moyer, D., Cornell Nicolai, J.

NOR PAR PARK PAY PEA ROC RODE SINC SM STJ SUN TBS TUCK TYE VIEL WALK WAT WHI ZIM

North, M. E. W., Cornell, NSA Parker, V., FITZ Parker, T. A., Cornell Payne, R. B., NSA Pearson, D. J. Roche´, J. C., NSA Rodewald, P. Sinclair, J. C. Smith, H., NSA Stjernstedt, R., NSA Sundler, J. Smith, T. B. Tucker, N. Tye, A., NSA Vielliard, J., NSA Walker, A., FITZ, NSA Watts, D. E., NSA White, T. C., NSA Zimmerman, D. and M.

ERRATA Volume VI p. vii, Authorship. We apologize for the omission of Deleornis from the list of genera written by M. P. S. Irwin and of Lamprotornis pulcher from the list of species accounts prepared by R. Wilkinson. p. 176. Cyanomitra obscura obscura is indistinguishable from C. o. cephaelis (see Tye, A. and Macaulay, L. R., 1991, Malimbus 14, 65–66). The taxon cephaelis Bates 1930 must therefore be subsumed in obscura Jardine 1843. pp. 387, 393. The remark under Malaconotus monteiri, p. 387, ‘‘1–2 seen at 1250 m at L. Edib, Bakossi Mts, 1997 (Williams 1997)’’ refers to M. gladiator and should have appeared on p. 393. Plate 26, opp. p. 399. Eyes of Laniarius mufumbiri and L. erythrogaster are nearly white, rather than the straw colour shown.

p. 522. Dicrurus ludwigii is mapped slightly too far north in Ghana. p. 604. Weights given under Lamprotornis chalybaeus nordmanni refer instead to L. c. sycobius. pp. 646–648 and 706. Three papers were overlooked in the account of Zavattariornis stresemanni: Dellelegn, Y. (1993). Observations on the Ethiopian Bush Crow Zavattariornis stresemanni in Yabello, southern Ethiopia. Proc. VIII Pan-Afr. Orn. Congr., 469–474. Londei, T. (1996). Sur corvide di Zavattari, Zavattariornis stresemanni Moltoni, 1938, in memoria del suo descrittore. Riv. Ital. Orn., Milano 65(2) 93–100. Tesfaye Hundessa. (1991). Survival status review of the Ethiopian Bushcrow (Zavattariornis stresemanni 1938) in the Boreno area, Ethiopia. Walia 13, 9–13.

653

INDEXES Plate key-page numbers in italics.

Scientific Names abyssinicus, Ploceus cucullatus 165 abyssinicus, Sporopipes frontalis 64 aenea, Vidua chalybeata 436 aethiopica, Quelea quelea 208 afer, Euplectes 228, 155 affinis, Emberiza 572, 523 afra, Pytilia 358, 266 africana, Fringilla coelebs 452 Alario 526 Alario alario 527, 474 Alario leucolaemus 529, 474 alario, Alario 527, 474 albifrons, Amblyospiza 198, 219 albifrons, Serinus burtoni 499 albinucha, Ploceus 182, 139 albirostris, Bubalornis 50, 26 albogularis, Serinus 494, 458 albonotatus, Euplectes 235, 203 alexanderi, Amadina fasciata 380 alexanderi, Nigrita luteifrons 254 alienus, Ploceus 121, 75 alienus, Rhodopechys sanguineus 553 Amadina 379 Amadina erythrocephala 381, 347 Amadina fasciata 379, 347 Amandava 398 Amandava amandava 398, 282 amandava, Amandava 398, 282 amaurocephalus, Ploceus bicolor 184 amauropteryx, Vidua chalybeata 437 Amblyospiza 198 Amblyospiza albifrons 198, 219 Anaplectes 97 Anaplectes rubriceps 97, 155 angolensis, Estrilda astrild 303 angolensis, Euplectes capensis 233 angolensis, Nigrita canicapillus 256 angolensis, Ploceus 192, 154 angolensis, Uraeginthus 325, 330 ankoberensis, Carduelis 549, 475 anomala, Brachycope 213, 155 Anomalospiza 417 Anomalospiza imberbis 417, 394 ansorgei, Euplectes gierowii 227 ansorgei, Lagonosticta rhodopareia 374 ansorgei, Nesocharis 268, 266

ansorgei, Ortygospiza atricollis 385 ansorgei, Parmoptila woodhousei 262 ansorgei, Plocepasser mahali 56 approximans, Euplectes capensis 233 ardens, Euplectes 239, 203 arnaudi, Pseudonigrita 67, 74 astrild, Estrilda 301, 282 asymmetrurus, Euplectes albonotatus 236 atricapilla, Estrilda 307, 282 atricollis, Ortygospiza 384, 330 atrogularis, Serinus 471, 411 aucupum, Vidua orientalis 423 aurantiiventris, Carduelis chloris 540 aurantius, Ploceus 130, 90 aureoflavus, Ploceus subaureus 127 aureonucha, Ploceus 180, 139 aureus, Euplectes 232, 155 australis, Cryptospiza reichenovii 284 avakubi, Estrilda atricapilla 307 axillaris, Euplectes 242, 203 axillaris, Ploceus badius 180

baddeleyi, Hypargos niveoguttatus 343 badius, Ploceus 179, 154 baglafecht, Ploceus 106, 74 balearica, Fringilla coelebs 453 ballmanni, Malimbus 78, 27 bannermani, Ploceus 109, 74 bansoensis, Nesocharis shelleyi 265 barbara, Petronia petronia 47 barbatus, Serinus mozambicus 480 bartletti, Malimbus rubricollis 96 basutoensis, Emberiza capensis 582 batesi, Ploceus 110, 75 beattyi, Ploceus intermedius 143 bella, Carduelis cannabina 545 belli, Pytilia melba 356 bengalus, Uraeginthus 326, 330 benguellensis, Estrilda paludicola 293 benguellensis, Passer motitensis 12 benguellensis, Serinus gularis 506 bertrandi, Ploceus 104, 75 bicolor, Nigrita 257, 219

655

bicolor, Ploceus 183, 139 bicolor, Spermestes 405, 347 bocagei, Coccopygia melanotis 272 bocagei, Euplectes axillaris 243 boehmi, Dinemellia dinemelli 54 bohndorffi, Ploceus cucullatus 165 bojeri, Ploceus 132, 90 bororensis, Petronia superciliaris 46 Brachycope 213 Brachycope anomala 213, 155 brachydactyla, Carpospiza 551, 522 brachypterus, Ploceus nigricollis 116 bradfieldi, Emberiza capensis 582 bradfieldi, Ortygospiza fuscocrissa 388 brevior, Ploceus ocularis 118 britannica, Carduelis carduelis 542 brittoni, Serinus hypostictus 465 brunneiceps, Lagonosticta senegala 362 brunneigularis, Uraeginthus bengalus 327 brunnescens, Nigrita bicolor 258 Bubalornis 49 Bubalornis albirostris 50, 26 Bubalornis niger 52, 26 Bucanetes 556 Bucanetes githagineus 556, 522 buchanani, Petronia dentata 42 buchanani, Serinus 488, 458 budongoensis, Ploceus baglafecht 106 burnieri, Ploceus 140, 91 burtoni, Serinus 498, 459 butleri, Anomalospiza imberbis 418 buvryi, Coccothraustes coccothraustes 560

cabanisi, Emberiza 565, 523 cabanisi, Pseudonigrita 70, 74 cabanisii, Ploceus intermedius 143 cabaret, Carduelis 545, 475 caerulescens, Estrilda 287, 267 caesia, Emberiza 594, 539 calandra, Miliaria 599, 539 camburni, Ploceus xanthops 129 camerunensis, Vidua 445, 410

656

INDEXES: SCIENTIFIC NAMES

cana, Spermophaga haematina 315 candidaus, Nigrita canicapillus 256 canicapilla, Serinus gularis 506 canicapillus, Nigrita 255, 219 caniceps, Serinus mozambicus 481 canicollis, Serinus 513, 459 cannabina, Carduelis 545, 475 cantans, Euodice 412, 347 capensis, Emberiza 582, 538 capensis, Euplectes 233, 155 capensis, Ploceus 122, 90 capistrata, Nesocharis 270, 266 capistratus, Serinus 465, 411 capitalba, Amblyospiza albifrons 199 capitalis, Ploceus melanocephalus 174 cardinalis, Quelea 204, 155 CARDUELINAE 455 Carduelis 535 Carduelis ankoberensis 549, 475 Carduelis cabaret 545, 475 Carduelis cannabina 545, 475 Carduelis carduelis 541, 475 Carduelis chloris 536, 475 Carduelis flammea 545, 475 Carduelis johannis 547, 475 Carduelis spinus 543, 475 carduelis, Carduelis 541, 475 Carpodacus 550 Carpodacus erythrinus 550, 522 Carpospiza 551 Carpospiza brachydactyla 551, 522 carychroa, Emberiza flaviventris 568 cassini, Malimbus 82, 27 castaneiceps, Ploceus 133, 90 castaneigula, Ploceus xanthopterus 136 castaneofuscus, Ploceus nigerrimus 161 castanops, Ploceus 137, 91 castanopterus, Passer 31, 11 caurinus, Ploceus velatus 146 cavandishi, Estrilda astrild 301 centralis, Estrilda rhodopyga 297 centralis, Hypargos niveoguttatus 343 centralis, Malimbus rubricollis 96 centralis, Vidua chalybeata 437 chalybeata, Vidua 436, 410 charmosyna, Estrilda 311, 282 chloris, Carduelis 536, 475 chlorotica, Carduelis chloris 540 chubbi, Mandingoa nitidula 276 chyuluensis, Cryptospiza salvadorii 280 cia, Emberiza 587, 538 cineracea, Emberiza 591, 539 cinerascens, Sporopipes frontalis 64 cinereovinacea, Euschistospiza 340, 267 cinnamomea, Emberiza capensis 582 cirlus, Emberiza 589, 538

citerior, Pytilia melba 355 citrinella, Emberiza 589, 538 citrinella, Serinus 518, 474 citrinelloides, Serinus 460, 411 citrinipectus, Serinus 477, 458 clanceyi, Pytilia melba 355 clarkei, Sporaeginthus subflavus 393 Clytospiza 348 Clytospiza monteiri 348, 267 Coccopygia 271 Coccopygia melanotis 272, 267 Coccopygia quartinia 273, 267 Coccothraustes 559 Coccothraustes coccothraustes 559, 522 coccothraustes, Coccothraustes 559, 522 codringtoni, Vidua 450, 410 coelebs, Fringilla 452, 523 cognominata, Emberiza cabanisi 566 Coliuspasser 214 collaris, Ploceus cucullatus 165 collaris, Serinus xantholaemus 486 concolor, Euplectes ardens 240 concolor, Neospiza 530, 475 confidens, Lagonosticta senegala 362 congica, Lagonosticta rubricata 371 conradsi, Euplectes macroura 238 contigua, Amadina fasciata 380 cordofanicus, Passer 15, 11 coronatus, Malimbus 87, 27 corsicana, Serinus 518 craspedopterus, Euplectes hordeaceus 225 crassirostris, Bucanetes githagineus 557 crassirostris, Euplectes capensis 233 crassirostris, Malimbus malimbicus 92 Crithagra 486 crocatus, Ploceus ocularis 118 crocopygia, Serinus albogularis 494 Cryptospiza 278 Cryptospiza jacksoni 279, 266 Cryptospiza reichenovii 281, 266 Cryptospiza salvadorii 280, 266 Cryptospiza shelleyi 278, 266 cucullatus, Ploceus 164, 138 cucullatus, Spermestes 401, 347 vcurvirostra, Loxia 562, 522 cyanocephalus, Uraeginthus 332, 330 cyanopleurus, Uraeginthus angolensis 325

