The Birds of Africa Volume IV 9781472927019, 9780121373047

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ACKNOWLEDGEMENTS Once again we wish to express our sincere appreciation of the authors. Researching their subjects in field, museum and library, and writing species accounts to exacting standards, are extremely time-consuming. Authors' drafts have been subjected to the seemingly endless process of revision by colleagues, referees and editors. Our authors have borne all this with patience and goodwill for which, and for their diligence, we warmly thank them. During the preparation of species accounts authors have drawn heavily upon the special knowledge of numerous colleagues, who have given them access to unpublished and pre-published works and field notes, provided measurements and weights, given information on specimens, distribution and habits, copied library materials, and responded to a host of queries. Many colleagues have also given of their valuable time to referee the accounts. For all their help we take great pleasure in thanking: Passeriformes: J. Cracraft, S. L. Olson and R. J. Raikow; Eurylaimidae, A. Brosset, T. Butynski, C. Erard, J. Kalina and E. F. G. Smith; Pittidae, P. R. Fogarty, J. Jones and A. N. B. Masterson; Alaudidae, D. G. Allan, J. P. Angle, J. S. Ash, D. R. Aspinwall, H. Boyer, R. K. Brooke, T. Cassidy, C. Chappuis, M. Charron, P. A. Clancey, C. Clinning, J. F . R. Colebrook-Robjent, P. R. Colston, R. J. Dowsett, F. Dowsett-Lemaire, E. K. Dunn, R. A. Earle, J. H. Elgood, L. Freysen, J. D . Gerhart, S. M. Goodman, A. Harris, D. L. Harrison, J. A. Harrison, P. A. R. Hockey, P. Hogg, K. Hustler, M. P. S. Irwin, H . D . Jackson, A. C. Kemp, J. Komen, R. Liversidge, I. A. W. Macdonald, G. L. Maclean, A. P. Martin, R. Martin, G. R. McLachlan, S. J. Milton, J. E. Miskell, J. M. Mendelsohn, D. C. Moyer, G. Nikolaus, S. L. Olson, P. Perry, D. Rockingham-Gill, D. Schultz, W. R. Siegfried, J. C. Sinclair, R. Stjernstedt, W. R. Tarboton, C. J . Vernon and D . A. Zimmerman; Hirundinidae, P . D. Alexander-Marrack, D. G. Allan, J. S. Ash, D . R. Aspinwall, P. Becker, L. Birch, A. Brosset, T. Butynski, M. Carswell, P. Christy, P. R. Colston, H . Q. P. Crick, R. J. Dowsett, M. Dyer, R. A. Earle, J. H. Elgood, C. Erard, J. Farrand Jr, K. Gamble, R. M. Glen, S. M. Goodman, L. Grant, L. G. Grimes, D. N. Johnson, L. B. Kiff, B. Lamarche, A. D . Lewis, M. Louette, R. D. Medland, J. Millward, H. Morand, G. J. Morel, D. C. Moyer, V. Parker, D. J. Pearson, E. H. Penry, R. Penry, D. E. Pomeroy, the late A. Prigogine, J. Schmidt, M. L. Schmidt, W. Serle, N. J. Skinner, S. N. Stuart, W. R. Tarboton, A. K . Turner, S. J. Tyler, L. L. Urban, D. M. Ward, R. T. Wilson and R. Zusi; Motacillidae, J. S. Ash, D. R. Aspinwall, G. C. Backhurst, P. R. Colston, R. J. Dowsett, F. Dowsett-Lemaire, A. J. Harris, M. P. S. Irwin, M. G. Kelsey, G. L. Maclean, R. A. M. McVicker, M. E. D. Nhlane, T. B. Oatley, S. E. Piper, D. K. Read , J . C. Sinclair, M. Thevenot, S. J. Tyler, A. Vale and B. Wood; Campephagidae, J. S. Ash, D. R. Aspinwall, A. Brosset, C. Chappuis, P. R. Colston, W. Gatter, M. Louette, the late A. Prigogine, N. J. Skinner and J.