The Birds of Africa [2] 0121373029, 9780121373023

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FRICA vlumell ...: , :_....: �;: ' �..... : ; ·--:>.·.:· ,_ : � -�- ; �- : ,_-·//.-: ;_:-�: . Edited by

EMIL K. URBAN Department of Biology, Augusta College, Augusta, Georgia, USA

C. HILARY FRY Department of Zoology, University of Aberdeen, Aberdeen, UK

STUART KEITH Department of Ornithology, American Museum of Natural History, New York, New York, USA

Colour Plates by Martin Woodcock Line Drawings by Ian Willis Acoustic References by Claude Chappuis

1986

@

ACADEMIC PRESS Harcourt Brace Jovanovich

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ACADEMIC PRESS INC. (LONDON) LTD. 24/28 Oval Road, London NWI 7DX

United States Edition published by ACADEMIC PRESS, INC. Orlando, Florida 32887

Copyright (C 1986 by ACADEMIC PRESS INC. (LONDON) LTD.

All rights reserved. No part of this book may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information retrieval system without permission in writing from the publisher

Birds of Africa Vol. II I. Birds-Africa I. Urban, Emil K. II. Fry, C. H. QL692.A I 598.296

III. Keith, Stuart

ISBN: 0-12-137302-9

Production services by Fisher Duncan Ltd, 10 Barley Mow Passage, London W4 4PH Printed in Hong Kong by Imago Publishing Ltd