damarensis, Estrilda astrild 301 damarensis, Passer melanurus 17 damarensis, Pytilia melba 356 damarensis, Serinus flaviventris 492 definita, Euplectes progne 247 delacouri, Euplectes progne 247

delamerei, Estrilda erythronotos 310 delamerei, Euplectes progne 247 Dendrospiza 460 dentata, Petronia 42, 10 deserti, Serinus atrogularis 472 diabolicus, Nigrita canicapillus 256 diadematus, Euplectes 230, 202 dicrocephalus, Ploceus 173, 138 diffusus, Passer 7, 10 digressa, Ortygospiza fuscocrissa 389 dilutescens, Ploceus cucullatus 165 dimidiata, Serinus xantholaemus 486 dimidiatus, Ploceus melanocephalus 174 dinemelli, Dinemellia 54, 26 Dinemellia 54 Dinemellia dinemelli 54, 26 dissita, Amadina erythrocephala 381 domesticus, Passer 20, 11 donaldsoni, Plocepasser 61, 26 donaldsoni, Serinus 487, 458 dorsalis, Pseudonigrita arnaudi 67 dorsomaculatus, Ploceus 187, 154 dorsostriata, Ortygospiza gabonensis 386 dorsostriatus, Serinus 483, 458 duboisi, Ploceus melanocephalus 175 dybowskii, Euschistospiza 338, 267 eisentrauti, Estrilda nonnula 306 elgonensis, Linurgus olivaceus 532 elgonensis, Serinus gularis 506 elizae, Estrilda nonnula 306

Emberiza 564 Emberiza cineracea 591, 539 Emberiza citrinella 589, 538 Emberiza affinis 572, 523 Emberiza cabanisi 565, 523 Emberiza caesia 594, 539 Emberiza capensis 582, 538 Emberiza cia 587, 538 Emberiza cirlus 589, 538 Emberiza flaviventris 567, 523 Emberiza forbesi 573, 523 Emberiza hortulana 592, 539 Emberiza impetuani 585, 538 Emberiza leucocephalos 588, 538 Emberiza melanocephala 598, 539 Emberiza poliopleura 571, 523 Emberiza pusilla 596, 539 Emberiza rustica 595, 539 Emberiza schoeniclus 596, 539 Emberiza socotrana 580, 538 Emberiza striolata 577, 538 Emberiza tahapisi 574, 523 Emberiza vincenti 583, 538 EMBERIZIDAE 564

INDEXES: SCIENTIFIC NAMES

emiliae, Nigrita canicapillus 256 emini, Ploceus baglafecht 106 emini, Pytilia phoenicoptera 352 emini, Sporopipes frontalis 64 eminibey, Passer 34, 10 endemion, Serinus gularis 507 eques, Euplectes albonotatus 236 eremica, Emberiza impetuani 585 eremobius, Ploceus baglafecht 107 erythrinus, Carpodacus 550, 522 erythrocephala, Amadina 381, 347 erythrogaster, Malimbus 86, 27 erythronotos, Estrilda 309, 282 erythrops, Quelea 205, 155 Estrilda 285 Estrilda atricapilla 307, 282 Estrilda astrild 301, 282 Estrilda caerulescens 287, 267 Estrilda charmosyna 311, 282 Estrilda erythronotos 309, 282 Estrilda kandti 308, 282 Estrilda melpoda 295, 267 Estrilda nigriloris 304, 282 Estrilda nonnula 305, 282 Estrilda paludicola 292, 267 Estrilda perreini 289, 267 Estrilda poliopareia 293, 267 Estrilda rhodopyga 297, 282 Estrilda thomensis 290, 267 Estrilda troglodytes 298, 282 ESTRILDIDAE 251 euchlorus, Passer 40, 10 Euodice 412 Euodice cantans 412, 347 Euplectes 214 Euplectes afer 228, 155 Euplectes albonotatus 235, 203 Euplectes ardens 239, 203 Euplectes aureus 232, 155 Euplectes axillaris 242, 203 Euplectes capensis 233, 155 Euplectes franciscanus 215, 202 Euplectes gierowii 227, 202 Euplectes hartlaubi 244, 218 Euplectes hordeaceus 224, 202 Euplectes jacksoni 249, 218 Euplectes macroura 237, 203 Euplectes nigroventris 223, 202 Euplectes orix 217, 202 Euplectes progne 246, 218 Euplectes psammocromius 246, 218 Euplects diadematus 230, 202 Euschistospiza 338 Euschistospiza cinereovinacea 340, 267 Euschistospiza dybowskii 338, 267

fagani, Malimbus erythrogaster 86

fasciata, Amadina 379, 347 finschi, Ploceus velatus 146 fischeri, Ploceus melanocephalus 174 fischeri, Vidua 433, 394 flammea, Carduelis 545, 475 flavicaudata, Pytilia melba 356 flaviceps, Ploceus manyar 195 flavigaster, Emberiza flaviventris 569 flavigula, Petronia superciliaris 46 flavigula, Serinus 484, 458 flavipes, Ploceus 104, 139 flaviventris, Emberiza 567, 523 flaviventris, Serinus 491, 458 flavivertex, Serinus 511, 459 flavodorsalis, Lagonosticta senegala 361 forbesi, Emberiza 573, 523 forbesi, Lagonosticta rara 369 fradei, Serinus rufobrunneus 501 franciscanus, Euplectes 215, 202 fricki, Ploceus baglafecht 106 friederichseni, Euplectes gierowii 227 Fringilla 451 Fringilla coelebs 452, 523 Fringilla montifringilla 454, 523 FRINGILLIDAE 451 fringilloides, Spermestes 407, 347 frobenii, Ploceus cucullatus 165 frommi, Estrilda rhodopyga 297 frommi, Pyrenestes minor 323 frommi, Serinus sulphuratus 489 frontalis, Serinus 463, 411 frontalis, Sporopipes 64, 26 fucata, Estrilda melpoda 295 fulgens, Passer castanopterus 32 funerea, Vidua 447, 410 furensis, Ploceus taeniopterus 142 fuscata, Ortygospiza gabonensis 387 fuscocrissa, Ortygospiza 388, 330 fusconotus, Nigrita 259, 219

gabonensis, Ortygospiza 386, 330 galbula, Ploceus 157, 91 gengleri, Fringilla coelebs 453 georgicus, Passer diffusus 8 gierowii, Euplectes 227, 202 githagineus, Bucanetes 556, 522 golandi, Ploceus 172, 154 gommaensis, Serinus mozambicus 481 gongonensis, Passer 6, 10 goslingi, Emberiza tahapisi 575 Granatina 333 Granatina granatina 335, 330 Granatina ianthinogaster 334, 330 granatina, Granatina 335, 330 grandis, Ploceus 170, 138 granti, Malimbus malimbicus 92

granti, Serinus mozambicus 481 graueri, Euschistospiza cinereovinacea 340 graueri, Ploceus cucullatus 165 graueri, Serinus striolatus 496 griseicapilla, Odontospiza 400, 347 griseitergum, Serinus canicollis 514 griseus, Passer 2, 10 grotei, Pytilia melba 356 grotei, Serinus mozambicus 481 guillarmodi, Serinus flaviventris 492 guineensis, Lagonosticta senegala 361 gularis, Serinus 506, 459 gurneti, Serinus burtoni 499

haematina, Spermophaga 314, 283 haematocephala, Lagonosticta rubricata 372 hartlaubi, Euplectes 244, 218 hawkeri, Granatina ianthinogaster 334 hemileucus, Passer insularis 17 hesperus, Serinus flaviventris 492 heuglini, Ploceus 153, 91 hewitti, Serinus albogularis 494 hildebrandti, Lagonosticta rubricata 372 hildegardae, Serinus capistratus 466 hispaniolensis, Passer 24, 11 Histurgops 62 Histurgops ruficaudus 62, 26 holomelas, Ploceus albinucha 183 homogenes, Cryptospiza reichenovii 284 hordeaceus, Euplectes 224, 202 hortulana, Emberiza 592, 539 huillensis, Serinus flavivertex 512 humeralis, Euplectes hartlaubi 245 humilis, Serinus gularis 507 hygrophila, Pytilia melba 356 Hypargos 341 Hypargos margaritatus 344, 283 Hypargos niveoguttatus 342, 283 Hypochera 420 Hypochera lorenzi 439 Hypocherina incognita 437 hypocherina, Vidua 431, 394 hypogrammica, Pytilia 349, 266 hypostictus, Serinus 464, 411

ianthinogaster, Granatina 334, 330 ibadanensis, Malimbus 84, 27 idius, Hypargos niveoguttatus 343 imberbis, Anomalospiza 417, 394 impetuani, Emberiza 585, 538 impiger, Serinus atrogularis 472