-P. Vande weghe; Pycnonotidae, especially F. Dowsett-Lemaire (referee) and C. Chappuis, also D. Amadon, D. R. Aspinwall, P. Becker, W. P. Cane, P. A. Clancey, P. R. Colston, G. Cowles, T. M. Crowe, W. R. J . Dean, R. W. Dickerman, R. J. Dowsett, W. Gatter, L. B. Kiff, M. LeCroy, R. Liversidge, B. L. Monroe Jr, T. B. Oatley, S. L. Olson, D . J. Pearson, J. Plunkett, the late R. Plunkett, the late A. Prigogine, L. L. Short, C. G. Sibley, S. N. Stuart, P. B. Taylor, F . Vuilleumier, M. Walters, M. W. Woodcock and D. A. Zimmerman; Cinclidae, S. J. Ormerod and S. J. Tyler; Troglodytidae, M. Thevenot; Prunellidae, N. J. Davies; Turdidae, J. S. Ash, G. C. Backhurst, P. Becker, A. Brosset, T. Butynski, T. Cassidy, C. Chappuis, W. R. J. Dean, P. Devillers, F. Dowsett-Lemaire, E. K. Dunn, A. D. Forbes-Watson, K. Gamble, P . Giraudoux, A. Harris, K. Joysey, J. Kalina, M. Louette, D. Merrie, J . E. Miskell, G. J. Morel, G. Nikolaus, T. B. Oatley, D. J . Pearson, P. Steyn, W. R. Tarboton, P. B. Taylor, K. Thangavelu, Y. Thonnerieux, M. A. Traylor, D. A. Turner, E. O. Willis, M. W. Woodcock and D . A. Zimmerman. During the preparation of their accounts authors have naturally consulted all the available literature, but we wish to single out for special mention the two major works on which they have drawn most heavily: Cramp (1988) for North Africa and Maclean (1985) for southern Africa. Special thanks are due to Dr Linda Birch, Alexander Librarian at the Edward Grey Institute, and to L. R. Macaulay, for extensive help with the literature. Authors also owe a special debt of gratitude to C. Chappuis and F. Dowsett-Lemaire for providing copies of tape recordings, to M. P. S. Irwin for providing copies of his extensive bibliography, to K. Garrett and M. C. Wimer for providing detailed printouts of specimens in the Los Angeles County Museum of Natural History, to L. B. Kiff for providing lists of specimens in the Western Foundation of Vertebrate Zoology, and to EURING for providing records of ringing recoveries. Stuart Keith also wishes to give special thanks to P. B. and C. Taylor for their hospitality and assistance during an extended visit to the Natural History Museum at Tring, to R. A. Ranney for extended hospitality and in particular for providing space in her house for the final co-ordinating of this volume, and to A. D. Forbes-Watson and D. A. Turner for their help in the field in Kenya. Again we are greatly indebted to the many museums holding African avian collections. For study facilities and specimen loans we thank the Trustees and staff of: The American Museum of Natural History, Department of Ornithology; Augusta College Reese Library (inter-library loan service); Natural History Museum (London), Department of Ornithology (Tring); Cambridge University Museum of Zoology; Carnegie Museum of Natural History; Durban Museum; East London Museum; Field Museum of Natural History, Bird Division; Los Angeles County Museum, Bird Division; Merseyside County Museum, Liverpool; Musee Royal de l' Afrique Centrale, v