ACIZNOWLEDGEMENTS We are most grateful to our authors for their untiring efforts and co-operation in preparing the species accounts, for providing us with unpublished notes and records, and for their patience during the protracted preparation of this volume for publication. We thank them all for their forbearance with editorial demands during a succession of drafts. We would like to acknowledge with thanks the contribution of referees, many of whom have gone to great lengths to ensure that the species accounts are as accurate and comprehensive as possible. Families have been refereed as follows: Phasianidae, R. K. Brooke, M.Ridley, the late B. W. H. Stronach and P. B. Taylor;Rallidae, P. B. Taylor; also A. Brosset, W. R. J. Dean, the late D. M. Skead and S. L. Olson; Gruidae, G. Archibald and P. Konrad; Heliornithidae, P. Ginn; Otididae, P. S. Jones, G. J. Morel, H. Schulz and W. R. Tarboton; Jacanidae W. R. Tarboton; Rostratulidae, M. de L. Brooke and D. J. Pearson; Dromadidae, C. J. Feare; Haematopodidae, J. Cooper; Recurvirostridae, D. J. Pearson and A. J. Tree; Burhinidae, J. M. Mendelsohn; Glareolidae (Egyptian Plover), J. R. Jehl, Jr (coursers), C. J. Vernon, (Glareolinae), C. Erard; Charadriidae (Charadriinae), G. L. Maclean, D. J. Pearson and A. J. Tree, (Vanellinae), C. J. Skead; Scolopacidae (Phalaropodinae), D. J. Pearson, (all others), A. J. Prater; Stercorariidae, J. C. Sinclair; Laridae, J. Cooper and P. J. Grant; Sternidae, P.R. Colston and C. J. Feare; Rynchopidae, M. J. Coe and D. J. Pearson; Alcidae, E. D. H. Johnson; Pteroclidae, D. H. Thomas; and Columbidae, A. Brosset, R. J. Dowsett, F. Dowsett-Lemaire, D. Goodwin andR. de Naurois. As well as writing species accounts or refereeing, G. Archibald, the late C. W. Benson, P, L. Britton, R. K. Brooke, A. Brosset, P. J. Grant, P. A. R. Hockey, P. Konrad, G. J. Morel, M.-Y. Morel, D. J. Pearson, A. J. Prater, P. B. Taylor, A. J. Tree and W.R. Tarboton have assisted in many additional ways and we are most thankful to them. We are greatly indebted to the Trustees and staff of: American Museum of Natural History, Department of Ornithology; British Library of Wildlife Sounds; British Museum (Natural History), Sub-Department of Ornithology; Carnegie Museum of Natural History; Cornell Laboratory for Ornithology; Durban Museum; Field Museum of Natural History, Bird Division; Los Angeles County Museum, Bird Division; MuseeRoyal de l'Afrique Centrale, 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; and Transvaal Museum, Bird Department, for repeatedly affording study facilities and for loans of numerous study skins. We especially thank P. J. K. Burton, P. A. Clancey, P. Colston, G.R. Cunningham-van Someren, the late B. G. Donnelly, J. Fitzpatrick, I. C. J. Galbraith, F. B. Gill, D. Goodwin, J. Gulledge, A. Harris, M. P. S. Irwin, H. D. Jackson, A. C. Kemp,R. Kettle, P. Lorber, M. Louette, J. Mendelsohn, J. P. Myers, S. L. Olson, K. C. Parkes, R. Paynter, D. Read, R. W. Schreiber, L. L. Short, D. W. Snow, W.R. Siegfried, D. Steadman, M. A. Traylor, M. Walters, G. E. Watson, and D. S. Wood. Further, we are indebted to G. Archibald and K.-L. Schuchmann for organizing meetings respectively at Bharatpur, India (International Crane Workshop) and Bonn, W. Germany (Symposium on African Vertebrates), and to the organizers of the Vth and Vlth Pan-African Ornithological Congresses at Lilongwe, Malawi, and Francistown, Botswana. We participated in all of these meetings, presenting papers stemming from study for The Birds of AJrica, and were afforded facilities to discuss the enterprise. In writing accounts of Palearctic birds in Africa we have drawn freely on the detailed information in Cramp and Simmons, The Birds of the Western Palearctic, and Glutz and Bauer, Handbuch der Vogel Mitteleuropas, and it is a pleasure to record our debt to the authors of these invaluable compilations. We are especially indebted to S. Cramp and M. Wilson for providing pre-publication information from Volume IV of the former work (now published). We have drawn heavily upon the generosity of correspondents, who have readily responded to queries affecting all aspects of African ornithology. G. C. Backhurst, G. R. Cunningham-van Someren, G. Nikolaus, A. J. Prater, M. D. Rae, T. Salinger, R. W. Schreiber, K.-L. Schuchmann and G. E. Watson provided us with thousands of bird measurements. We have benefitted greatly from the kindness, in providing us with their books and papers in manuscript prior to publication, of W.R. J. Dean and M. A. Huntley (Angola), P. Colston and K. Curry-Lindahl (Mt Nimba), M. K.Rowan (Columbidae), J. M. Thiollay (Ivory Coast) and F. Dowsett-Lemaire (Columbidae). S. M. Goodman, R. W. Storer and Sherif Baha El Din have provided us with manuscripts, records, food data and measurements from Egypt, andR. W. Schreiber and K. L. Garrett gave us complete data on African specimens in the Los Angeles County Museum. J. D.R. Vernon has provided distributional summaries for Morocco, M. Smart for Tunisia, M. W. Pienkowski for northwest Africa, P. W. P. Browne and B. Lamarche for Mauritania, G. Nikolaus and M. D. Rae for Sudan, and P. Becker for Namibia. M. de L. Brooke, J. Hinshaw, P. C. Lack, H. Lapman, E. Little (and colleagues at Augusta College's Reese Library and Aberdeen University's Queen Mother Library, Inter-Library Loan Departments), M. W. Pienkowski, A. J. Prater, C. Sibley, G. E. Watson, M. Wilson and staff of Oxford University Edward Grey Institute have been most helpful in obtaining library materials. We are no less indebted for help given in many ways by J. S. Ash, D.R. Aspinwall, P. C. Beaubrun, D. Baird, P. Becker, P. Bergier, V