657

658

INDEXES: SCIENTIFIC NAMES

incana, Estrilda perreini 289 incognita, Hypocherina 437 indicus, Passer domesticus 22 inornata, Euodice cantans 413 insignis, Ploceus 186, 154 insularis, Emberiza tahapisi 575 insularis, Passer 16, 11 interior, Hypargos niveoguttatus 343 interjecta, Vidua 424, 395 intermedia, Emberiza schoeniclus 597 intermedia, Euplectes macroura 237 intermedia, Quelea quelea 208 intermedius, Bubalornis niger 52 intermedius, Ploceus 142, 91 interscapularis, Ploceus tricolor 181 italiae, Passer hispaniolensis 25

jacksoni, Cryptospiza 279, 266 jacksoni, Euplectes 249, 218 jacksoni, Ploceus 175, 138 jagoensis, Estrilda astrild 301 jamesoni, Lagonosticta rhodopareia 374 jamesoni, Parmoptila 264, 219 jamesoni, Ploceus xanthops 129 jebelmarrae, Emberiza striolata 579 jessei, Pytilia melba 356 johannis, Carduelis 547, 475 jubaensis, Anaplectes rubriceps 98 jubaensis, Pytilia melba 356

kalaharica, Emberiza flaviventris 568 kandti, Estrilda 308, 282 kasaica, Amblyospiza albifrons 199 kasamaensis, Serinus atrogularis 472 katangae, Ploceus 151, 91 katangae, Uraeginthus bengalus 327 kavirondensis, Ploceus luteolus 111 kempi, Estrilda astrild 303 kersteni, Ploceus bicolor 184 kigomaensis, Ploceus bicolor 184 kikuyuensis, Serinus citrinelloides 461 kilimensis, Coccopygia quartinia 274 kilimensis, Cryptospiza salvadorii 280 kilimensis, Euplectes capensis 233 kilimensis, Linurgus olivaceus 532 kilimensis, Serinus burtoni 499 kiwanukae, Estrilda charmosyna 312 knysnae, Euplectes capensis 233 koliensis, Serinus 467, 411 kubaensis, Carpodacus erythrinus 551

ladoensis, Euplectes afer 228 laeneni, Passer griseus 3 Lagonosticta 359

Lagonosticta rubricata 371, 346 Lagonosticta larvata 367, 331 Lagonosticta nitidula 366, 331 Lagonosticta rara 369, 331 Lagonosticta rhodopareia 373, 346 Lagonosticta rufopicta 364, 331 Lagonosticta sanguinodorsalis 376, 346 Lagonosticta senegala 360, 331 Lagonosticta umbrinodorsalis 378, 346 Lagonosticta virata 375, 346 landanae, Lagonosticta rubricata 372 languens, Serinus sulphuratus 490 larvata, Lagonosticta 367, 331 larvaticola, Vidua 444, 410 laterita, Lagonosticta rufopicta 365 lathami, Quelea quelea 208 laticauda, Euplectes ardens 240 lebomboensis, Ploceus bicolor 184 leucocephalos, Emberiza 588, 538 leucolaemus, Alario 529, 474 leuconotus, Anaplectes rubriceps 98 leucopterus, Serinus 502, 459 leucopygius, Serinus 470, 411 limpopoensis, Emberiza capensis 582 lineata, Pytilia 354, 266 Linurgus 531 Linurgus olivaceus 531, 474 litoris, Euplectes capensis 233 littoralis, Uraeginthus bengalus 327 locustella, Paludipasser 390, 330 loitanus, Sporopipes frontalis 64 Lonchura malabarica 412, 413 lopezi, Pytilia 351 lorenzi, Hypochera 439 louisae, Rhynchostruthus 535, 475 loveridgei, Serinus sulphuratus 489 Loxia 562 Loxia curvirostra 562, 522 luangwae, Passer diffusus 8 luteifrons, Nigrita 253, 219 luteolus, Ploceus 111, 75 luteus, Passer 36, 10 lwenarum, Serinus atrogularis 472

macmillani, Estrilda astrild 303 macrocerca, Euplectes macroura 237 macrorhynchus, Euplectes capensis 233 macrospilotus, Hypargos niveoguttatus 343 macroura, Euplectes 237, 203 macroura, Vidua 420, 394 maculicollis, Serinus dorsostriatus 483 mahali, Plocepasser 56, 26 malabarica, Lonchura 412, 413 malensis, Ploceus nigricollis 116 malimbicus, Malimbus 89, 27

Malimbus 77 Malimbus ballmanni 78, 27 Malimbus cassini 82, 27 Malimbus coronatus 87, 27 Malimbus erythrogaster 86, 27 Malimbus ibadanensis 84, 27 Malimbus malimbicus 89, 27 Malimbus nitens 93, 27 Malimbus racheliae 80, 27 Malimbus rubricollis 95, 27 Malimbus scutatus 81, 27 Mandingoa 275 Mandingoa nitidula 275, 283 manuensis, Ploceus xanthops 129 manyar, Ploceus 195, 154 margaritatus, Hypargos 344, 283 marleyi, Ploceus xanthopterus 136 marshalli, Serinus flaviventris 492 marunguensis, Estrilda atricapilla 307 marwitzi, Estrilda paludicola 293 maryae, Vidua 442, 410 massaica, Estrilda astrild 303 maxima, Amblyospiza albifrons 199 maxwelli, Ploceus albinucha 183 mechowi, Euplectes axillaris 243 mediterranea, Carduelis cannabina 545 melanocephala, Emberiza 598, 539 melanocephalus, Ploceus 174, 138 melanochrous, Serinus 499, 459 melanogaster, Ploceus 120, 75 melanorhynchus, Plocepasser mahali 56 melanota, Amblyospiza albifrons 199 melanotis, Coccopygia 272, 267 melanoxanthus, Ploceus nigricollis 116 melanurus, Passer 17, 11 melba, Pytilia 354, 266 melpoda, Estrilda 295, 267 mendosus, Serinus gularis 507 mennelli, Serinus 504, 459 mentalis, Ploceus bicolor 184 meridionalis, Amadina fasciata 380 microrhynchus, Malimbus nitens 93 Miliaria 598 Miliaria calandra 599, 539 militaris, Bubalornis niger 52 miniscula, Ortygospiza fuscocrissa 388 minor, Estrilda astrild 303 minor, Pyrenestes 323, 283 minor, Spermestes bicolor 406 monacha, Ploceus pelzelni 113 montana, Amblyospiza albifrons 199 montanorum, Serinus gularis 506 montanus, Passer 32, 11 monteiri, Clytospiza 348, 267 montifringilla, Fringilla 454, 523 moreaui, Malimbus nitens 93 mosambicus, Passer diffusus 8

INDEXES: SCIENTIFIC NAMES

motitensis, Passer 9, 11 mozambicus, Serinus 479, 458 muelleri, Ortygospiza fuscocrissa 388

natalensis, Uraeginthus angolensis 325 nebularum, Emberiza capensis 582 neglecta, Lagonosticta rubricata 371 neidiecki, Estrilda astrild 301 Neospiza 530 Neospiza concolor 530, 475 Nesocharis 265 Nesocharis capistrata 270, 266 Nesocharis ansorgei 268, 266 Nesocharis shelleyi 265, 266 neumanni, Ploceus baglafecht 106 neumanni, Vidua chalybeata 437 niassensis, Euplectes afer 228 niassensis, Uraeginthus angolensis 325 nicolli, Ploceus 191, 154 niedecki, Carduelis carduelis 542 niethammeri, Sporaeginthus subflavus 393 niger, Bubalornis 52, 26 nigeriae, Emberiza affinis 573 nigeriae, Malimbus rubricollis 96 nigeriae, Vidua 441, 410 nigerrima, Vidua funerea 447 nigerrimus, Ploceus 161, 139 nigriceps, Ploceus cucullatus 165 nigriceps, Serinus 519, 474 nigriceps, Spermestes bicolor 406 nigricollis, Lagonosticta larvata 368 nigricollis, Ploceus 115, 75 nigrifrons, Euplectes orix 220 nigrifrons, Malimbus malimbicus 92 nigrifrons, Ploceus velatus 146 nigriloris, Estrilda 304, 282 nigrimentus, Ploceus 108, 74 Nigrita 253 Nigrita bicolor 257, 219 Nigrita canicapillus 255, 219 Nigrita fusconotus 259, 219 Nigrita luteifrons 253, 219 nigrotemporalis, Ploceus baglafecht 106 nigroventris, Euplectes 223, 202 niloticus, Passer domesticus 21 nitens, Malimbus 93, 27 nitidula, Lagonosticta 366, 331 nitidula, Mandingoa 275, 283 nivalis, Plectrophenax 601, 539 nivenorum, Emberiza tahapisi 575 niveoguttatus, Hypargos 342, 283 nonnula, Estrilda 305, 282 Notiospiza 192 nyansae, Estrilda astrild 303 nyanzae, Coccopygia quartinia 274 nyikae, Ploceus baglafecht 107

obsoleta, Rhodospiza 555, 522 obtusa, Vidua 427, 395 occidentalis, Estrilda astrild 303 ochrogaster, Estrilda paludicola 293 Ochrospiza 469 ocularis, Cryptospiza reichenovii 284 ocularis, Ploceus 117, 75 Odontospiza 400 Odontospiza griseicapilla 400, 347 okavangoensis, Vidua chalybeata 437 olivaceiceps, Ploceus 189, 154 olivaceus, Linurgus 531, 474 olivaceus, Ploceus capensis 123 omoensis, Emberiza affinis 573 orangensis, Serinus albogularis 494 orientalis, Emberiza cabanisi 566 orientalis, Euodice cantans 413 orientalis, Vidua 423, 395 orix, Euplectes 217, 202 Ortygospiza 383 Ortygospiza atricollis 384, 330 Ortygospiza fuscocrissa 388, 330 Ortygospiza gabonensis 386, 330 oryzivora, Padda 415, 347 ostrinus, Pyrenestes 318, 283 ovambensis, Serinus atrogularis 472

Pachyphantes 196 Pachyphantes superciliosus 196, 154 Padda 415 Padda oryzivora 415, 347 pallens, Serinus leucopygius 470 pallida, Ortygospiza fuscocrissa 389 pallida, Petronia pyrgita 44 pallida, Quelea cardinalis 204 pallidicrissa, Lagonosticta senegala 362 pallidior, Estrilda charmosyna 311 pallidior, Serinus tristriatus 510 pallidior, Sporopipes frontalis 64 paludicola, Estrilda 292, 267 Paludipasser 390 Paludipasser locustella 390, 330 paradisaea, Vidua 428, 395 Parmoptila 260 Parmoptila jamesoni 264, 219 Parmoptila rubrifrons 263, 219 Parmoptila woodhousei 261, 219 paroptus, Ploceus cucullatus 165 parva, Carduelis carduelis 542 Passer 1 Passer castanopterus 31, 11 Passer cordofanicus 15, 11 Passer diffusus 7, 10 Passer domesticus 20, 11 Passer eminibey 34, 10 Passer euchlorus 40, 10 Passer gongonensis 6, 10

Passer griseus 2, 10 Passer hispaniolensis 24, 11 Passer insularis 16, 11 Passer luteus 36, 10 Passer melanurus 17, 11 Passer montanus 32, 11 Passer motitensis 9, 11 Passer rufocinctus 13, 11 Passer shelleyi 14, 11 Passer simplex 29, 11 Passer suahelicus 7, 10 Passer swainsonii 4, 10 PASSERIDAE 1 peasei, Estrilda astrild 303 pectoralis, Plocepasser mahali 56 peguensis, Ploceus manyar 195 pelzelni, Ploceus 112, 75 percivali, Pytilia melba 356 perreini, Estrilda 289, 267 Petronia 41 Petronia dentata 42, 10 Petronia petronia 47, 10 Petronia pyrgita 44, 10 Petronia superciliaris 45, 10 petronia, Petronia 47, 10 Philetairus 72 Philetairus socius 72, 74 phoeniceus, Euplectes axillaris 243 phoenicomerus, Euplectes capensis 234 phoenicoptera, Pytilia 352, 266 Phormoplectes 192 pica, Spermestes fringilloides 408 Plectrophenax 601 Plectrophenax nivalis 601, 539 PLOCEIDAE 48 Plocepasser 55 Plocepasser donaldsoni 61, 26 Plocepasser mahali 56, 26 Plocepasser rufoscapulatus 60, 26 Plocepasser superciliosus 58, 26 Ploceus 101 Ploceus albinucha 182, 139 Ploceus alienus 121, 75 Ploceus angolensis 192, 154 Ploceus aurantius 130, 90 Ploceus aureonucha 180, 139 Ploceus badius 179, 154 Ploceus baglafecht 106, 74 Ploceus bannermani 109, 74 Ploceus batesi 110, 75 Ploceus bertrandi 104, 75 Ploceus bicolor 183, 139 Ploceus bojeri 132, 90 Ploceus burnieri 140, 91 Ploceus capensis 122, 90 Ploceus castaneiceps 133, 90 Ploceus castanops 137, 91 Ploceus cucullatus 164, 138