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ACKNOWLEDGEMENTS

Tervuren; Museum of Comparative Zoology, Harvard; Museum Alexander Koenig, Bonn; National Museum of Zimbabwe, Bulawayo; National Museums of Kenya, Nairobi; Percy Fitzpatrick Institute of African Ornithology; Philadelphia Academy of Natural Sciences; Smithsonian Institution, US National Museum of Natural History, Department of Ornithology; South African Museum; State Museum (Namibia); and Transvaal Museum, Bird Department. For copious assistance with literature we thank especially the Edward Grey Institute, Oxford University. For the loan of bird sound recordings we are grateful to the British Library of Wildlife Sounds, Cornell University Laboratory of Ornithology, and the Transvaal Museum Bird Department. We owe particular thanks to the following staff members of these institutions: A. Andors, G. Barrowclough, C. Blake, P. J. K. Burton, P. A. Clancey, P. Colston, G. Cowles, T. M. Crowe, J. Fitzpatrick, K. Garrett, F. B. Gill, J. Gulledge, A. Harris, M. P. S. Irwin, H. Adan Isack, H. D. Jackson, A. C. Kemp, R. Kettle, A. Knox, M. LeCroy, P. Lorber, M. Louette, J. Mendelsohn, P. Morgan, J. P. Myers, S. L. Olson, K. C. Parkes, R. A. Paynter, D. Read, H. Schifter, the late R. W. Schreiber, K.-L. Schuchmann, L. L. Short, W . R. Siegfried, J. C. Sinclair, D. Steadman, M. A. Traylor, F. Vuilleumier, M. Walters and D . S. Wood. We would like to single out for special mention P. R. Colston of the Natural History Museum at Tring, who has willingly supplied authors with information on numerous occasions. We acknowledge with thanks the following sources of material for the preparation of black and white line drawings: R. M. Bloomfield in 'Complete Book of Southern African Birds' (1989) (Oenanthe pileata); G. J. Broekhuysen in Bokmakierie (1961 ) (Hirundo rustica ); M. Brown (Ammomanes cincturus and Cercomela sordida); J. P. Busby in 'Birds of the Western Palearctic' (1988) (Alaemon alaudipes, Galerida cristata); J. P. Chapin in 'Birds of the Belgian Congo' (1953) (Smithornis rufolateralis); R. A. Cheke in Malimbus (1982) (Hirundo preussi); W. R. J. Dean (Pinarornis plumosus); A. R . Devez in Brosset and Erard (1986) (Smithornis rufolateralis), in Bioi. Gabonica (1971 ) (Andropadus virens, Phyllastrephus icterinus, Criniger calurus, Chlorocichla simplex), and in Terre Vie (1974) (Bieda eximia, Criniger barbatus); M. Dyer in Nyala (1988) (Hirundo atrocaerulea ); R. A. Earle (Hirundo cucullata) and in Navors. Mus. Bloemfontein (1985) (Hirundo spilodera); M. D. England in 'Birds of the Western Palearctic' (1988) (Cercotrichas galactotes); H. Eriksen and J. Eriksen (loan of photo library); P. R. Fogarty (Pitta angolensis); D. Foster (Alaemon alaudipes); C. H. Fry (Ammomanes spp. , Ramphocoris clotbey, Hirundo rustica, H. lucida, Neocossyphus spp., Cercomela spp.); P. J. Ginn in 'Birds of Botswana' (1979) (Anthus novaeseelandiae); E. Hosking in 'Birds of the World' (IPC) 6, Part 5 (Eremophila bilopha); R . A. Jensen in 'Birdlife of Southern Africa' (1979) (Smithornis capensis) and in 'Complete Book of Southern African Birds' (1989) (Namibornis herero); J. Jones (Pitta angolensis); G. Langsbury in 'Complete Book of Southern African Birds' (1989) (Tmetothylacus tenellus); C. Laubscher in 'Complete Book of Southern African Birds' (1989) (Coracina pectoralis); R. Liversidge, unpublished PhD thesis (1970) (Pycnonotus capensis); A. N. B. Masterson (Pitta angolensis); W . G. McIlleron in 'Complete Book of Southern African Birds' (1989) (Cercotrichas leucophrys ); H . Morand (Pseudochelidon eurystomina ); 'Newman's Birds of Southern Africa' (1983) (Cercomela spp.); D. J. Pearson (Luscinia spp., Phoenicurus spp. ); C. Rose in 'Birds of the Western Palearctic' (1988) (Cercomela melanura ); J. C. Sinclair: 'Field Guide to Birds of Southern Africa' (1984) (Oenanthe bifasciata , Cercomela sinuata); P. Steyn in Ostrich (Pinarornis plumosus), in 'Birdlife in Southern Africa' (1979) (Macrony x capensis) and in 'Complete Book of Southern African Birds' (1989) (Smithornis capensis); W. R. Tarboton in Sinclair: 'Field Guide to Birds of Southern Africa' (1984) (Oenanthe bifasciata); P. B. Taylor (Macronyx spp. ); 1. Trotignon/Jacana, in 'Birds of the World' (IPC) 6, Part 5 (Chersophilus duponti ); C. J. Uys in 'Halcyon Days' (1963) (Galerida magnirostris, Pycnonotus capensis); J . L. Viljoen in 'Complete Book of Southern African Birds' (1989) (Campephaga flava); and D. Waters (Hypocolius ampelinus, Cercotrichas podobe) . All colour plates in this volume are the work of Martin Woodcock, also the monochrome paintings illustrating plumage topography and bill shapes and details of head feathering and ornamentation in the Introduction. All black and white line illustrations except as noted below were drawn by Ian Willis. Many drawings are based on sketches he made in the field. Like the authors, the artists have accommodated all referees' and editors' demands, and we are particularly grateful to them for their skill and ready co-operation. The following line drawings in the text are by Martin Woodcock: Ixonotus guttatus, and tail patterns of some cuckoo-shrikes Campephaga . Martin Woodcock has made further extensive studies in the field in Africa, and these have been invaluable during the preparation of the colour plates. He is grateful to all those friends and correspondents who have kindly given advice, criticism, help and photographs; in particular to the Editors and authors, and to his wife Barbara for her support both at home and in the field. Many people in Africa have generously offered hospitality and support in many ways, especially A. Boswell, N. and E. Baker, W. and N. Cooper, M. and E. Coverdale, Mrs J. Hartley, K. Howell, and D. and J. Moyer. He is especially indebted to the Trustees and staff of the Natural History Museum, Department of Ornithology at Tring for the loan of specimens and for their help. Ian Willis likewise wishes to thank R. McGowan of the Royal Museums of Scotland for his help in providing specimens. Again it is our pleasure to thank C. Chappuis for preparing the acoustic references, Lois Urban for indexing the book, our wives Sallyann Keith, Lois Urban and Kathie Fry for their ever-present support and encouragement, and lastly Andrew Richford, a Senior Editor at Academic Press Ltd, who continues to oversee the project with his customary skill, enthusiasm and tact. May 1992