VI

ACKNOWLEDGEMENTS

R. K. Brooke, A. Brosset, L. Bortoli, N. Bryant, B. Campbell, J. Carver, M.A. Casado, R. Chancellor, C. Chappuis, D.R. Collins, H. Q. P. Crick, A. A. Crowe, J.E. W. Dixon, R.J. Douthwaite, R.J. Dowsett, F. Dowsett-Lemaire, E. K. Dunn, J. H. Elgood, C. C. H. Elliott, H. F. I. Elliott, C. Erard, H. Fanshawe, G. D. Field, D. Fisher, R. W. Furness, J. P. Gee, P. Geroudet, P. J. Ginn, H. H. Gray, L. Grimes, J. Gulledge, S. I. Guttman, D. B. Hanmer, P. Hogg, D.J. Holmes, J. H. Hosken, S. Howe, R. Hudson, J. L. Ingold, M. P. S. Irwin, R. A. C.Jensen, M. C. Jennings, R.Johns, E. D. H.Johnson, M. Kelsey, P.J. Knight, B. Lamarche, C. Levassor, A. Lewis, Mrs. W. Licll-Cock, P. Lorber, A. Morris, R. de Naurois, J. H. van Niekerk, B. Ochando, R. Osborne, J. R. Peek, E. H. Pcnry, R. and J. Plunkett, D. Pomeroy, A. Prigogine, R. Ralph, J. F. Reynolds, M. Ridley, D. V. Rockingham-Gill, M. K. rowan, A. Sala, D. Schmid), J. T. R. Sharrock, W. R. Siegfried, C. J. Skead, G. S. Smith, t. Stevenson, P. Steyn, R. Stjernstedt, the late B. W. H. Stronach, C. D. Taylor, K. Thangavelu, M. Thevenot, D. H. Thomas, P. Thomsen, R. Thomsen, D. A. Turner, J. P. Vande weghe, L. Walkinshaw, A. J. S. Weaving, G. R. Welch, H.J. Welch and J. B. Wood. We should like to thank the following for permission to redraw black and white line illustrations from their own published or personal references: G. Archibald (Anthropoides paradisea, A. virgo, Balearica regulorum, B. pavonina, Bugeranus carunculatus and Grus grus); C. T. Astley Maberly in Bokmakierie (1967) (Eupodotis ruficrista), in Honeyguide (1979) (Neotis denhami); G. W. Begg (Vanellus a/biceps); G. R. Cunningham van Someren (Actophilornis africana); H. Deetjen in Journal fi.ir Ornithologie (1969) (Fulica cristata and F. atra); M. P. L. Fogden in Ibis (1964) (Larus leucopthalmus and L. cirrocephalus); P. G. H. Frost in the Bulletin of the British Ornithologists' Club (1975) (downy young of various Phasianidae); C. H. Fry (Podica senegalensis); D. Goodwin in 'Pigeons of the World', Cornell University Press ( 1983) ( Columba livia), in Avicultural Magazine (1956) ( Columba Livia, C. guinea, Streptopelia roseogrisea and S. semitorquata); C. J. 0. Harrison in Sandgrouse (1983) (Sterna albifrons and S. saundersi); P. A. R. Hockey (Hae111a10pus bachmani and Charadrius marginatus); J. B. D. Hopcraft (Actophilornis africana); J. Horsfall (Chlamydotis undulata); T. R. Howell in University of California Publications in Zoology (1979) (Pluvianus aegyptius); J. H. Van Niekerk in Bokmakierie (1983) (Francolinus swainsonii); G. L. Maclean ( Vanellus melanopterus, V. coronatus, Pterocles burchelli, P. namaqua, Burhinus oedicnemus, B. senegalensis, B. vermiculatus, B. capensis, Cursorius africanus) and in The Living Bird (1968) (Pterocles namaqua and P. bicinctus); K.A. Muller in Emu (1975) (Rostratula benghalensis); D. Pearson (Gallinago gallinago, G. nigripennis and G. stenura); G. P. Schaller (Ardeotis lwri); W. Tarboton (Actophilornis africana); J. Terres (Rynchops fl.avirostris); N. Tinbergen in Behaviour (1959) (Larus argentatus); N. Tinbergen & G. Broekhuysen in Ostrich (1925) (Larus hartlaubii); J. Trollope in the Avicultural Magazine (1970) (Turnix sylvatica); A. J. Weaving (Actophilornis africana); B. Wood in the Bulletin of the British Ornithologists' Club (1975) (Streptopelia hypopyrrha). We are especially pleased to have worked with Martin Woodcock who prepared the colour and black and white plates, Ian Willis who drew the black and white illustrations, Claude Chappuis who prepared the list of acoustic references, and Lois Urban who prepared the indexes. We wish to thank the staff of Academic Press (London) Ltd and Fisher Duncan Ltd, particularly Jane Duncan, Moira Fisher, Jennie Morley and Andrew Richford for helping us through an increasingly arduous task. With Volume III already in press, each Editor has accumulated vast files of correspondence and papers. We hope that we have acknowledged all of our numerous sources of help and apologize to-and ask the forbearance of­ anyone whose name has been inadvertently omitted. Finally, we would like to thank our three wives-Kathie Fry, Sallyann Keith and Lois Urban-for their patience, tolerance and understanding during the long hours we have spent preparing this volume.