659

660

INDEXES: SCIENTIFIC NAMES

Ploceus dicrocephalus 173, 138 Ploceus dorsomaculatus 187, 154 Ploceus flavipes 104, 139 Ploceus galbula 157, 91 Ploceus golandi 172, 154 Ploceus grandis 170, 138 Ploceus heuglini 153, 91 Ploceus insignis 186, 154 Ploceus intermedius 142, 91 Ploceus jacksoni 175, 138 Ploceus katangae 151, 91 Ploceus luteolus 111, 75 Ploceus manyar 195, 154 Ploceus melanocephalus 174, 138 Ploceus melanogaster 120, 75 Ploceus nicolli 191, 154 Ploceus nigerrimus 161, 139 Ploceus nigricollis 115, 75 Ploceus nigrimentus 108, 74 Ploceus ocularis 117, 75 Ploceus olivaceiceps 189, 154 Ploceus pelzelni 112, 75 Ploceus preussi 188, 154 Ploceus princeps 134, 90 Ploceus reichardi 150, 91 Ploceus rubiginosus 177, 154 Ploceus ruweti 152, 91 Ploceus sanctithomae 193, 139 Ploceus spekei 158, 138 Ploceus spekeoides 160, 138 Ploceus subaureus 126, 90 Ploceus subpersonatus 114, 75 Ploceus taeniopterus 141, 91 Ploceus temporalis 125, 90 Ploceus tricolor 181, 139 Ploceus velatus 145, 91 Ploceus vitellinus 148, 91 Ploceus weynsi 171, 139 Ploceus xanthops 128, 90 Ploceus xanthopterus 135, 90 plowesi, Emberiza capensis 582 po, Ploceus nigricollis 116 poensis, Spermestes bicolor 406 poliogastra, Estrilda perreini 289 poliogenys, Spermophaga 313, 283 poliogyna, Loxia curvirostra 562 polionota, Lagonosticta rubricata 371 poliopareia, Estrilda 293, 267 poliopleura, Emberiza 571, 523 Poliospiza 503 praedi, Malimbus rubricollis 96 preussi, Ploceus 188, 154 prigoginei, Linurgus olivaceus 532 princeps, Emberiza flaviventris 569 princeps, Ploceus 134, 90 progne, Euplectes 246, 218 psammocromius, Euplectes 246, 218

pseudobarbatus, Serinus mozambicus 480 Pseudochloroptila 520 Pseudochloroptila symonsi 525, 474 Pseudochloroptila totta 521, 474 Pseudonigrita 67 Pseudonigrita arnaudi 67, 74 Pseudonigrita cabanisi 70, 74 punctigula, Serinus mozambicus 481 purpurascens, Vidua 448, 410 pusilla, Emberiza 596, 539 pusillus, Euplectes franciscanus 216 pusillus, Serinus 520, 474 pustulata, Spermophaga haematina 314 Pyrenestes 316 Pyrenestes minor 323, 283 Pyrenestes ostrinus 318, 283 Pyrenestes sanguineus 322, 283 pyrgita, Petronia 44, 10 Pyrrhula 559 Pyrrhula pyrrhula 559, 522 pyrrhula, Pyrrhula 559, 522 Pytilia 349 Pytilia afra 358, 266 Pytilia lineata 354, 266 Pytilia hypogrammica 349, 266 Pytilia melba 354, 266 Pytilia phoenicoptera 352, 266

quanzae, Euplectes axillaris 243 quartinia, Coccopygia 273, 267 Quelea 204 Quelea cardinalis 204, 155 Quelea quelea 207, 155 Quelea erythrops 205, 155 quelea, Quelea 207, 155 quintoni, Serinus flaviventris 492

racheliae, Malimbus 80, 27 rara, Lagonosticta 369, 331 raricola, Vidua 443, 410 regia, Vidua 434, 394 reichardi, Ploceus 150, 91 reichardi, Serinus 509, 459 reichenovii, Cryptospiza 281, 266 reichenowi, Ploceus baglafecht 106 reichenowi, Serinus 473, 411 reidi, Emberiza capensis 582 rendalli, Lagonosticta senegala 362 retusus, Granatina granatina 336 rex, Ploceus aurantius 131 Rhinoploceus 104 rhodesiae, Quelea cardinalis 204 rhodopareia, Lagonosticta 373, 346 Rhodopechys 553

Rhodopechys sanguineus 553, 522 rhodopsis, Lagonosticta senegala 361 rhodopyga, Estrilda 297, 282 Rhodospiza 555 Rhodospiza obsoleta 555, 522 Rhynchostruthus 533 Rhynchostruthus louisae 535, 475 Rhynchostruthus socotranus 534, 475 riggenbachi, Serinus leucopygius 470 roosevelti, Granatina ianthinogaster 334 roseicrissa, Estrilda paludicola 293 ruberrima, Lagonosticta senegala 362 rubiginosus, Ploceus 177, 154 rubricata, Lagonosticta 371, 346 rubriceps, Anaplectes 97, 155 rubriceps, Malimbus malimbicus 89 rubricollis, Malimbus 95, 27 rubricromus, Ploceus capensis 123 rubrifrons, Parmoptila 263, 219 rubriventris, Estrilda astrild 303 ruficapilla, Spermophaga 315, 283 ruficaudus, Histurgops 62, 26 rufidorsalis, Passer domesticus 21 rufitergum, Petronia superciliaris 45 rufobrunneus, Serinus 500, 459 rufocinctus, Passer 13, 11 rufopicta, Lagonosticta 364, 331 rufoscapulatus, Plocepasser 60, 26 rufovelatus, Malimbus rubricollis 96 rustica, Emberiza 595, 539 ruthae, Estrilda paludicola 293 ruwenzori, Cryptospiza salvadorii 280 ruweti, Ploceus 152, 91

sabinjo, Euplectes capensis 233 saharae, Passer simplex 30 sahari, Emberiza striolata 578 salvadorii, Cryptospiza 280, 266 samaliyae, Serinus mozambicus 481 sanctithomae, Ploceus 193, 139 sanghae, Emberiza striolata 579 sanguineus, Pyrenestes 322, 283 sanguineus, Rhodopechys 553, 522 sanguinodorsalis, Lagonosticta 376, 346 santhome, Serinus mozambicus 481 sassi, Serinus flavivertex 512 saturata, Amblyospiza albifrons 199 saturatior, Emberiza striolata 579 saturiator, Nigrita bicolor 258 schistaceus, Nigrita canicapillus 256 schlegeli, Mandingoa nitidula 276 schoeniclus, Emberiza 596, 539 sclateri, Ploceus bicolor 184 scotops, Serinus 468, 411 scutatus, Malimbus 81, 27

INDEXES: SCIENTIFIC NAMES

scutatus, Spermestes cucullatus 402 scutopartitus, Malimbus scutatus 81 semenovi, Emberiza cineracea 591 semideserti, Serinus atrogularis 472 senegala, Lagonosticta 360, 331 septemstriata, Emberiza tahapisi 575 Serinus 455, 511 Serinus donaldsoni 487, 458 Serinus albogularis 494, 458 Serinus atrogularis 471, 411 Serinus buchanani 488, 458 Serinus burtoni 498, 459 Serinus canicollis 513, 459 Serinus capistratus 465, 411 Serinus citrinella 518, 474 Serinus citrinelloides 460, 411 Serinus citrinipectus 477, 458 Serinus dorsostriatus 483, 458 Serinus flavigula 484, 458 Serinus flaviventris 491, 458 Serinus flavivertex 511, 459 Serinus frontalis 463, 411 Serinus gularis 506, 459 Serinus hypostictus 464, 411 Serinus koliensis 467, 411 Serinus leucopterus 502, 459 Serinus leucopygius 470, 411 Serinus melanochrous 499, 459 Serinus mennelli 504, 459 Serinus mozambicus 479, 458 Serinus nigriceps 519, 474 Serinus pusillus 520, 474 Serinus reichardi 509, 459 Serinus reichenowi 473, 411 Serinus rufobrunneus 500, 459 Serinus scotops 468, 411 Serinus serinus 516, 474 Serinus striolatus 495, 459 Serinus sulphuratus 489, 458 Serinus syriacus 518, 474 Serinus tristriatus 510, 459 Serinus whytii 497, 459 Serinus xantholaemus 486, 458 Serinus xanthopygius 476, 411 serinus, Serinus 516, 474 seshekeensis, Serinus atrogularis 472 sharpei, Serinus sulphuratus 489 Sharpia 192 sharpii, Ploceus baglafecht 107 shelleyi, Cryptospiza 278, 266 shelleyi, Nesocharis 265, 266 shelleyi, Passer 14, 11 shelleyi, Ploceus velatus 146 shelleyi, Serinus sulphuratus 489 siccata, Granatina granatina 336 simplex, Passer 29, 11 sloggetti, Emberiza impetuani 585 smithersi, Ortygospiza fuscocrissa 389

smithersii, Emberiza capensis 582 socius, Philetairus 72, 74 socotrana, Emberiza 580, 538 socotranus, Rhynchostruthus 534, 475 soligena, Estrilda erythronotos 309 somaliensis, Lagonosticta senegala 362 somereni, Serinus atrogularis 472 sordahlae, Serinus albogularis 494 soror, Euplectes macroura 237 soudanensis, Pytilia melba 356 spekei, Ploceus 158, 138 spekeoides, Ploceus 160, 138 Spermestes 401 Spermestes bicolor 405, 347 Spermestes cucullatus 401, 347 Spermestes fringilloides 407, 347 Spermophaga 312 Spermophaga haematina 314, 283 Spermophaga poliogenys 313, 283 Spermophaga ruficapilla 315, 283 spilootus, Ploceus cucullatus 165 spinus, Carduelis 543, 475 spodiogenys, Fringilla coelebs 453 spoliator, Quelea quelea 208 Sporaeginthus 392 Sporaeginthus subflavus 392, 282 Sporopipes 63 Sporopipes frontalis 64, 26 Sporopipes squamifrons 65, 26 squamifrons, Sporopipes 65, 26 stentor, Plocepasser mahali 56 stephanophorus, Ploceus melanogaster 121 stictifrons, Ploceus bicolor 184 stigmatophorus, Spermestes bicolor 406 striatipectus, Serinus reichardi 509 strictus, Euplectes afer 229 striolata, Emberiza 577, 538 striolatus, Serinus 495, 459 stuartirwini, Coccopygia quartinia 274 stuhlmanni, Ploceus baglafecht 107 stygiceps, Passer diffusus 8 suahelica, Euplectes ardens 240 suahelicus, Passer 7, 10 suahelicus, Ploceus ocularis 118 subaureus, Ploceus 126, 90 subflavus, Sporaeginthus 392, 282 subpersonatus, Ploceus 114, 75 subsolanus, Passer motitensis 12 sulphuratus, Serinus 489, 458 sundevalli, Euplectes orix 220 superciliaris, Petronia 45, 10 superciliosus, Pachyphantes 196, 154 superciliosus, Plocepasser 58, 26 swainsonii, Passer 4, 10 sylvanus, Ploceus bicolor 184 symonsi, Pseudochloroptila 525, 474 syriacus, Serinus 518, 474

