The Birds of Africa [3, 1 ed.] 0121373037, 9780121373030

Table of contents :
Cover
CONTENTS
ACKNOWLEDGEMENTS
LIST OF PLATES
INTRODUCTION
ORDER PSITTACIFORMES
Psittacidae, parrots, parakeets and lovebirds (the late C. W. Benson, F. M. Benson, S. N. Stuart and C. H. Fry)
ORDER MUSOPHAGIFORMES
Musophagidae, turacos, go-away birds and plantain-eaters (A. Brosset and C. H. Fry)
ORDER CUCULIFORMES
Cuculidae, cuckoos, malkohas and coucals (M. P. S. Irwin)
ORDER STRIGIFORMES
Tytonidae, barn owls and bay owls (R. T. Wilson, M. P. Wilson and C. H. Fry)
Strigidae, typical owls (A. C. Kemp)
ORDER CAPRIMULGIFORMES
Caprimulgidae, nightjars (C. H . Fry and R. M. Harwin)
ORDER APODIFORMES
Apodidae, spinetails and swifts (C.
H. Fry)
ORDER COLIIFORMES
Coliidae, mousebirds or colies (J. P.
Decoux)
ORDER CORACIIFORMES
Trogonidae, trogons (G. R. Cunningham-van Someren and C. H. Fry)
Alcedinidae, kingfishers (C.
H. Fry)
Meropidae, bee-eaters (C.
H. Fry)
Coraciidae, rollers (C.
H. Fry)
Phoeniculidae, wood-hoopoes (D. Ligon and N . C. Davidson)
Upupidae, Hoopoe (C.
H. Fry)
Bucerotidae, hornbills (A. C. Kemp)
ORDER PICIFORMES
Capitonidae, barbets and tinkerbirds (L. L. Short and J. F.
M. Horne)
Indicatoridae, honeyguides (L. L. Short and J. F. M. Horne)
Picidae, woodpeckers, piculets and wrynecks (L. L. Short)
BIBLIOGRAPHY
1. General and Regional References
2. References for Each Family
3. Acoustic References
ERRATA, VOLUME III
INDEXES
1. Scientific Names
2. English Names
3. French Names

Citation preview

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Edited by

C. HILARY FRY

Department of Biology, Sultan Qaboos University, Muscat, Sultanate of Oman

STUART KEITH

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

EMIL K. URBAN

Department of Biology, Augusta College, Augusta, Georgia, USA

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

ACADEMIC PRESS Harcourt Brace & Company, Publishers

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ACADEMIC PRESS LIMITED 24/28 Oval Road, London NWl 7DX

United States Edition published by ACADEMIC PRESS, INC. San Diego, CA 92101

Copyright© 1988 by ACADEMIC PRESS LIMITED Second printing 1993

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. III I. Birds-Africa I. Fry, C. Hilary. II. Keith, S. QL692.Al 598.296

III. Urban, E. K.

ISBN: 0-12-137303-7

Colour plates printed in Great Britain by George Over Ltd, Rugby Printed and bound in Great Britain by Mackays of Chatham PLC, Chatham, Kent