July 1985

Emil K. Urban C. Hilary Fry Stuart Keith

CONTENTS ACKNOWLEDGEMENTS

V lX

LIST OF PLATES INTRODUCTION

Xl

Illustrations Superspecies Nomenclature and Systematics Range and Status Maps Description and Field Characters Voice General Habits and Breeding Habits .. Reference System References

Xl Xll Xlll Xlll

xv xv xv xv

XVl

ORDER GALLIFORMES Phasianidae, guineafowl, Congo Peacock, quail, partridges and francolins (T. M. Crowe, S. Keith and the late L. H. Brown) ORDER GRUIFORMES Turnicidae, button-quail (P. B. Taylor) Rallidae, rails, flufftails, crakes, gallinules, moorhens and coots (S. Keith; Crex egregia coauthored by P. B. Taylor) Gruidae, cranes (E. K. Urban) Heliornithidae, finfoots (C. H. Fry) Otididae, bustards (N. J. Collar, P. D. Goriup and P. E. Osborne) ORDER CHARADRIIFORMES Jacanidae, jacanas (C. H. Fry) Rostratulidae, painted-snipe (E. K. Urban) Dromadidae, Crab Plover (the late C. W. Benson and F. M. Benson) Haematopodidae, oystercatchers (P. A. R. Hockey) .. Recurvirostridae, stilts and avocets (E. K. Urban) .. Burhinidae, thick-knees, stone curlews and dikkops (G. L. Maclean and E. K. Urban) Glareolidae Cursoriinae .. Pluvianus, Egyptian Plover (T. R. Howell) Cursorius, coursers (G. L. Maclean and E. K. Urban) .. Glareolinae, pratincoles (A. Brosset) Charadriidae Charadriinae, plovers (E. K. Urban) Vanellinae, lapwings (S. Keith) Scolopacidae Calidridinae, Gallinagininae, Scolopacinae, Tringinae, Arenariinae, sandpipers and snipe (D. J. Pearson) .. Phalaropodinae, phalaropes (E. K. Urban) Stercorariidae, skuas (B. L. Furness and E. K. Urban) Laridae, gulls (P. L. Britton) .. Sternidae, terns (P. L. Britton) Rynchopidae, skimmers (C. H. Fry) Alcidae, auks (E. K. Urban) ORDER PTEROCLIFORMES Pteroclidae, sandgrouse (G. L. Maclean and C. H. Fry) vii

1 76 84 131 145 148 180 185 188 190 193 198 206 206 206 209 218 224 225 250 283 283 328 332 340 374 412 415 422

Vlll

CONTENTS

ORDER COLUMBIFORMES Columbidae, pigeons and doves (G.

J. Morel, M.-Y. Morel and C. H. Fry)

442

BIBLIOGRAPHY 1. General and Regional References 2. References for Each Family 3. Acoustic References Section A: Discs and Cassettes Section B: Most Important Discs and Cassettes by Region Section C: Institutions with Sound Libraries Section D: Individual Recordists