taeniopterus, Ploceus 141, 91 taha, Euplectes afer 229 tahapisi, Emberiza 574, 523 tahatali, Ploceus velatus 146 tandae, Amblyospiza albifrons 199 tando, Serinus mozambicus 481 tanganjicae, Serinus burtoni 499 taruensis, Serinus dorsostriatus 483 temporalis, Ploceus 125, 90 tenebridorsa, Estrilda astrild 301 tenuirostris, Ploceus ocularis 118 tephronotus, Ploceus bicolor 184 terricolor, Plocepasser mahali 56 tessellatus, Spermestes cucullatus 402 thamnophila, Pytilia melba 356 theresae, Emberiza striolata 578 Thomasophantes 193 thomensis, Estrilda 290, 267 thomensis, Serinus rufobrunneus 501 thompsonae, Serinus canicollis 513 tingitanus, Passer domesticus 21 togoensis, Spermophaga haematina 314 togoensis, Vidua 426, 395 tongensis, Ploceus subaureus 126 torrida, Estrilda perreini 289 totta, Pseudochloroptila 521, 474 transcaspicus, Passer hispaniolensis 25 transvaalensis, Euplectes capensis 233 transvaalensis, Serinus scotops 468 traversii, Euplectes axillaris 243 tricolor, Ploceus 181, 139 tristriatus, Serinus 510, 459 troglodytes, Estrilda 298, 282 trothae, Ploceus rubiginosus 177 tschadensis, Estrilda melpoda 295 tschusii, Emberiza schoeniclus 597 turgidus, Euplectes orix 220

uelensis, Paludipasser locustella 391 ugandae, Lagonosticta rubricata 371 ugandae, Ortygospiza atricollis 385 ugandae, Passer griseus 3 ugogoensis, Uraeginthus bengalus 327 ultramarina, Vidua chalybeata 437 uluensis, Ploceus vitellinus 149 umbrinodorsalis, Lagonosticta 378, 346 umbrosus, Serinus scotops 468 unicolor, Amblyospiza albifrons 199 upembae, Ploceus katangae 152 Uraeginthus 324 Uraeginthus angolensis 325, 330 Uraeginthus bengalus 326, 330 Uraeginthus cyanocephalus 332, 330 uropygialis, Nigrita fusconotus 260

vanmarli, Carduelis chloris 540

661

662

INDEXES: ENGLISH NAMES

vansoni, Serinus mozambicus 481 velatus, Ploceus 145, 91 vicarius, Ploceus olivaceiceps 189 vicinus, Passer melanurus 17 victoriae, Ploceus (castanops) 137 Vidua 419 Vidua paradisaea 428, 395 Vidua camerunensis 445, 410 Vidua chalybeata 436, 410 Vidua codringtoni 450, 410 Vidua fischeri 433, 394 Vidua funerea 447, 410 Vidua hypocherina 431, 394 Vidua interjecta 424, 395 Vidua larvaticola 444, 410 Vidua macroura 420, 394 Vidua maryae 442, 410 Vidua nigeriae 441, 410 Vidua obtusa 427, 395 Vidua orientalis 423, 395

Vidua purpurascens 448, 410 Vidua raricola 443, 410 Vidua regia 434, 394 Vidua togoensis 426, 395 Vidua wilsoni 439, 410 VIDUIDAE 416 vinacea, Emberiza capensis 582 vinacea, Lagonosticta larvata 367 vincenti, Emberiza 583, 538 viniceps, Quelea erythrops 206 virata, Lagonosticta 375, 346 virginiae, Mandingoa nitidula 276 vitellinus, Ploceus 148, 91 voousi, Carduelis chloris 536 vulpecula, Emberiza affinis 573

wilsoni, Vidua 439, 410 witherbyi, Emberiza schoeniclus 597 woltersi, Amblyospiza albifrons 200 woltersi, Spermestes bicolor 406 woodhousei, Parmoptila 261, 219

xantholaemus, Serinus 486, 458 xanthomelas, Euplectes capensis 233 xanthops, Ploceus 128, 90 xanthopterus, Ploceus 135, 90 xanthopygius, Serinus 476, 411

zambesiensis, Euplectes capensis 233 zanzibaricus, Euplectes axillaris 243 zedlitzi, Bucanetes githagineus 557

weynsi, Ploceus 171, 139 whytii, Serinus 497, 459 wilsoni, Serinus sulphuratus 490

English Names Antpecker, Jameson’s 264, 219 Antpecker, Red-fronted 263, 219 Antpecker, Woodhouse’s 261, 219 Avadavat, Red 398, 282 Bishop, Black 227, 202 Bishop, Black-winged 224, 202 Bishop, Fire-fronted 230, 202 Bishop, Golden-backed 232, 155 Bishop, Northern Red 215, 202 Bishop, Southern Red 217, 202 Bishop, Yellow 233, 155 Bishop, Yellow-crowned 228, 155 Bishop, Zanzibar 223, 202 Bluebill, Grant’s 313, 283 Bluebill, Red-headed 315, 283 Bluebill, Western 314, 283 Brambling 454, 523 Buffalo-Weaver, Red-billed 52, 26 Buffalo-Weaver, White-billed 50, 26 Buffalo-Weaver, White-headed 54, 26 Bullfinch, Eurasian 559, 522 Bunting, African Rock 573 Bunting, Black-headed 598, 539 Bunting, Brown-rumped 572, 523 Bunting, Cabanis’s 565, 523 Bunting, Cape 582, 538 Bunting, Cinereous 591, 539 Bunting, Cinnamon-breasted 573, 523 Bunting, Cirl 589, 538 Bunting, Common Reed 596, 539 Bunting, Corn 599, 539

Bunting, Cretzschmar’s 594, 539 Bunting, Golden-breasted 567, 523 Bunting, House 577, 538 Bunting, Lark-like 585, 538 Bunting, Little 596, 539 Bunting, Ortolan 592, 539 Bunting, Pine 588, 538 Bunting, Rock 587, 538 Bunting, Rustic 595, 539 Bunting, Snow 601, 539 Bunting, Socotra 580, 538 Bunting, Somali 571, 523 Bunting, Somali Goldenbreasted 571 Bunting, Vincent’s 583, 538

Canary, Black-faced 465, 411 Canary, Black-headed 527, 474 Canary, Black-throated 471, 411 Canary, Brimstone 489, 458 Canary, Bully 489 Canary, Cape 513, 459 Canary, Damara 529, 474 Canary, Forest 468, 411 Canary, Grey 470 Canary, Lemon-breasted 477, 458 Canary, Northern Grosbeak- 487, 458 Canary, Papyrus 467, 411 Canary, Protea 502, 459 Canary, Southern Grosbeak- 488, 458 Canary, White-bellied 483, 458

Canary, White-throated 494, 458 Canary, Yellow 491, 458 Canary, Yellow-crowned 511, 459 Canary, Yellow-eyed 479 Canary, Yellow-fronted 479, 458 Chaffinch 452, 523 Citril, African 460, 411 Citril, Southern 464, 411 Citril, Western 463, 411 Cordon-bleu, Blue-capped 334, 330 Cordon-bleu, Red-cheeked 332, 330 Crimsonwing, Abyssinian 280, 266 Crimsonwing, Dusky 279, 266 Crimsonwing, Red-faced 281, 266 Crimsonwing, Shelley’s 278, 266 Crossbill, Common 562, 522 Cuckoo-Finch 417, 394

Finch, Citril 518, 474 Finch, Crimson-winged 553, 522 Finch, Cut-throat 379, 347 Finch, Desert 555, 522 Finch, Melba 354 Finch, Red-headed 381, 347 Finch, Trumpeter 556, 522 Firefinch, African 371, 346 Firefinch, Bar-breasted 364, 331 Firefinch, Black-bellied 369, 331 Firefinch, Black-faced 367, 331 Firefinch, Blue-billed 371 Firefinch, Brown 366, 331 Firefinch, Chad 378, 346 Firefinch, Jameson’s 373, 346

662

INDEXES: ENGLISH NAMES

vansoni, Serinus mozambicus 481 velatus, Ploceus 145, 91 vicarius, Ploceus olivaceiceps 189 vicinus, Passer melanurus 17 victoriae, Ploceus (castanops) 137 Vidua 419 Vidua paradisaea 428, 395 Vidua camerunensis 445, 410 Vidua chalybeata 436, 410 Vidua codringtoni 450, 410 Vidua fischeri 433, 394 Vidua funerea 447, 410 Vidua hypocherina 431, 394 Vidua interjecta 424, 395 Vidua larvaticola 444, 410 Vidua macroura 420, 394 Vidua maryae 442, 410 Vidua nigeriae 441, 410 Vidua obtusa 427, 395 Vidua orientalis 423, 395

Vidua purpurascens 448, 410 Vidua raricola 443, 410 Vidua regia 434, 394 Vidua togoensis 426, 395 Vidua wilsoni 439, 410 VIDUIDAE 416 vinacea, Emberiza capensis 582 vinacea, Lagonosticta larvata 367 vincenti, Emberiza 583, 538 viniceps, Quelea erythrops 206 virata, Lagonosticta 375, 346 virginiae, Mandingoa nitidula 276 vitellinus, Ploceus 148, 91 voousi, Carduelis chloris 536 vulpecula, Emberiza affinis 573

wilsoni, Vidua 439, 410 witherbyi, Emberiza schoeniclus 597 woltersi, Amblyospiza albifrons 200 woltersi, Spermestes bicolor 406 woodhousei, Parmoptila 261, 219

xantholaemus, Serinus 486, 458 xanthomelas, Euplectes capensis 233 xanthops, Ploceus 128, 90 xanthopterus, Ploceus 135, 90 xanthopygius, Serinus 476, 411

zambesiensis, Euplectes capensis 233 zanzibaricus, Euplectes axillaris 243 zedlitzi, Bucanetes githagineus 557

weynsi, Ploceus 171, 139 whytii, Serinus 497, 459 wilsoni, Serinus sulphuratus 490