ACKNOWLEDGEMENTS We wish to express, first, 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 then 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. C. W. Benson, among the greatest ofAfrican ornithologists, died whilst researching and writing the Psittacidae, and we are particularly grateful to his widow, F. M. Benson, and to S. N. Stuart, for completing his work. We are much indebted to referees, whose input has been meticulous and invaluable. Referees for this volume have been: Psittacidae, G.A. Smith;Musophagidae,M.Candy, C. Erard, G. J.Morel, D.K.Read and the late M. K. Rowan; Cuculidae, J. F. R. Colebrook-Robjent, C. J. Vernon and S. N. Stuart; Tytcinidae, A. C. Kemp; Strigidae, N.Collar, P. Colston, B. King, T. J. Roberts and P. Steyn; Caprimulgidae, the late C. W.Benson, J. F. R. Colebrook-Robjent and D. H. Jackson; Apodidae, J. S.Ash, R. K. Brooke, A. Brosset, R. M. Harwin, R. de Naurois, A. Prigogine, R. K. Schmidt, W. Serie, P. Steyn and E. K. Urban (we are especially grateful to R. K. Brooke, who commented painstakingly on all 22 species); Coliidae, H. Schifter; Trogonidae, A. Brosset; Alcedinidae, G.Arkell, D. Boag,A. Brosset, P. J. K. Burton, P.A. Clancey, R.J.Douthwaite, P.W.Greig-Smith, D. B.Hanmer, P.J. Jones, B.S.Meadows, R.de Naurois, H. U.Reyer and A.Tye; Meropidae, J.S. Ash, P. L. Britton, H. Q.P.Crick, G. R.Cunningham-van Someren,M.Dyer, D.B. Hanmer, R.M.Harwin, R.E.Hegner, J. Krebs and T. J. Roberts; Coraciidae, J. S. Ash,M. P. S. Irwin, J.M. Thiollay and A.Tye; Phoeniculidae, C.H. Fry and S. Keith; Upupidae, D.M. Skead; Bucerotidae, C. H.Fry, J. Kalina andM. I. Kemp; Capitonidae, G.R. Cunningham-van Someren, also D. R.Aspinwall and H. Winkler; Indicatoridae, H. Friedmann, also G. R. Cunningham-van Someren, A. W. Diamond and H.Adan !sack; Picidae, G. R. Cunningham-van Someren, F. Dowsett-Lemaire, T. Oatley and H. Winkler. 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; British Museum (Natural History), Sub-Department of Ornithology; Carnegie Museum of Natural History; DurbanMuseum; FieldMuseum of Natural History, Bird Division; Los Angeles County Museum, Bird Division; Merseyside County Museum, Liverpool; Musee Royal de l'Afrique Centrale, Tervuren; Museum of Comparative Zoology, Harvard; Museum Alexander Koenig, Bonn; National Museum of Zimbabwe, Bulawayo; NationalMuseums 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 TransvaalMuseum, 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, the Cornell University Laboratory of Ornithology, and the Transvaal Museum Bird Department.We owe particular thanks to the following staff members of these institutions: P. J. K. Burton, P. A. Clancey, P. Colston, G. Cowles, G. R. Cunningham-van Someren, }."Fitzpatrick, I.C.J. Galbraith, F.B.Gill, J. Gulledge, A. Harris,M. P. S. Irwin, H. D. Jackson, A. C. Kemp, R. Kettle, A. Knox, P.Lorber, M.Louette, J. Mendelsohn, P. Morgan, J. P. Myers, S. L. Olson, K. C. Parkes, R. Paynter, D. Read, H. Schifter, R. W. Schreiber, K.-L. Schuchmann, L. L. Short, W. R. Siegfried, D. Steadman, M. A. Traylor, M. Walters, G. E. Watson and D. S. Wood. During the preparation of species accounts, we have drawn heavily upon the special knowledge of numerous colleagues, who have given us access to unpublished and pre-published works, provided measurements,. weights, information on distribution and food, and library materials, and have responded to a host of queries. For their help we have pleasure in thanking: J. Anderson, J. S.Ash, D.R. Aspinwall, G. C. Backhurst, S. Baha el Din, D.Baird, P.C. Beaubrun, P. Becker, the late C. W. Benson, P. Bergier, L.Bortoli, P.St.J.Bowen, J.Bradbury, R. Brereton, S.Brereton,M.de L. Brooke, R. K. Brooke, D. Brooks,A. Brosset, P.W.P.Browne, M.D.Bruce, N.Bryant, P.J. K. Burton, M. Byrne, B. Campbell, M. Candy, J. Carver, M. A. Casado, R. Chancellor, K.A. Chapman, C. Chappuis, J. F.R. Colebrook-Robjent, N. Collar, D.R. Collins, P.R. Colston, J. Cracraft, the late S. Cramp, H. Q.P.Crick,A.A. Crowe, G. R. Cunningham-van Someren, W. R. J.Dean,M.J. Delany, I.Dillingham, 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. Eriksen, H.Fanshawe, C. J. Feare, G. D. Field, D. Fisher,M. Fisher, R.W.Furness, K. L. Garrett, J. P. Gee, P. Geroudet, P. J. Ginn, S.M. Goodman, D. Gordon, H. H. Gray,A.A.Green, L. Grimes, J. Gulledge, S. I. Guttman, D. B. Hanmer,A. Harper,A. Harris, J. Harwin, W. J. Haswell, C. J.Hazevoet, S. Head, J. Hinshaw, P. Hogg, D. J.Holmes, J.H.Hosken, S.Howe, R.Hudson,M.A.Huntley, L. Hyde, J.L. Ingold, M. P.S. Irwin, H.D. Jackson,M. C. Jennings, R.A. C. Jensen, R. Johns, 3..D.H.Johnson, K.A.Joysey, J.Kalina, M. Kelsey,A. C. Kemp,M. I. Kemp, B. King, P. J. Knight, P. C. Lack, B. Lamarche, V. Lambert, H. Lapham, C. Levassor, A. Lewis, W. Liell-Cock, E. Little, G. Lodge, T. Lodge, P. Lorber, M. Louette, G. L.Maclean, C. Marais, G. J. Morel, A. Morris, R. de Naurois, J. H. van Niekerk, G. Nikolaus, T. B. Oatley, B. Ochando, R. Osborne, P. Pattie, R.B. Payne, J. R. Peek, E. H.Penry,M. W. Pienkowski, J.Plunkett, R. Plunkett, D.Pomeroy, V