Stuart Keith Emil K. Urban C. Hilary Fry

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CONTENTS

Authors' Contributions to Shared Families ALAUDIDAE Dean and Keith: Mirafra (except M. pulpa, M. hypermetra, M. ashi, M. degodiensis and M. poecilosterna), Heteromirafra ruddi and H. archeri, Certhilauda, Pinarocorys, Chersomanes, Ammomanes deserti and A. grayi, Calandrella cinerea, Spizocorys, Eremalauda, Pseudalaemon, Galerida modesta and G. magnirostris, E remopterix australis, E. leucotis and E. verticalis. Fry: Mirafra ashi and M. degodiensis, Heteromirafra sidamoensis, Alaemon, Rhamphocoris, Melanocorypha, Calandrella brachydactyla and C. rufescens, Chersophilus, Galerida cristata and G. malabarica, Lullula, Alauda and Eremophila. Keith: Calandrella somalica. Lack: Mirafra pulpa, M. hypermetra and M. poecilosterna, Eremopterix nigriceps, E. signata and E. leucopareia.

HIRUNDINIDAE Fry: Pseudochelidon, Phedina, Riparia, Pseudhirundo, Hirundo daurica, H. fuliginosa, H. preussi, H. rufigula, H. spilodera, H. perdita, H. fuligula, H. rupestris, H. rustica and Delichon. Urban and Keith: Psalidoprocne, Hirundo semirufa, H. senegalensis, H. abyssinica, H. cucullata, H. atrocaerulea, H. nigrorufa, H. smithii, H. nigrita, H. leucosoma, H. megaensis, H. dimidiata, H. aethiopica, H. albigularis, H. angolensis and H. lucida.

MOTACILLIDAE Fry: Motacilla. Pearson: Tmetothylacus, Anthus. Taylor: Macronyx.

TURDIDAE Dean: Cichladusa. Erard: Neocossyphus. Fry: Erithacus, Cossypha albicapilla, Pinarornis, Cercotrichas galactotes and C. podobe, Cercomela, Myrmecocichla cinnamomeiventris and M. semirufa. Keith: Alethe. Keith and Jensen: Namibornis. Lack: Myrmecocichla (except M. cinnamomeivenrtis and M. semirufa). Lack and Haas: Myrmecocichla aethiops. Oatley, Fry, Keith and Tye: Pogonocichla, Swynnertonia, Stiphrornis, Sheppardia (except S. montana, S. lowei), Cossyphicula, Cossypha (except C. isabellae, C. archeri, C. anomala and C. albicapilla), Xenocopsychus, Cercotrichas (except C. galactotes and C. podobe). Pearson: Luscinia, Irania, Phoenicurus, Saxicola torquata and S. rubetra. Stuart: Sheppardia montana and S. lowei, Cossypha isabellae, C. archeri and C. anomala, Modulatrix, Arcanator. Tye: Saxicola bifasciata, Oenanthe.

LIST OF PLATES Plate

Facing Page

Plate

Facing Page

1 Broadbills and pittas

32

17 Greenbuls (Andropadus)

304

2 Larks (northern Africa)

33

18 Greenbuls (Andropadus, Phyllastrephus)

305

3 Larks (West, central and East Africa)

48

19 Greenbuls (Phyllastrephus)

320

4 Larks (West, central and East Africa)

49

20 Greenbuls (Chlorocichla, Criniger, BIeda)

321

5 Larks (southern Africa)

96

21

Greenbuls (remainder) . .

352

6 Larks (remainder)

97

22

Thrushes (mainly Sheppardia)

353

7 Swallows, martins

112

23 Thrushes (robins, Alethe)

368

8 Swallows ..

113

24 Thrushes (robin-chats) . .

369

9 Swallows, saw-wings

160

25 Thrushes (robin-chats and others)

400

10

Swallows, martins

161

26 Thrushes ( juveniles)

401

11

Wagtails ..

176

27 Thrushes (scrub-robins and others)

416

177

28 Thrushes (Phoenicurus, Saxicola)

417

Pipits

240

29 Wheatears

448

14 Longclaws

241

30 Wheatears

449

15 Cuckoo-Shrikes

256

31 Chats (Cercomela)

464

16 Waxwing, Hypocolius, Dipper, Wren,

257

32 Chats (Myrmecocichla )

465

12 Pipits, Sharpe's Longclaw l3

accentors, Pycnonotus bulbuls

IX

Plate 1

s.

Smithornis rufolateralis bUdOngo7~ ~S iS

s.

c. capensis

c. capensis

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African Green Broadbill (p. 2) Pseudocalyptomena graueri

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Grey-headed Broadbill (p . 4)

32 1 5cm

Plate 2

~

Dunn's Lark (p . 89) Eremalauda dunni dunni

Bar-tailed Lark (p. 71) Ammomanes cincturus arenicolor

~/? Kordofan Lark (p . 20)

Desert Lark (p. 73) Ammomanes deserti deserti

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Lesser Short-toed Lark (p. 80) Calandrel/a rufescens minor

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Rusty Buoh.Lark (p . 36)

~~;~;ya

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Thick-billed Lark (p . 65) Ramphocoris clot-bey