English Names Antpecker, Jameson’s 264, 219 Antpecker, Red-fronted 263, 219 Antpecker, Woodhouse’s 261, 219 Avadavat, Red 398, 282 Bishop, Black 227, 202 Bishop, Black-winged 224, 202 Bishop, Fire-fronted 230, 202 Bishop, Golden-backed 232, 155 Bishop, Northern Red 215, 202 Bishop, Southern Red 217, 202 Bishop, Yellow 233, 155 Bishop, Yellow-crowned 228, 155 Bishop, Zanzibar 223, 202 Bluebill, Grant’s 313, 283 Bluebill, Red-headed 315, 283 Bluebill, Western 314, 283 Brambling 454, 523 Buffalo-Weaver, Red-billed 52, 26 Buffalo-Weaver, White-billed 50, 26 Buffalo-Weaver, White-headed 54, 26 Bullfinch, Eurasian 559, 522 Bunting, African Rock 573 Bunting, Black-headed 598, 539 Bunting, Brown-rumped 572, 523 Bunting, Cabanis’s 565, 523 Bunting, Cape 582, 538 Bunting, Cinereous 591, 539 Bunting, Cinnamon-breasted 573, 523 Bunting, Cirl 589, 538 Bunting, Common Reed 596, 539 Bunting, Corn 599, 539

Bunting, Cretzschmar’s 594, 539 Bunting, Golden-breasted 567, 523 Bunting, House 577, 538 Bunting, Lark-like 585, 538 Bunting, Little 596, 539 Bunting, Ortolan 592, 539 Bunting, Pine 588, 538 Bunting, Rock 587, 538 Bunting, Rustic 595, 539 Bunting, Snow 601, 539 Bunting, Socotra 580, 538 Bunting, Somali 571, 523 Bunting, Somali Goldenbreasted 571 Bunting, Vincent’s 583, 538

Canary, Black-faced 465, 411 Canary, Black-headed 527, 474 Canary, Black-throated 471, 411 Canary, Brimstone 489, 458 Canary, Bully 489 Canary, Cape 513, 459 Canary, Damara 529, 474 Canary, Forest 468, 411 Canary, Grey 470 Canary, Lemon-breasted 477, 458 Canary, Northern Grosbeak- 487, 458 Canary, Papyrus 467, 411 Canary, Protea 502, 459 Canary, Southern Grosbeak- 488, 458 Canary, White-bellied 483, 458

Canary, White-throated 494, 458 Canary, Yellow 491, 458 Canary, Yellow-crowned 511, 459 Canary, Yellow-eyed 479 Canary, Yellow-fronted 479, 458 Chaffinch 452, 523 Citril, African 460, 411 Citril, Southern 464, 411 Citril, Western 463, 411 Cordon-bleu, Blue-capped 334, 330 Cordon-bleu, Red-cheeked 332, 330 Crimsonwing, Abyssinian 280, 266 Crimsonwing, Dusky 279, 266 Crimsonwing, Red-faced 281, 266 Crimsonwing, Shelley’s 278, 266 Crossbill, Common 562, 522 Cuckoo-Finch 417, 394

Finch, Citril 518, 474 Finch, Crimson-winged 553, 522 Finch, Cut-throat 379, 347 Finch, Desert 555, 522 Finch, Melba 354 Finch, Red-headed 381, 347 Finch, Trumpeter 556, 522 Firefinch, African 371, 346 Firefinch, Bar-breasted 364, 331 Firefinch, Black-bellied 369, 331 Firefinch, Black-faced 367, 331 Firefinch, Blue-billed 371 Firefinch, Brown 366, 331 Firefinch, Chad 378, 346 Firefinch, Jameson’s 373, 346

INDEXES: ENGLISH NAMES

Firefinch, Firefinch, Firefinch, Firefinch, Firefinch, Firefinch, Firefinch,

Kulikoro 375 Mali 375, 346 Pink-backed 373 Red-billed 360, 331 Rock 376, 346 Senegal 360 Vinaceous 367

Goldbreast 392 Goldfinch, European 541, 475 Greenfinch, European 536, 475 Grenadier, Purple 335, 230 Grosbeak, Arabian Goldenwinged 534, 475 Grosbeak, Sa˜o Tome´ 530, 475 Grosbeak, Somali Goldenwinged 535, 475

Hawfinch 559, 522

Indigobird, Bar-breasted Firefinch 439, 410 Indigobird, Barka 444, 410 Indigobird, Black-faced Firefinch 444 Indigobird, Cameroon 445 Indigobird, Dusky 447, 410 Indigobird, Fonio 445, 410 Indigobird, Goldbreast 443 Indigobird, Jambandu 443, 410 Indigobird, Peter’s Twinspot 450 Indigobird, Plateau 442, 410 Indigobird, Purple 448, 410 Indigobird, Quailfinch 441, 410 Indigobird, Village 436, 410 Indigobird, Wilson’s 439 Indigobird, Zambezi 450, 410

Linnet, Common 545, 475 Linnet, Warsangli 547, 475 Locust-Finch 390, 330

Malimbe, Blue-billed 93, 27 Malimbe, Cassin’s 82, 27 Malimbe, Crested 89, 27 Malimbe, Gola 78, 27 Malimbe, Ibadan 84, 27 Malimbe, Rachel’s 80, 27 Malimbe, Red-bellied 86, 27 Malimbe, Red-crowned 87, 27 Malimbe, Red-headed 95, 155 Malimbe, Red-vented 81, 27

Mannikin, Black-and-White 405, 347 Mannikin, Bronze 401, 347 Mannikin, Magpie 407, 347 Mannikin, Red-backed 405

Negrofinch, Chestnut-breasted 257, 219 Negrofinch, Grey-headed 255, 219 Negrofinch, Pale-fronted 253, 219 Negrofinch, White-breasted 259, 219 Oliveback, Grey-headed 270, 266 Oliveback, Shelley’s 265, 266 Oliveback, White-collared 268, 266 Oriole-Finch 531, 474

Paradise-Whydah, Broad-tailed 427, 395 Paradise-Whydah, Exclamatory 424, 395 Paradise-Whydah, Long-tailed 428, 395 Paradise-Whydah, Sahel 434, 394 Paradise-Whydah, Togo 426, 395 Petronia, Bush 42, 10 Petronia, Yellow-spotted 44, 10 Petronia, Yellow-throated 45, 10 Pytilia, Golden-backed 358, 266 Pytilia, Green-winged 354, 266 Pytilia, Orange-winged 358, 266 Pytilia, Red-billed 354, 266 Pytilia, Red-winged 352, 266 Pytilia, Yellow-winged 349, 266 Quailfinch, African 388, 330 Quailfinch, Black-chinned 386, 330 Quailfinch, Black-faced 384, 330 Quelea, Cardinal 204, 155 Quelea, Red-billed 207, 155 Quelea, Red-headed 205, 155 Redpoll, Common 545, 475 Rockfinch, Pale 551, 522 Rosefinch, Scarlet 550, 475 Sakabula 246 Seedcracker, Black-bellied 318, 283 Seedcracker, Crimson 322, 283 Seedcracker, Lesser 323, 283 Seedeater, Black-eared 504, 459 Seedeater, Black-throated 471

Seedeater, Brown-rumped 510, 459 Seedeater, Mennell’s 504 Seedeater, Prı´ncipe 500, 459 Seedeater, Reichard’s 509, 459 Seedeater, Reichenow’s 473, 411 Seedeater, Salvadori’s 486, 458 Seedeater, Streaky 495, 459 Seedeater, Streaky-headed 506, 459 Seedeater, Stripe-breasted 509 Seedeater, Tanzania 499, 459 Seedeater, Thick-billed 498, 459 Seedeater, White-rumped 470, 411 Seedeater, Yellow-browed 497, 459 Seedeater, Yellow-rumped 476, 411 Seedeater, Yellow-throated 484, 458 Serin, Ankober 549, 475 Serin, European 516, 474 Serin, Red-fronted 520, 474 Serin, Reichenow’s 473 Serin, Syrian 518, 474 Serin, Tristram’s 518 Serin, White-throated 470 Serin, White-throated 476 Silverbill, African 412, 347 Silverbill, Grey-headed 400, 347 Siskin, Black-headed 519 Siskin, Cape 521, 474 Siskin, Drakensberg 525, 474 Siskin, Ethiopian 519, 474 Siskin, Eurasian 543, 475 Sparrow, Arabian Golden 40, 10 Sparrow, Cape 17, 11 Sparrow, Chestnut 34, 10 Sparrow, Desert 29, 11 Sparrow, Eurasian Tree 32, 11 Sparrow, Great 9, 11 Sparrow, House 20, 11 Sparrow, Java 415, 347 Sparrow, Kenya Rufous 13, 11 Sparrow, Kordofan Rufous 15, 11 Sparrow, Northern Grey-headed 2, 10 Sparrow, Pale Rock 551 Sparrow, Parrot-billed 6, 10 Sparrow, Rock 47, 10 Sparrow, Shelley’s Rufous 14, 11 Sparrow, Socotra 16, 11 Sparrow, Somali 31, 11 Sparrow, Southern Grey-headed 7, 10 Sparrow, Spanish 24, 11 Sparrow, Sudan Golden 36, 10 Sparrow, Swahili 7, 10 Sparrow, Swainson’s 4, 10 Sparrow-Weaver, Chestnut-backed 60, 26 Sparrow-Weaver, Chestnutcrowned 58, 26

663

664

INDEXES: FRENCH NAMES

Sparrow-Weaver, DonaldsonSmith’s 61, 26 Sparrow-Weaver, White-browed 56, 26

Twinspot, Brown 348, 267 Twinspot, Dusky 340, 267 Twinspot, Dybowski’s 338, 267 Twinspot, Green 275, 283 Twinspot, Peters’s 342, 283 Twinspot, Pink-throated 344, 267 Twinspot, Red-throated 342, 283

Waxbill, Anambra 293, 267 Waxbill, Black-cheeked 311, 202 Waxbill, Black-crowned 305, 202 Waxbill, Black-faced 309, 202 Waxbill, Black-headed 307, 202 Waxbill, Black-lored 304, 202 Waxbill, Black-rumped 298, 282 Waxbill, Blue 325, 330 Waxbill, Cinderella 290, 267 Waxbill, Common 301, 202 Waxbill, Crimson-rumped 297, 282 Waxbill, Fawn-breasted 292, 267 Waxbill, Grey 289, 267 Waxbill, Kandt’s 308, 202 Waxbill, Lavender 287, 267 Waxbill, Orange-breasted 392 Waxbill, Orange-cheeked 295, 267 Waxbill, Southern Blue 326, 330 Waxbill, Swee 272, 267 Waxbill, Violet-eared 335, 230 Waxbill, Yellow-bellied 273, 267 Waxbill, Zebra 392, 282 Weaver, Baglafecht 106, 74 Weaver, Bannerman’s 109, 74 Weaver, Bar-winged 192, 154 Weaver, Bates’s 110, 75 Weaver, Bertram’s 104, 75 Weaver, Black-billed 120, 75 Weaver, Black-capped Social 70, 74 Weaver, Black-chinned 108, 74 Weaver, Black-headed 164 Weaver, Black-necked 115, 75 Weaver, Bob-tailed 213, 155 Weaver, Bocage’s 125, 90 Weaver, Brown-capped 186, 154 Weaver, Cape 122, 90