498 501 534 534 536 536 536

ERRATA, VOLUME I

538

INDEXES 1. Scientific Names 2. English Names 3. French Names

539 547 550

1

32

17

304

2

33

18

305

3

48

19

4

49

5

96

6

97

7

112

8

113

9

160 161 176

12 13

177 240

14

241

15

256

16

257

320

20

321

21

352

22

353

23

368

24

369

25

400

26

401

27

416

28

417

29

448

30

449

31

464

32

465

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INTRODUCTION Superspecies

The Birds of Africa was the brainchild of Leslie Brown, and his death necessitated a wholly new approach to authorship. It has become a multi-author work, under the joint editorship of one of the original authors, Emil K. Urban, co-equally with two members of the former Advisory Editorial Panel, C. Hilary Fry and Stuart Keith. Species accounts have been written by the best authority available for each family, and refereed by equally competent authorities, many, like the authors themselves, presently living and working in Africa. The numerous published reviews of Volume I have been in general laudatory. The Editors have benefitted from constructively critical comments of reviewers and correspondents; they have also been their own severest critics, and have sought to introduce such improve­ ments as could be made without materially altering the format established for the series with the first volume. With a large team of authors, the Editors have been obliged to ensure uniformity of treatment, which re­ quirement has inevitably resulted in a long and detailed set of instructions to authors. During the course of preparation of this volume, the instructions themselves have changed and evolved, so that some inconsistencies may have slipped through uncorrected. The main departures from the format of Volume I are in the layout of legends interpreting the colour plates, and in adop­ tion of the superspecies principle; these, and several less obvious innovations, are detailed below. Innovations notwithstanding, the treatment of species is in much the same format as in Volume I (in which see pp. 24-26: Content and Layout of the Text). The Birds of Africa was originally conceived as a four volume work. However, to accomodate the mass of information on African birds that is continually coming to light, both published and unpublished, the work has been expanded to six volumes, three non-passerine and three passerine. The present volume covers galliforms to columbiforms. The cut-off date for information included in this volume is November 1983, although many significant 1984 and 1985 references have been included.

The concept of the superspecies is invaluable in zoology to both museum systematists and field naturalists. The term was coined by Ernst Mayr ( 1931) as a translation of the Artenkreis of Rensch (l 929). Mayr later (1963) defined it as "a monophyletic group of entirely or essentially allopatric species that are too distinct to be included in a single species". The definition was further refined by Amadon (l 966) as "a group of entirely or essentially allopatric taxa that were once races of a single species but which now have achieved species status". The first works to apply the superspecies concept to a whole continent were Mayr and Short ( 1970) for North America and Hall and Moreau (1970) for Africa. The former had a more rigid definition of superspecies as "groups of taxa which have barely crossed the threshold of species status" (p. 100). The concept was more loosely used by Hall and Moreau, and some of their superspecies comprise species that clearly crossed this threshold some time ago and now exhibit very consider­ able differences. A brief explanation of the superspecies concept and its application in Africa and in this work may be fitting. In this huge continent numerous kinds of birds have vast ranges, and they vary geographically in plumage, size or behaviour. Some variation is gradual or clinal, and is a balance between differing selection pressures at each geographical extreme (producing differentiation) and unimpeded gene flow in between (promoting uniformity and countering differentiation). Commonly, variation is abrupt, particularly when different popula­ tions of a species are allopatric (isolated from each other by considerable distances). In many other instances isolates have expanded and met again, producing a spatially coherent species with abrupt changes at quite well marked boundaries between contiguous (para­ patric) subspecies. If the ranges of two essentially parapatric, closely related populations are beginning to overlap and they remain reproductively discrete despite frequent encounters, systematists must treat them as separate species, for that is how they treat each other. But well differentiated populations may hybridize where they meet, although the hybrid zone seems to be narrow, stable and contained (Prigogine 1980). The question then arises, should they be treated as the same or different species? The difficulty is even more acute when populations which are moderately well different­ iated, yet sufficiently alike to be certainly of immediate common descent, are either allopatric or strictly para­ patric. For practical purposes, such as the treatment which they will be accorded in this work, a decision has to be made as to whether they have differentiated to specific level; and the decision has to be based on an untestable prediction - how the two populations would behave reproductively were they ever to meet naturally.

Illustrations All of the Plates in this volume are by Martin Wood­ cock. As before, he has painted from skins, but he has also been able to familiarize himself with numerous birds in the field, by means of a three-month sketching visit to East and southern Africa. A new departure, intended to facilitate the identification of individual bird illustrations in each Plate, is that their names appear on full-sized outlines on the facing page. Black­ and-white illustrations in the text, mainly of behaviour, were drawn by Ian Willis from photographs and other original sources. xi