Lesser Hoopoe-Lark (p . 64) Alaemon hamertoni hamertoni Bimaculated Lark (p . 69)

MS)~ roj",,",

~

\

Temm;nck' o Homed Eremophila bilopha r-/

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~

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Calandra Lark (p . 67) M elanocorypha calandra calandra

~\ #-_ -__ _

Thekla Lark (p. 103)

Gal~ro ,upaflu~ a ~

)

Horned Lark (p . 121) Eremophila alpestris atlas

\~

~

~

Cre"ed Lark (p. 100) -

~

~/

Woodlark (p.105) Lul/ula arborea pal/ida

/

~

Eurasian Skylark (p . 107) Alauda arvensis harterti

~ .~

Black-crowned Sparrow-Lark (p. 1 Eremopterix nigriceps albi/rons I

6 in

, ----------------------------~I 15 em

33

Plate 3 White-tai~ed

Bush-Lark (p. 18) Mlrafra albicauda

~

~

Friedmann's Lark (p.21) Mirafra pulpa

Wil1ia~s ' s

Lark (p.20) Mlrafra williamsi

M. a. athi

Red-winged B us h -Lark \ (p . 24) . Mlrafra hypermetra hypermetra

Ash's . Lark (p. 26) M lrafra ashi

/

~omali

Lark (p . 25) Mlrafra somalica somalica

~

)7~7

I

c~:~a~; 34)~

Archer's Lark (p. 44) H eteromirafra archeri (

~~\"

~\

\

,~ Glllet~

Pink-~reasted Lark (p. 39)

M,,"Fa po"i/a"",,"

I.

s Lark (p. 37)

M"aj," gilktti

38)~

Degodi Lark (p . Mirafra degodiensis

~~

~C;£~~di ' -z/l;: ~

Rufous-romped Lark (p 56) Pin", a,ary' ,-......, ._.":~ .......~./"y.,. , ," ~

tAMEROCJ

f2 Bioko. \. ..-! ~ 'UGANDA) \Iv \. EQUATORIAL .:::---------: . . . .I R Con 0 GUINEA """t.? 0 f KEN YA SAO TOME GABON ~ 0 RWtXNDA-+- ''Y ........ .J1.-' ~ 15',f,: \. ..,. ' . Pogolu · (,0 . 0$0/ ZAIRE ~ . .P \. BURUNDI 'Pem¢a

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~ANZANIA

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SOUTH AFRICA

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o Fig. 2. Updated political map.

200 400 600 800 1000 miles I

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500

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1000

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1500km

INTRODUCTION

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ETHIOPIA

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MOZAMBIQUE

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Fig. 3. Climatic zones of East Africa (Uganda, Kenya, Tanzania), referred to under specific Laying Dates (from Brown and Britton 1980 after J. F. Griffiths, Climatic zones of East Africa, E. Afr. Agrie.]. 1958, 179-185).

Xlll

Topography of African Passerine Birds I

Uppertail-

Crown Forecrown

T6 Lores

Secondaries

Chin

Median coverts Greater primary Lower breast _ UPPERWINGCOVERTS

Primari es

Greater coverts

daries

Upper belly----

UNDERWING-COVERTS under primary

pg

P8

P5 Superciliary stripe Undertail -

T6

T5

Wattled orbital ring (skin)

Topography of African Passerine Birds II Forecrown crest

Mouth spots Orbital ring (featherlets)

Recurved loral crest

/

Filoplumes

T5

T2

Eye wattle

Tl

Ornamental plumes

Breast band or pectoral band

mandible knob Bare skin Cutting edges

Median crown stripe Lateral crown stripe

\

. u:..+-_-Malar stripe

Central streamers (Tl )

This page intentionally left blank

Order

PASSERIFORMES 'Perching birds', mostly 'songbirds', much the largest order of birds, embracing approximately 5300 species worldwide, or well over half of all living birds; commonly called passerines for short. Cosmopolitan. Land-birds, only a very few species feeding in (fresh) water; mainly small, a few medium-sized - ravens Corvus spp. among the largest. Foot adapted for perching; always 4 toes, the hallux enlarged, directed backwards and not reversible; toes never webbed. In most species, syrinx adapted for singing; syrinx usually tracheo-bronchial with complex and varied musculature, but tracheal in 4 Neotropical families, and simple in a few groups. Wing eutaxic. Primaries 10, the outer 1 reduced or vestigial, or 9 in pipits, swallows, finches, buntings, some larks, and a few New World families . 12 tailfeathers. Aegithognathous palate, 'passerine' tensor propatagialis brevis, bundled spermatozoa with coiled head and large acrosome, and type VII deep plantar tendons. Young nidicolous, hatching blind, naked and helpless. Parental care well developed; but 3 groups brood-parasitic (2 in Africa). Passeriformes are a monophyletic group (Raikow 1982). Plumages, bill form and life-styles are very varied; 82 families recognized by Campbell and Lack (1985), many biologically distinctive. Evolutionary convergence rife, hence relationships, boundaries and true composition of families often questionable. Morphological and DNA-hybridization studies confirm that pittas, broadbills, tyrant-flycatchers, antbirds, ovenbirds, gnateaters and tapaculos form a natural assemblage (Suborder Tyranni - most families New World). They also suggest that 'true' songbirds of Suborder Passeri (Oscines) are related among themselves in ways radically different from those recognized conventionally (Sibley and Ahlquist 1990). In the main, classification of Passeriformes and sequence of passerine families adopted here follow those of Mayr and Greenway (1956) and Campbell and Lack (1985 ). However, subordinal nomenclature, and classification of Suborder Tyranni, follow Sibley et al. (1988 ).