VI

ACKNOWLEDGEMENTS

A. J.Prater, A.Prigogine,M. D.Rae, R. Ralph,M. I.Rawahy, J. F. Reynolds,M. Ridley, A. A.Ridley, D.Robel, T. J. Roberts, D. V. Rockingham-Gill, the lateM. K. Rowan, R. G. Ryan, A. Sala, D. Schmidl, R. W. Schreiber, K.-L. Schuchmann, W. Serie, J. T.R. Sharrock, C. Sibley, W.R. Siegfried, C. J. Skead, N. J. Skinner,M. Smart, G. S. Smith, D. W. Snow, W. Stanford, T. Stevenson,P. Steyn, R. Stjernstedt, the late B. W. H. Stronach, S. N. Stuart, C. D. Taylor, R. B. Taylor, K. Thangavelu, M. Thevenot, J.M. Thiollay, D. H. Thomas,P. Thomsen, R. Thomsen, D. A. Turner, A. Tye, J.P. Vande weghe, C. A. Vaucher, C. Vernon, J. D.R. Vernon, C. Violani, L. Walkinshaw,M. Walters, G. E. Watson, A. J. S. Weaving, G. R. Welch, H.J. Welch, R. Wilkinson,M. Wilson and J. B. Wood. We acknowledge with thanks the following sources of material for the preparation of black and white line illustrations: F. G. H. Allen in Birds of the World (1970) (Cypsiurus parvus, Apus caffer); F. V. Blackburn in British Birds (1975) (Caprimulgus europaeus); P. Blasdale (Caprimulgus tristigma); R. Bloomfield in Birds of the World (1970) (Lybius torquatus); D. Boag in 'The Kingfisher' (1982) (Alcedo atthis); D. Bromhall in 'Devil Birds' (1980) (Apus apus); A. Brosset and R. Darchen in Biologia Gabonica (1967) (Halcyon badia); J. Carr in Birds of the World (1970) (Bucorvus abyssinicus); R. W. Carroll (Caprimulgus binotatus); J.P. Chapin in 'Birds of the Belgian Congo' (1954) (Neafrapus cassini, Cypsiurus parvus and Tachymarptis aequatorialis); A. Christiansen in Birds of the World (1970) (Psittacus erithacus); D. M. Cottridge in British Birds (1981) (Apus pallidus); H. Q.P. Crick (Merops bullocki); J.P. Decoux (Coliidae); N. J. Demong (Merops bullockoides); A. Devez in 'Vie de Ja Foret Equatoriale' ( I 976) (Scolopelia bouvieri, Apaloderma aequatoriale, Campethera caroli and C. nivosa); M. Dizerens in Tierwelt (1982) ( Tachymarptis melba); R.J. Douthwaite in Journal of the East African Natural History Society (1978) (Ceryle rudis); J. Echevarri andM. Olano in Natural History (1981) (Merops apiaster); F. Erize in Birds of the World (1970) (Lybius leucomelas);M. Fogden in Birds of the World (1970) (Gymnobucco bonapartei); R. Fothergill in Honeyguide (1984) (Agapornis lilianae); 0. von Frisch in Bonner Zoologische Beitriige (1966) (Coracias garrulus); C. H. Fry (Cenlropus senegalensis, Merops oreobates, M. breweri, M. albicollis, M. malimbicus), in Journal of Zoology (1969) (Macrodipteryx longipennis), in Ibis (1980) (Halcyon leucocephala and H. chelicuti) and in Ostrich (1982) (Merops nubicus); M. Gartshore (Merops breweri"); U. N. Glutz von Blotzheim and K. M. Bauer in 'Handbuch der Vogel Mitteleuropas' (1980) (Apus apus, Cuculus canorus and Jynx torquilla); M. Goetz (Bubo africanus, Otus leucotis);P. W. Greig-Smith in Ostrich (1978) (Halcyon senegalensis); M. Herremans and J. Stevens in Malimbus (1983) (Caprimulgus climacurus); F. Hoffer (Apaloderma narina); E. Hosking in Birds of the World (1970) (Trachyphonus darnaudii); R. A. C.Jensen in 'Birdlife of Southern Africa' (1979) (Clamator glandarius);P. Johnson in Birds of the World (1970) (Phoeniculus purpureus, Lybius leucomelas); J. Karmali in 'Birds of Africa' (1980) (Bucorvus cafer); A. Kemp (Strix woodfordi, Bubo lacteus, B. africanus, Otus scops, 0. senegalensis); J. D. Ligon in The Living Bird (1978) (Phoeniculus purpureus); H. Lohrl in Der