XII

INTRODUCTION

To that extent the recognition of species is arbitrary, and if the choice were solely between a subspecies or a full species the consequences for a work such as this would be profound. Had we decided, for instance, that Black and Grey Crowned Cranes were conspecific, Balearica pavonina sensu law would have received little more than half the text allocation given to what we in fact regard as two allopatric species, B. pavonina sensu stricto and B. regulorum. Such geographical replacement species, or allo­ species, of which there may be two or more, together constitute a superspecies. The use of this taxon not only emphasizes close affinity between congeneric species, but also reduces the practical difficulty stemming from limited taxonomic choice. Ideally, every sort of bird would receive textual treatment in direct proportion with its degree of distinction, brief if it is an ill-defined subspecies, and lengthy if it is a distinctive monotypic species without close relatives, like the Egyptian Plover Pluvianus aegypcius. For reasons of space, that ideal cannot be achieved here; but recognizing superspecies has permitted some textual abbreviation as a result of cross-reference between allospecies. It also allows the reader to extrapolate data about a well known bird to its poorly known allospecies with some confidence. Most birds considered to constitute a superspecies have their breeding ranges shown in the generic diag­ noses, using one map of Africa per superspecies, and their allospecies are also listed in the text, immediately after the species heading. Since their ranges do not overlap they are shown on the superspecies map simply in outline. In a few cases two birds may have the main attributes of a superspecies but their ranges are beginn­ ing to overlap (without hybridization); with them the area of overlap (e.g. the doves Strepiopelia vinacea and S. capicola) or of possible overlap (e.g. the bustards Ardeotis arabs and A. kori) are made plain. Where an African bird has a European or Asiatic allospecies which is a non-breeding visitor to Africa, the latter is identified as such in the text but its breeding range is not shown in the superspecies map. Allospecies never recorded in Africa are ignored (e.g. the Asiatic Little Pratincole Glareola lactea, an allospecies of the African Grey Pratincole G. cinerea).

Nomenclature and Systematics Authors have been required to assess critically the generic affiliations of a species, superspecific affiliations, specific boundaries, and the validity of described races. This has led to a few systematic innovations, although we have favoured conventional treatments and as a rule taxonomy follows that of Snow (1978) for species and White (1965) for subspecies. Family sequence is that of Voous (1973). Species sequence was decided by the Editors and authors, and is intended to express evolutionary progression from primitive to derived species wherever possible.

English Names Many African birds are known by different English names in different parts of the continent. The Editors, in selecting a single preferred name for each species, have had to make some difficult decisions. (No more than a single alternative name is provided for any bird, and that only when it is widely used.) For a full account of the principles used in selecting names, see S. Keith and L. Short, 'The English names of African birds' (Proc. VI Pan-Afr. Orn. Congr., in press). The main points are as follows: 1. We take a world view of English names; it is our intention that every African bird shall have a name that clearly distinguishes it from every other bird in the world. In cases of synonymy we either change the name of our bird or add 'African' to it. For instance, the name Purple Gallinule has been given to both Porphyrio martinica of the New World and P. porphyrio of the Old World; for the latter species we decided to use another available name, Purple Swamphen. 'White-winged Dove' is used for both Zenaida asiatica of the New World and Streptopelia reichenowi of Africa; we dis­ tinguish our bird by calling it African White-winged Dove. In another move toward global conformity, all members of the genus Vanellus are here called Lap­ wings, following Johnsgard (1981). For African species which go by different names in other parts of the world, we use the name most appropriate to its world range; if the names seem equally valid, we use the African name; e.g. for Fulica cristata we prefer Red-knobbed Coot (African) to Crested Coot (European). In the case of Palearctic species we use the European name, e.g. Kentish Plover for Charadrius alexandrinus, rather than Snowy Plover (North America) or Red-capped Dotterel (Australia). 2. Certain species which are locally the sole repre­ sentatives of their genus or family-particularly in Britain-have arrogated the genus or family name to themselves, without a modifier. To these we give modifiers, e.g. Coot ('The' Coot) becomes Eurasian Coot, 'The' Knot becomes Red Knot, 'The' Turnstone becomes Ruddy Turnstone. 3. We tend to be conservative and to favour old, traditional names even if not completely accurate or appropriate. Usage has an important role, and we coin new names only for the most difficult cases, and then only after searching for already-published alternatives. 4. For some purely African species we replace restrictive regional names with descriptive ones appro­ priate for the species as a whole. 'Uganda Spotted Woodpecker' is inappropriate for a bird that occurs from Nigeria to Kenya, and we adopt the name coined for it by Short (1982), Speckle-breasted Woodpecker. But we favour geographical names when they describe all or nearly all of a bird's range, as Djibouti Francolin for Francolinus ochropectus. 5. Changes in taxonomy sometimes dictate changes in English names. Upon systematic reappraisal of fran­ colins, Francolinus psilolaemus emerged as a full species; its component races had previously been assigned to other species and consequently no English name was

Dense breeding Sparse breeding []]]Il]] Migration Dense non-breeding Sparse non-breeding 7 Occurrence uncertain X Extralimital record / Points to isolated / occurrence Migration route N B This tone � shows where breeding and non-breeding ranges overlap