Suborder TYRANNI (Suboscines) More or less synonymous with Deutero-oscines, Clamatores, Suboscines, Oligomyodi and Tyranniformes: 10- 14 families embracing c. 1100 species. Syrinx mesomyodian, with not more than 3 pairs of intrinsic muscles (Ames 1971); form of ear ossicle or stapes unique (except in New Zealand wrens: Feduccia 1980). Contains 3 infraorders - 1 large and Neotropical (Tyrannides) and 2 small and Paleotropical (Acanthisittides, New Zealand wrens, and Eurylaimides).

Infraorder EURYLAIMIDES Pittas Pittidae, broadbills Eurylaimidae, and asities Philepittidae (confined to Madagascar) (Olson 1971, Raikow 1982).

Superfamily EURYLAIMOIDEA: broadbills, aSlt1es Family EURYLAIMIDAE: broadbills Small to medium birds (weight range 20-160 g) with strikingly-coloured plumage. Bill moderately or extremely wide; outermost primary long, wing rounded; tail short or medium, more or less graduated. Tarsus with scutes anteriorly and small hexagonal scales posteriorly; toes 3 and 4 united basally. Main artery of leg is ischiatic. Non-furcate sterna spinalis. Tendon of flexor hallucis longus connected by vinculum to tendon of flexor digitorum longus. Inhabit humid forests; sedentary, somewhat sluggish; eat plant and some animal material. 1

Order

PASSERIFORMES 'Perching birds', mostly 'songbirds', much the largest order of birds, embracing approximately 5300 species worldwide, or well over half of all living birds; commonly called passerines for short. Cosmopolitan. Land-birds, only a very few species feeding in (fresh) water; mainly small, a few medium-sized - ravens Corvus spp. among the largest. Foot adapted for perching; always 4 toes, the hallux enlarged, directed backwards and not reversible; toes never webbed. In most species, syrinx adapted for singing; syrinx usually tracheo-bronchial with complex and varied musculature, but tracheal in 4 Neotropical families, and simple in a few groups. Wing eutaxic. Primaries 10, the outer 1 reduced or vestigial, or 9 in pipits, swallows, finches, buntings, some larks, and a few New World families . 12 tailfeathers. Aegithognathous palate, 'passerine' tensor propatagialis brevis, bundled spermatozoa with coiled head and large acrosome, and type VII deep plantar tendons. Young nidicolous, hatching blind, naked and helpless. Parental care well developed; but 3 groups brood-parasitic (2 in Africa). Passeriformes are a monophyletic group (Raikow 1982). Plumages, bill form and life-styles are very varied; 82 families recognized by Campbell and Lack (1985), many biologically distinctive. Evolutionary convergence rife, hence relationships, boundaries and true composition of families often questionable. Morphological and DNA-hybridization studies confirm that pittas, broadbills, tyrant-flycatchers, antbirds, ovenbirds, gnateaters and tapaculos form a natural assemblage (Suborder Tyranni - most families New World). They also suggest that 'true' songbirds of Suborder Passeri (Oscines) are related among themselves in ways radically different from those recognized conventionally (Sibley and Ahlquist 1990). In the main, classification of Passeriformes and sequence of passerine families adopted here follow those of Mayr and Greenway (1956) and Campbell and Lack (1985 ). However, subordinal nomenclature, and classification of Suborder Tyranni, follow Sibley et al. (1988 ).

Suborder TYRANNI (Suboscines) More or less synonymous with Deutero-oscines, Clamatores, Suboscines, Oligomyodi and Tyranniformes: 10- 14 families embracing c. 1100 species. Syrinx mesomyodian, with not more than 3 pairs of intrinsic muscles (Ames 1971); form of ear ossicle or stapes unique (except in New Zealand wrens: Feduccia 1980). Contains 3 infraorders - 1 large and Neotropical (Tyrannides) and 2 small and Paleotropical (Acanthisittides, New Zealand wrens, and Eurylaimides).

Infraorder EURYLAIMIDES Pittas Pittidae, broadbills Eurylaimidae, and asities Philepittidae (confined to Madagascar) (Olson 1971, Raikow 1982).