Weaver, Chestnut 177, 138 Weaver, Chestnut-backed Sparrow60, 26 Weaver, Chestnut-crowned Sparrow- 58, 26 Weaver, Cinnamon 179, 154 Weaver, Clarke’s 172, 154 Weaver, Compact 196, 154 Weaver, Dark-backed 183, 139 Weaver, Donaldson-Smith’s Sparrow- 61, 26 Weaver, Eastern Golden 126, 90 Weaver, Fox’s 160, 138 Weaver, Giant 170, 138 Weaver, Golden Palm- 132, 90 Weaver, Golden-backed 175, 138 Weaver, Golden-naped 180, 154 Weaver, Grey-headed Social 67, 74 Weaver, Grosbeak 198 Weaver, Heuglin’s Masked 153, 91 Weaver, Holub’s Golden 128, 90 Weaver, Juba 173, 138 Weaver, Katanga Masked 151, 91 Weaver, Kilombero 140, 91 Weaver, Lake Lufira 152, 91 Weaver, Lesser Masked 142, 91 Weaver, Little 111, 75 Weaver, Loango 114, 75 Weaver, Maxwell’s Black 182, 139 Weaver, Northern Brownthroated 137, 91 Weaver, Northern Masked 141, 91 Weaver, Olive-headed 189, 154 Weaver, Orange 130, 90 Weaver, Parasitic 417 Weaver, Preuss’s 188, 154 Weaver, Prı´ncipe Golden 134, 90 Weaver, Red-billed Buffalo- 52, 26 Weaver, Red-headed 97, 155 Weaver, Rufous-tailed 62, 26 Weaver, Ru ¨ ppell’s 157, 91 Weaver, Sa˜o Tome´ 193, 139 Weaver, Scaly-fronted 65, 26 Weaver, Slender-billed 112, 75 Weaver, Sociable 72, 74 Weaver, Southern Brownthroated 135, 90 Weaver, Southern Masked 145, 91 Weaver, Speckle-fronted 64, 26 Weaver, Spectacled 117, 75 Weaver, Speke’s 158, 138

Weaver, Spotted-backed 164 Weaver, Strange 121, 75 Weaver, Streaked 195, 154 Weaver, Tanganyika Masked 150, 91 Weaver, Taveta Golden 133, 90 Weaver, Thick-billed 198, 219 Weaver, Usambara 191, 154 Weaver, Vieillot’s Black 161, 139 Weaver, Village 164, 138 Weaver, Vitelline Masked 148, 91 Weaver, Weyns’s 171, 139 Weaver, White-billed Buffalo- 50, 26 Weaver, White-browed Sparrow- 56, 26 Weaver, White-headed Buffalo- 54, 26 Weaver, Yellow-backed 174, 138 Weaver, Yellow-capped 187, 154 Weaver, Yellow-legged 104, 139 Weaver, Yellow-mantled 181, 139 Whydah, Straw-tailed 433, 394 Whydah, Broad-tailed Paradise- 427, 395 Whydah, Exclamatory Paradise- 424, 395 Whydah, Long-tailed Paradise- 428, 395 Whydah, Pin-tailed 420, 394 Whydah, Sahel Paradise- 423, 395 Whydah, Shaft-tailed 434 Whydah, Steel-blue 431, 394 Whydah, Togo Paradise- 426, 395 Widowbird, Fan-tailed 242, 203 Widowbird, Hartlaub’s Marsh 244, 218 Widowbird, Jackson’s 249, 218 Widowbird, Long-tailed 246, 218 Widowbird, Mountain Marsh 246, 218 Widowbird, Red-collared 239, 203 Widowbird, White-winged 235, 203 Widowbird, Yellow-mantled 237, 203 Widowfinch, Steel-blue 436

Amadine `a teˆte rouge 381, 347 Amadine cou-coupe´ 379, 347 Amarante `a ventre noir 369, 331

Amarante de Jameson 373, 346 Amarante de Reichenow 378, 346 Amarante de roche 376, 346

Yellowhammer 589, 538

French Names Alecto `a bec blanc 50, 26 Alecto `a bec rouge 52, 26 Alecto `a teˆte blanche 54, 26

664

INDEXES: FRENCH NAMES

Sparrow-Weaver, DonaldsonSmith’s 61, 26 Sparrow-Weaver, White-browed 56, 26

Twinspot, Brown 348, 267 Twinspot, Dusky 340, 267 Twinspot, Dybowski’s 338, 267 Twinspot, Green 275, 283 Twinspot, Peters’s 342, 283 Twinspot, Pink-throated 344, 267 Twinspot, Red-throated 342, 283

Waxbill, Anambra 293, 267 Waxbill, Black-cheeked 311, 202 Waxbill, Black-crowned 305, 202 Waxbill, Black-faced 309, 202 Waxbill, Black-headed 307, 202 Waxbill, Black-lored 304, 202 Waxbill, Black-rumped 298, 282 Waxbill, Blue 325, 330 Waxbill, Cinderella 290, 267 Waxbill, Common 301, 202 Waxbill, Crimson-rumped 297, 282 Waxbill, Fawn-breasted 292, 267 Waxbill, Grey 289, 267 Waxbill, Kandt’s 308, 202 Waxbill, Lavender 287, 267 Waxbill, Orange-breasted 392 Waxbill, Orange-cheeked 295, 267 Waxbill, Southern Blue 326, 330 Waxbill, Swee 272, 267 Waxbill, Violet-eared 335, 230 Waxbill, Yellow-bellied 273, 267 Waxbill, Zebra 392, 282 Weaver, Baglafecht 106, 74 Weaver, Bannerman’s 109, 74 Weaver, Bar-winged 192, 154 Weaver, Bates’s 110, 75 Weaver, Bertram’s 104, 75 Weaver, Black-billed 120, 75 Weaver, Black-capped Social 70, 74 Weaver, Black-chinned 108, 74 Weaver, Black-headed 164 Weaver, Black-necked 115, 75 Weaver, Bob-tailed 213, 155 Weaver, Bocage’s 125, 90 Weaver, Brown-capped 186, 154 Weaver, Cape 122, 90

Weaver, Chestnut 177, 138 Weaver, Chestnut-backed Sparrow60, 26 Weaver, Chestnut-crowned Sparrow- 58, 26 Weaver, Cinnamon 179, 154 Weaver, Clarke’s 172, 154 Weaver, Compact 196, 154 Weaver, Dark-backed 183, 139 Weaver, Donaldson-Smith’s Sparrow- 61, 26 Weaver, Eastern Golden 126, 90 Weaver, Fox’s 160, 138 Weaver, Giant 170, 138 Weaver, Golden Palm- 132, 90 Weaver, Golden-backed 175, 138 Weaver, Golden-naped 180, 154 Weaver, Grey-headed Social 67, 74 Weaver, Grosbeak 198 Weaver, Heuglin’s Masked 153, 91 Weaver, Holub’s Golden 128, 90 Weaver, Juba 173, 138 Weaver, Katanga Masked 151, 91 Weaver, Kilombero 140, 91 Weaver, Lake Lufira 152, 91 Weaver, Lesser Masked 142, 91 Weaver, Little 111, 75 Weaver, Loango 114, 75 Weaver, Maxwell’s Black 182, 139 Weaver, Northern Brownthroated 137, 91 Weaver, Northern Masked 141, 91 Weaver, Olive-headed 189, 154 Weaver, Orange 130, 90 Weaver, Parasitic 417 Weaver, Preuss’s 188, 154 Weaver, Prı´ncipe Golden 134, 90 Weaver, Red-billed Buffalo- 52, 26 Weaver, Red-headed 97, 155 Weaver, Rufous-tailed 62, 26 Weaver, Ru ¨ ppell’s 157, 91 Weaver, Sa˜o Tome´ 193, 139 Weaver, Scaly-fronted 65, 26 Weaver, Slender-billed 112, 75 Weaver, Sociable 72, 74 Weaver, Southern Brownthroated 135, 90 Weaver, Southern Masked 145, 91 Weaver, Speckle-fronted 64, 26 Weaver, Spectacled 117, 75 Weaver, Speke’s 158, 138

Weaver, Spotted-backed 164 Weaver, Strange 121, 75 Weaver, Streaked 195, 154 Weaver, Tanganyika Masked 150, 91 Weaver, Taveta Golden 133, 90 Weaver, Thick-billed 198, 219 Weaver, Usambara 191, 154 Weaver, Vieillot’s Black 161, 139 Weaver, Village 164, 138 Weaver, Vitelline Masked 148, 91 Weaver, Weyns’s 171, 139 Weaver, White-billed Buffalo- 50, 26 Weaver, White-browed Sparrow- 56, 26 Weaver, White-headed Buffalo- 54, 26 Weaver, Yellow-backed 174, 138 Weaver, Yellow-capped 187, 154 Weaver, Yellow-legged 104, 139 Weaver, Yellow-mantled 181, 139 Whydah, Straw-tailed 433, 394 Whydah, Broad-tailed Paradise- 427, 395 Whydah, Exclamatory Paradise- 424, 395 Whydah, Long-tailed Paradise- 428, 395 Whydah, Pin-tailed 420, 394 Whydah, Sahel Paradise- 423, 395 Whydah, Shaft-tailed 434 Whydah, Steel-blue 431, 394 Whydah, Togo Paradise- 426, 395 Widowbird, Fan-tailed 242, 203 Widowbird, Hartlaub’s Marsh 244, 218 Widowbird, Jackson’s 249, 218 Widowbird, Long-tailed 246, 218 Widowbird, Mountain Marsh 246, 218 Widowbird, Red-collared 239, 203 Widowbird, White-winged 235, 203 Widowbird, Yellow-mantled 237, 203 Widowfinch, Steel-blue 436

Amadine `a teˆte rouge 381, 347 Amadine cou-coupe´ 379, 347 Amarante `a ventre noir 369, 331

Amarante de Jameson 373, 346 Amarante de Reichenow 378, 346 Amarante de roche 376, 346

Yellowhammer 589, 538

French Names Alecto `a bec blanc 50, 26 Alecto `a bec rouge 52, 26 Alecto `a teˆte blanche 54, 26

INDEXES: FRENCH NAMES

Amarante du Kulikoro 375, 346 Amarante du Se´ne´gal 360, 331 Amarante fonce´ 371, 346 Amarante masque´ 367, 331 Amarante nitidule 366, 331 Amarante pointe´ 364, 331 Astrild `a croupion rose 297, 282 Astrild `a joues noires 272, 267 Astrild `a joues orange 295, 267 Astrild `a masque noir 304, 282 Astrild `a moustaches 309, 282 Astrild `a poitrine fauve 292, 267 Astrild `a queue noire 289, 267 Astrild `a teˆte noire 307, 282 Astrild `a ventre jaune 273, 267 Astrild cendre´ 298, 282 Astrild cendrillon 290, 267 Astrild de Kandt 308, 282 Astrild des fe´es 311, 282 Astrild du Niger 293, 267 Astrild nonnette 305, 282 Astrild ondule´ 301, 282 Astrild queue-de-vinaigre 287, 267 Astrild-caille `a face noire 384, 330 Astrild-caille `a gorge noire 386, 330 Astrild-caille `a gorge rouge 390, 330 Astrild-caille `a lunettes 388, 330