I

K ·_1

E

N Y A

XVI

INTRODUCTION

References Amadon, D. (1966).The superspecies concept. Sysl. Zool. 15, 245-249. Brown, L. H. and Britton, P. L. (1980). 'The Breeding Seasons of East African Birds'. East African Natural History Society, Nairobi. Chappuis, C. (1980). List of sound-recorded Ethiopian birds. Malimbus 2, 1-15, 82-98. Hall, B. P. and Moreau, R. E. (1970). 'An Atlas of Speciation in African Passerine Birds'. British Museum (Natural History), London. Johnsgard, P.A. (1981). 'The Plovers, Sandpipers and Snipes of the World'. University of Nebraska Press, Lincoln, Nebraska. Mayr, E. ( 1931 ). Birds collected during the Whitney South Sea Expedition. XII. Notes on Halcyon chloris and some of its subspecies. Amer. Mus. Novil. 469, 1-10.

Mayr, E. (1963). 'Animal Species and Evolution'. Belknap Press (Harvard University), Cambridge, Mass. Mayr, E. and Short, L. L. (1970). Species taxa of North American birds.A contribution to comparative systemat­ ics. Publ. Nuuall Orn. Club No. 9; Cambridge, Mass. Prigogine, A. (1980). Etude de quelques contacts secondaires au Zai:re oriental. Gerfaut 70, 305-384. Rensch, B. (1929). 'Das Prinzip Geographischer Rassenkreise und das Problem der Artbildung'. Borntraeger Verlag, Berlin. Short, L. L. (1982). 'Woodpeckers of the World'. Delaware Museum of Natural History, Greenville, Delaware. Snow, D. W. (ed) (1978). 'An Atlas of Speciation in African Non-passerine Birds'. British Museum (Natural His­ tory), London. Voous, K. H. (1973). List of recent Holarctic bird species. Non-passerines. Ibis 115, 612-638. White, C. M. N. (1965). 'A Revised Check List of African Non-passerine Birds'. Government Printer, Lusaka.

Order GALLIFORMES A large cosmopolitan order of small to very large, mainly terrestrial birds. Earliest fossils date from mid-Eocene, i.e. c. 45 million years ago. The only taxonomic character which unambiguously indicates that the order is monophyletic is presence of a lateral foramen delimited by fused manubrial spines of sternum. There are 2 superfamilies: Megapodioidea (Australasian megapodes) and Phasianoidea (South American Cracidae, and the bulk of the world's galliforms, the Phasianidae). Nearest relatives appear to be the Anseriformes, perhaps via the screamers (Anhimidae) (Prager and Wilson 1981, Cracraft 1981, Dzerzhinsky 1982, C. G. Sibley, pers. comm.); but see Olson and Feduccia (1980) for another view. Only one family in Africa, the Phasianidae. This family may be divided into 6 subfamilies (T. M. Crowe et al. unpub., Mainardi 1963, Stock and Bunch 1982): Numidinae (guineafowl), Pavoninae (peafowl, Argus and peacock pheasants, Congo Peacock), Gallinae (junglefowl, Old World quail, francolins and partridges), Tetraoninae (grouse), Meleagridinae (turkeys), and Phasianinae (true pheasants and New World quail). Of these, the Numidinae, Pavoninae and Gallinae occur in Africa.

Family PHASIANIDAE: guineafowl, Congo Peacock, quail, partridges and francolins A large and diverse family, including the smallest (quail), largest (turkeys), and most spectacular (peafowl) galliforms. 4 species, Junglefowl (Gallus gal/us), Turkey (Meleagris gallopavo), Helmeted Guineafowl (Numida meleagris) and Common Quail (Coturnix coturnix) are important domesticated animals. Many phasianids arc popular gamebirds and/or are prized by aviculturalists. Represented in Africa by 10 genera. Phasianids inhabit every African vegetation type from desert to dense forest, and from sea level to the upper limits of vegetation on high mountains. All are terrestrial, with rather short, stout, sometimes strongly spurred, tarsi. J J average larger than 9'i?. Heads usually small; bills short and robust, well suited for digging; necks short; bodies stout, rounded, with very large breast muscles; wings short, rounded, with little emargination of primaries; tails short, but long in Congo Peacock and Vulturine Guineafowl. Sedentary except for quail, which are migratory or at least nomadic. Prefer to escape by running; fly only reluctantly and when pressed, or to reach elevated roosts; flight direct, fast, with rapid wing-beats, usually low and not long sustained. Feed on a variety of animal and vegetable matter found on or immediately above or below ground level (although Crested and Vulturine Guineafowl Guttera and Acryllium spp. have been observed eating fruit in trees). All have well developed crops, powerful gizzards and extensive caeca. Syrinx relatively simple; voices loud, often raucous, but calls not elaborate. Most lay large clutches on the ground, and all produce precocial young, capable of limited flight within 2 weeks of hatching.