Superfamily EURYLAIMOIDEA: broadbills, aSlt1es Family EURYLAIMIDAE: broadbills Small to medium birds (weight range 20-160 g) with strikingly-coloured plumage. Bill moderately or extremely wide; outermost primary long, wing rounded; tail short or medium, more or less graduated. Tarsus with scutes anteriorly and small hexagonal scales posteriorly; toes 3 and 4 united basally. Main artery of leg is ischiatic. Non-furcate sterna spinalis. Tendon of flexor hallucis longus connected by vinculum to tendon of flexor digitorum longus. Inhabit humid forests; sedentary, somewhat sluggish; eat plant and some animal material. 1

Order

PASSERIFORMES 'Perching birds', mostly 'songbirds', much the largest order of birds, embracing approximately 5300 species worldwide, or well over half of all living birds; commonly called passerines for short. Cosmopolitan. Land-birds, only a very few species feeding in (fresh) water; mainly small, a few medium-sized - ravens Corvus spp. among the largest. Foot adapted for perching; always 4 toes, the hallux enlarged, directed backwards and not reversible; toes never webbed. In most species, syrinx adapted for singing; syrinx usually tracheo-bronchial with complex and varied musculature, but tracheal in 4 Neotropical families, and simple in a few groups. Wing eutaxic. Primaries 10, the outer 1 reduced or vestigial, or 9 in pipits, swallows, finches, buntings, some larks, and a few New World families . 12 tailfeathers. Aegithognathous palate, 'passerine' tensor propatagialis brevis, bundled spermatozoa with coiled head and large acrosome, and type VII deep plantar tendons. Young nidicolous, hatching blind, naked and helpless. Parental care well developed; but 3 groups brood-parasitic (2 in Africa). Passeriformes are a monophyletic group (Raikow 1982). Plumages, bill form and life-styles are very varied; 82 families recognized by Campbell and Lack (1985), many biologically distinctive. Evolutionary convergence rife, hence relationships, boundaries and true composition of families often questionable. Morphological and DNA-hybridization studies confirm that pittas, broadbills, tyrant-flycatchers, antbirds, ovenbirds, gnateaters and tapaculos form a natural assemblage (Suborder Tyranni - most families New World). They also suggest that 'true' songbirds of Suborder Passeri (Oscines) are related among themselves in ways radically different from those recognized conventionally (Sibley and Ahlquist 1990). In the main, classification of Passeriformes and sequence of passerine families adopted here follow those of Mayr and Greenway (1956) and Campbell and Lack (1985 ). However, subordinal nomenclature, and classification of Suborder Tyranni, follow Sibley et al. (1988 ).

Suborder TYRANNI (Suboscines) More or less synonymous with Deutero-oscines, Clamatores, Suboscines, Oligomyodi and Tyranniformes: 10- 14 families embracing c. 1100 species. Syrinx mesomyodian, with not more than 3 pairs of intrinsic muscles (Ames 1971); form of ear ossicle or stapes unique (except in New Zealand wrens: Feduccia 1980). Contains 3 infraorders - 1 large and Neotropical (Tyrannides) and 2 small and Paleotropical (Acanthisittides, New Zealand wrens, and Eurylaimides).

Infraorder EURYLAIMIDES Pittas Pittidae, broadbills Eurylaimidae, and asities Philepittidae (confined to Madagascar) (Olson 1971, Raikow 1982).

Superfamily EURYLAIMOIDEA: broadbills, aSlt1es Family EURYLAIMIDAE: broadbills Small to medium birds (weight range 20-160 g) with strikingly-coloured plumage. Bill moderately or extremely wide; outermost primary long, wing rounded; tail short or medium, more or less graduated. Tarsus with scutes anteriorly and small hexagonal scales posteriorly; toes 3 and 4 united basally. Main artery of leg is ischiatic. Non-furcate sterna spinalis. Tendon of flexor hallucis longus connected by vinculum to tendon of flexor digitorum longus. Inhabit humid forests; sedentary, somewhat sluggish; eat plant and some animal material. 1