Beaumarquet `a ailes jaunes 349, 266 Beaumarquet `a bec rouge 354, 266 Beaumarquet `a dos jaune 358, 266 Beaumarquet aurore 352, 266 Beaumarquet melba 354, 266 Bec-croise ´ des sapins 562, 522 Bengali rouge 398, 282 Bengali ze´bre´ 392, 282 Bouvreuil pivoine 559, 552 Bruant `a calotte blanche 588, 538 Bruant `a poitrine dore´e 567, 523 Bruant `a ventre jaune 572, 523 Bruant cannelle 573, 523 Bruant cendre´ 591, 539 Bruant cendrillard 594, 539 Bruant de Cabanis 565, 523 Bruant de Socotra 580, 538 Bruant de Somalie 571, 523 Bruant de Vincent 583, 538 Bruant des neiges 601, 539 Bruant des rochers 585, 538 Bruant des roseaux 596, 539 Bruant du Cap 582, 538 Bruant fou 587, 538 Bruant jaune 589, 538 Bruant melanocephale 598, 539 Bruant nain 596, 539

Bruant Bruant Bruant Bruant Bruant

ortolan 592, 539 proyer 599, 539 rustique 595, 539 striole´ 577, 538 zizi 589, 538

Capucin `a teˆte grise 400, 347 Capucin bec d’argent 412, 347 Capucin bicolore 405, 347 Capucin nonnette 401, 347 Capucin pie 407, 347 Chardonneret ´ele´gant 541, 475 Combassou d’Astrild-caille 441, 410 Combassou de Barka 444, 410 Combassou de Codrington 450, 410 Combassou de Jambandu 443, 410 Combassou de Jos 442, 410 Combassou de Se´ne´ga 436, 410 Combassou de Wilson 439, 410 Combassou du Cameroun 445, 410 Combassou noir 447, 410 Combassou pourpre´ 448, 410 Cordonbleu `a joues rouges 326, 330 Cordonbleu cyanoce´phale 332, 330 Cordonbleu de l’Angola 325, 330 Cordonbleu grenadin 335, 330 Cordonbleu violace´ 334, 330

Dos-vert `a collier 268, 266 Dos-vert `a joues blanches 270, 266 Dos-vert `a teˆte noire 265, 266

Euplecte `a croupion jaune 233, 155 Euplecte `a diade`me 230, 202 Euplecte `a dos d’or 237, 203 Euplecte `a ´epaules blanches 235, 203 Euplecte `a ´epaules orange´es 242 Euplecte `a longue queue 246, 218 Euplecte de Gierow 227, 202 Euplecte de Jackson 249, 218 Euplecte de Zanzibar 223, 202 Euplecte des marais 244, 218 Euplecte dore´ 232, 155 Euplecte franciscain 215, 202 Euplecte ignicolore 217, 202 Euplecte monseigneur 224, 202 Euplecte montagnard 246, 218 Euplecte veuve-noire 239, 203 Euplecte vorabe´ 228, 155

Grand-verdier `a ailes d’or d’Arabie 534, 475 Grand-verdier `a ailes d’or de Somalie 535, 475 Grosbec `a front blanc 198, 219 Gros-bec cassenoyaux 559, 522

Histurgopse `a queue rouge 62, 26

Linotte de Warsangli 547, 475 Linotte me´lodieuse 545, 475 Linurge loriot 531, 474

Mahali `a calotte marron 58, 26 Mahali `a dos roux 60, 26 Mahali `a sourcils blancs 56, 26 Mahali de Donaldson 61, 26 Malimbe `a bec bleu 93, 27 Malimbe `a queue rouge 81, 27 Malimbe `a teˆte rouge 95, 27 Malimbe `a ventre rouge 86, 27 Malimbe couronne´ 87, 27 Malimbe d’Ibadan 84, 27 Malimbe de Cassin 82, 27 Malimbe de Gola 78, 27 Malimbe de Rachel 80, 27 Malimbe huppe´ 89, 27 Moineau `a point jaune 44, 10 Moineau blanc 29, 11 Moineau bride´ 45, 10 Moineau d’Arabie 40, 10 Moineau d’Emin 34, 10 Moineau de Socotra 16, 11 Moineau de Somalie 31, 11 Moineau de Swainson 4, 10 Moineau domestique 20, 11 Moineau dore´ 36, 10 Moineau espagnol 24, 11 Moineau friquet 32, 11 Moineau gris 2, 10 Moineau me´lanure 17, 11 Moineau paˆle 551, 522 Moineau perroquet 6, 10 Moineau roux de Kenya 13, 11 Moineau roux de Kordofan 15, 11 Moineau roux de Shelley 14, 11 Moineau soulcie 47, 10 Moineau soulcie, Petit 42, 10 Moineau sud-africain 7, 10 Moineau swahili 7, 10 Moineau, Grand 9, 11

Ne ´ospize de Sa˜o Tome´ 530, 475 Nigrette `a calotte grise 255, 219

665

666

INDEXES: FRENCH NAMES

Nigrette `a front jaune 253, 219 Nigrette `a ventre blanc 259, 219 Nigrette `a ventre roux 257, 219

Padda de Java 415, 347 Parmoptile `a front rouge 263, 219 Parmoptile `a gorge rousse 261, 219 Parmoptile de Jameson 264, 219 Pinson des arbres 452, 523 Pinson du Nord 454, 523 Pyre´neste gros-bec 322, 283 Pyre´neste ponceau 318, 283 Pyre´neste, Petit 323, 283

Re´publicain d’Arnaud 67, 74 Re´publicain de Cabanis 70, 74 Re´publicain social 72, 74 Roselin `a ailes roses 553, 522 Roselin cramoisi 550, 475 Roselin de Lichtenstein 555, 522 Roselin githagene 556, 552

Se´ne ´gali `a bec bleu 313, 283 Se´ne´gali de Jackson 279, 266 Se´ne´gali `a teˆte rouge 315, 283 Se´ne´gali `a ventre noire 338, 267 Se´ne´gali brun 348, 267 Se´ne´gali de Reichenow 281, 266 Se´ne´gali de Salvadori 280, 266 Se´ne´gali de Shelley 278, 266 Se´ne´gali de Verreaux 344, 283 Se´ne´gali enflamme´ 342, 283 Se´ne´gali sanguin 314, 283 Se´ne´gali sombre 340, 267 Se´ne´gali vert 275, 283 Serin `a calotte jaune 511, 459 Serin `a croupion blanc 470, 411 Serin `a croupion jaune 476, 411 Serin `a diade`me 463, 411 Serin `a face noire 465, 411 Serin `a front d’or 520, 474 Serin `a gorge blanche 494, 458 Serin `a gorge jaune 484, 458 Serin `a gorge noire 471, 411 Serin `a gros bec 487, 458 Serin `a poitrine citron 477, 458 Serin `a teˆte noire 519, 474 Serin `a trois raies 510, 459 Serin `a ventre blanc 483, 458 Serin alario du Cap 527, 474 Serin alario du Nord 529, 474 Serin bifascie´ 502, 459 Serin bride´ 497, 459

Serin cini 516, 474 Serin d’Abyssinie 460, 411 Serin d’Ankober 549, 475 Serin de Buchanan 488, 458 Serin de Burton 498, 459 Serin de Reichard 509, 459 Serin de Reichenow 473, 411 Serin de Sainte-He´le`n 491, 458 Serin de Salvadori 486, 458 Serin de Symons 525, 474 Serin des Kipe´nge´re 499, 459 Serin du Cap 513, 459 Serin du Koli 467, 411 Serin du Mozambique 479, 458 Serin est-africain 464, 411 Serin forestier 468, 411 Serin gris 506, 459 Serin oreillard 504, 459 Serin roux 500, 459 Serin soufre´ 489, 458 Serin strie´ 495, 459 Serin syriaque 518, 474 Serin totta 521, 474 Sizeron flamme´ 545, 475 Sporopipe quadrille 64, 26 Sporopipe squameux 65, 26

Tarin des aulnes 545, 475 Tisserin `a bec greˆle 114, 75 Tisserin `a cap jaune 187, 154 Tisserin `a cape brune 186, 154 Tisserin `a cou noir 115, 75 Tisserin `a dos d’or 175, 138 Tisserin `a gorge brune 135, 90 Tisserin `a gorge noire 137, 91 Tisserin `a lunettes 117, 75 Tisserin `a menton noir 108, 74 Tisserin `a nuque d’or 180, 154 Tisserin `a pieds jaunes 104, 139 Tisserin `a teˆte jaune 120, 75 Tisserin `a teˆte noire 174, 138 Tisserin `a teˆte olive 189, 154 Tisserin baglafecht 106, 74 Tisserin bicolore 183, 139 Tisserin cannelle 179, 154 Tisserin de Bannerman 109, 74 Tisserin de Bates 110, 75 Tisserin de Bertrand 104, 75 Tisserin de Bocage 125, 90 Tisserin de Clarke 172, 154 Tisserin de Fox 160, 138 Tisserin de Katanga 151, 91 Tisserin de Kilombero 140, 91 Tisserin de Maxwell 182, 139 Tisserin de montagne 121, 75

Tisserin de Pelzeln 112, 75 Tisserin de Preuss 188, 154 Tisserin de Prı´ncipe 134, 90 Tisserin de Reichard 150, 91 Tisserin de Ru ¨ ppell 157, 91 Tisserin de Ruwet 152, 91 Tisserin de Salvadori 173, 138 Tisserin de Sa˜o Tome´ 193, 139 Tisserin de Speke 158, 138 Tisserin de Taveta 133, 90 Tisserin de Vieillot 161, 139 Tisserin de Weyns 171, 139 Tisserin des Usambara 191, 154 Tisserin du Cap 122, 90 Tisserin du Nil 141, 91 Tisserin ´ecarlate 97, 155 Tisserin ge´ant 170, 138 Tisserin gendarme 164, 138 Tisserin gros-bec 196, 154 Tisserin jaune 126, 90 Tisserin malimbe 192, 154 Tisserin manyar 195, 154 Tisserin masque´ 145, 91 Tisserin masque´ austral 142, 91 Tisserin masque´ d’Heuglin 153, 91 Tisserin minule 111, 75 Tisserin orange 130, 90 Tisserin palmiste 132, 90 Tisserin roux 177, 138 Tisserin safran 128, 90 Tisserin tricolore 181, 139 Tisserin vitellin 148, 91 Tisserin-coucou 417, 394 Travailleur `a bec rouge 207, 155 Travailleur `a queue courte 213, 155 Travailleur `a teˆte rouge 205, 155 Travailleur cardinal 204, 155

Venturon montagnard 518, 474 Verdier d’Europe 536, 475 Veuve `a collier d’or 423, 395 Veuve de Chapin 427, 395 Veuve de Fischer 433, 394 Veuve de paradis 428, 395 Veuve dominicaine 420, 394 Veuve du Togo 426, 395 Veuve me´tallique 431, 394 Veuve nige´rienne 424, 395 Veuve royale 434, 394