Subfamily NUMIDINAE: guineafowl A well defined endemic African subfamily comprising 4 genera: Agelastes (2 spp.), Guttera (2), Acryllium (I) and Numida (1). Morphological (Cracraft 1981, T. M. Crowe et al., in prep), immunological (Mainardi 1963) and karyological (Stock and Bunch 1982) evidence suggests that guineafowl link Phasianidae and Cracidae. Large galliforms with bare heads and necks, often brightly coloured; most with feathers or a casque on crown. Heads relatively small in comparison with body. Bills short, stout, upper mandible arched. Necks relatively long, bodies stout, thickset. Wings short, rounded. Tails short (long in Acryllium). Tarsi short, stout, unspurred in most species, well adapted for running, walking and scratching. 3 forward and 1 strong hind toe. Almost exclusively ground foragers, using feet and bill to excavate food items. Plumage black, spotted and/or vermiculated with white. No obvious sexual dimorphism, although cS average larger than 9 , and cScS of at least I species (Helmeted Guineafowl Numida meleagris) possess a cloaca! protuberance in the breeding season. Voices raucous, often high-pitched; more musical and whistling in Agelastes spp. All appear sedentary. Inhabit nearly every type of African vegetation from dense tropical forest to subdesert steppe. The Helmeted Guincafowl has been widely domesticated and feral populations have become established in some areas, e.g. Cuba and W Cape Province (South Africa). Generally gregarious when not breeding; flocks fragment into monogamous pairs at onset of breeding season.

Genus Agelastes Bonaparte Small guineafowl inhabiting dense primary forest. Head and neck bare, skin dull to bright pink; no tufts or plumes on crown. Body plumage black, or black with white collar, with varying amount of white vermiculation. 2 monotypic

2

PHASIANIDAE spp., both very little known. JJ and some 99 have 1 or 2 tarsal spurs. Spurs are bony outgrowths from tarsometatarsus itself (as in Gallus), and not from the hypotarsus as in most phasianids. Tarsal scales imbricated and in rows, as in francolins, and unlike other guineafowl genera. The most primitive members of the subfamily, nearest to an ancestral francolin-like stock (Ghigi 1936, Crowe 1978a), or perhaps from the same stock which gave rise to the junglefowl.

Plate 5 (Opp. p. 96)

Agelastes meleagrides Bonaparte. White-breasted Guineafowl. Pintade blanche.

a poitrine

Agelastes meleagrides 'Temm.' Bonaparte, 1849 (1850). Proc. Zool. Soc. London, p. 145; no locality given = Ghana. Forms a superspecies with A. niger. Range and Status. Resident in primary forests of W Africa from Liberia to Ghana. Formerly occurred Sierra Leone and probably S Guinea. Not seen Ghana since 1963, may be extinct there. Very rare Liberia; formerly occurred Mt Nimba but rapidly became extinct when area opened up for iron ore mining. Status in Ivory Coast unclear; still some numbers at Tai National Park, but generally uncommon to rare. Severely threatened by hunting pressure and habitat destruction and will probably disappear except from a few protected areas. Possibly one of the most endan­ gered birds in Africa.

Agelosfes meleogndes

Description. ADULT J: head and upper neck naked and red, pinkish on lower neck with scanty whitish filaments. A broad white collar on hindneck and breast. Rest of body plumage black, finely vermiculated white. Primaries dark brown, edged grey on outer webs; secondaries dull brown, vermiculated white on outer webs. Bill greenish brown; eye brown; legs greyish brown or greyish black; 1-2 spurs 5-8·5 mm. Sexes alike, JJ average larger. SIZE: (7 JJ, 4