Order

PASSERIFORMES 'Perching birds', mostly 'songbirds', much the largest order of birds, embracing approximately 5300 species worldwide, or well over half of all living birds; commonly called passerines for short. Cosmopolitan. Land-birds, only a very few species feeding in (fresh) water; mainly small, a few medium-sized - ravens Corvus spp. among the largest. Foot adapted for perching; always 4 toes, the hallux enlarged, directed backwards and not reversible; toes never webbed. In most species, syrinx adapted for singing; syrinx usually tracheo-bronchial with complex and varied musculature, but tracheal in 4 Neotropical families, and simple in a few groups. Wing eutaxic. Primaries 10, the outer 1 reduced or vestigial, or 9 in pipits, swallows, finches, buntings, some larks, and a few New World families . 12 tailfeathers. Aegithognathous palate, 'passerine' tensor propatagialis brevis, bundled spermatozoa with coiled head and large acrosome, and type VII deep plantar tendons. Young nidicolous, hatching blind, naked and helpless. Parental care well developed; but 3 groups brood-parasitic (2 in Africa). Passeriformes are a monophyletic group (Raikow 1982). Plumages, bill form and life-styles are very varied; 82 families recognized by Campbell and Lack (1985), many biologically distinctive. Evolutionary convergence rife, hence relationships, boundaries and true composition of families often questionable. Morphological and DNA-hybridization studies confirm that pittas, broadbills, tyrant-flycatchers, antbirds, ovenbirds, gnateaters and tapaculos form a natural assemblage (Suborder Tyranni - most families New World). They also suggest that 'true' songbirds of Suborder Passeri (Oscines) are related among themselves in ways radically different from those recognized conventionally (Sibley and Ahlquist 1990). In the main, classification of Passeriformes and sequence of passerine families adopted here follow those of Mayr and Greenway (1956) and Campbell and Lack (1985 ). However, subordinal nomenclature, and classification of Suborder Tyranni, follow Sibley et al. (1988 ).

Suborder TYRANNI (Suboscines) More or less synonymous with Deutero-oscines, Clamatores, Suboscines, Oligomyodi and Tyranniformes: 10- 14 families embracing c. 1100 species. Syrinx mesomyodian, with not more than 3 pairs of intrinsic muscles (Ames 1971); form of ear ossicle or stapes unique (except in New Zealand wrens: Feduccia 1980). Contains 3 infraorders - 1 large and Neotropical (Tyrannides) and 2 small and Paleotropical (Acanthisittides, New Zealand wrens, and Eurylaimides).

Infraorder EURYLAIMIDES Pittas Pittidae, broadbills Eurylaimidae, and asities Philepittidae (confined to Madagascar) (Olson 1971, Raikow 1982).

Superfamily EURYLAIMOIDEA: broadbills, aSlt1es Family EURYLAIMIDAE: broadbills Small to medium birds (weight range 20-160 g) with strikingly-coloured plumage. Bill moderately or extremely wide; outermost primary long, wing rounded; tail short or medium, more or less graduated. Tarsus with scutes anteriorly and small hexagonal scales posteriorly; toes 3 and 4 united basally. Main artery of leg is ischiatic. Non-furcate sterna spinalis. Tendon of flexor hallucis longus connected by vinculum to tendon of flexor digitorum longus. Inhabit humid forests; sedentary, somewhat sluggish; eat plant and some animal material. 1

2

EURYLAIMIDAE

W Africa to Philippines. 14 species in 8 genera; in Africa, 4 species in 2 endemic genera. The 2 African genera have generally been allocated to separate subfamilies, but 'recognition of subfamilies within the Eurylairnidae is discouraged' (Olson 1971).

Genus Pseudocalyptomena Rothschild Endemic, monotypic. Differs from Calyptomena (3 spp.; SE Asia) in lacking dense feather fan concealing base of bill and nostrils. Bill flat; rictal bristles < 5 mm. Tarsus slender and quite long, covered behind with soft skin with small rounded scales. First joint of toes 3 and 4 united. Outer primaries stiff and straight, with tip of inner web slightly twisted down. Carotids paired.

Plate 1 (Opp. p. 32)

Pseudocalyptomena graueri Rothschild. African Green Broadbill; Grauer's Broadbill. Eurylaime de Grauer. Pseudocalyptomena graueri Rothschild, 1909. Ibis, p. 690; west of Ruzizi R., north of L. Tanganyika, 2000 m.

Range and Status. Endemic resident. Main population in Itombwe Mts, ZaIre, where known at 5 localities (Ibachilo, Miki, Luvumba, Karungu, Muusi) between 3° 41'S, 28° 31 'E and 3° 4'S, 28° 48'E and from 1760 to 2480 m; common around Miki. 80% of c. 30 birds collected were at 1940- 2390 m. Known also from mountains west ofL. Kivu, ZaIre (Mt Kahuzi: Nyawaronga, 2° O'S, 28° 49 'E) and Impenetrable Forest, Uganda (Bwindi, Ruhizha, 2100- 2200 m). Threatened by forest clearance around villages; L. Kivu population probably safeguarded in Kahuzi-Biega Nat. Park; other 2 areas need protection (Collar and Stuart 1985). Description. ADULT cJ: forehead and crown greenish buff, thickly covered with small black streaks; nape, back, wings, rump and tail green; lores black, with very narrow black line over and behind eye; ear-coverts, chin, throat and upper breast pale blue; a few black or dusky spots form irregular moustachial streak; lower breast, belly and flanks green (paler and yellower than upperparts); undertail-coverts blue, or green with blue tips. Blue areas clearly demarcated from green ones. Rerniges blackish; underwing and undertail black. Bill black; eye very dark brown , eye-rim black; legs and feet greyish green to blackish. Impenetrable Forest population may be narrowerbilled than others (Friedmann and Williams 1968). Sexes alike. SIZE: (3 cJcJ, 2