Ornithologische Beobachter (1978) (Jynx torquilla); J.Markham in British Birds (1961) (Caprimulgus europaeus); H.Mester and W. Priinte in Ornithologische Mitteilungen (1957) (Apus apus); N. Myburgh in 'Birdlife of Southern Africa' (1979) (Geocolaptes olivaceus); R. de Naurois in Alauda (1985) (Zoonavena thomensis); G. Newlands (Bubo africanus); K. Newman (Caprimulgus fossii), in 'Birdlife of Southern Africa' (1979) (Cuculus solitarius, Dendropicos fuscescens) and in Birds of the World (1970) (Trachyphonus levaillantii); K. Newman and P. Steyn in Bokmakierie (1970) (Cuculus solitarius); L. Norstrom in Birds of the World (1970) (Apus affinis); C. E. Palmer in British Birds (1981) (Apus apus); M.Pforr and A. Limbrunner in 'Breeding Birds of Europe', Vol. 2 (1982) (Coracias garrulus) and in 'Bird Life' (1979) (Upupa epops); J.Pick in British Birds (1981) (Apus pallidus); D. C. H.Plowes in Birds of the World (1970) (Cypsiurus parvus and Phoeniculus cyanomelas); R.Reucassel in Bokmakierie (1970) ( Scotopelia peli); H. U.Reyer in Ostrich (1980) (Halcyon chelicuti); J. F. Reynolds (Caprimulgus pectoralis), and in British Birds (1975) (Coracias garrulus); H. M. Salmon in British Birds (1969) (Apus caffer); H. Schouten in British Birds (1981) (Apus apus); L. L. Short in American Museum Novitates (1985) (Indicator variegatus, Trachyphonus erythrocephalus, T. darnaudii); G. A. Smith in Avicultural Magazine (1972) (Psittacula krameri) and in Ibis (1975) (Poicephalus spp. and Psittacula krameri); V. G. L. van Someren in 'Days with Birds' (1956) (Centropus superciliosus); J. Stannard in Bokmakierie (1971) (Tauraco persa); P. Steyn (Caprimulgusfossii and Centropus superciliosus), in Ostrich Supplement (1971) (Caprimulgus tristigma), in Bokmakierie (1973 and 1976) (Upupa epops), and in 'Delight of Owls' (1984) (Tyto alba, T. capensis, Bubo africanus, B. capensis, B. lacteus, Scotopelia peli, Glaucidium perlatum and Asio capensis); J. F. andM. Terrasse in Birds of the World (1970) ( Tachymarptis melba); W. Tarboton in Bokmakierie (1970) (Dendropicos namaquus); J. M. Thiollay in Annales d'Universite d'Abidjan, series E, Ecologie (1971) (Halcyon malimbicus); R. Thompson in British Birds (1961) (Caprimulgus europaeus); K. Voous in 'Atlas of European Birds' (1960) (Tachymarptis melba); D. Waters (Psittacula krameri); A. J. S. Weaving (Macrodipteryx vexillaria) and in Bokmakierie (1973) (Upupa epops); and M. C. Wilkes in British Birds (1981) (Apus apus). All colour plates in this volume are the work of Martin Woodcock, and all black and white line illustrations except those given below were drawn by Ian Willis. 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 C. H. Fry: Caprimulgus binotatus (A, B), C. europaeus (B), C. pectoralis (A), Macrodipteryx longipennis (B), Neafrapus cassini (A), Cypsiurus parvus (A), Apus apus (B), A. pallidus (A), A. caffer (A- right), A. affinis (A), Tachymarptis aequatorialis (A), T. melba (A), Colius striatus (C), C. colius (A), Apaloderma narina (A), Merops nubicus (A) and Upupa epops (A-D). Martin Woodcock has spent many months on field sketching safaris in Africa, expressly to improve his

ACKNOWLEDGEMENTS

portraiture of birds in this work. He is grateful to all those friends and correspondents who have kindly offered criticism, help and information and especially photographs, in particular: the Editors and authors and, in Africa, N. E. Baker, Block Hotels, E. and A. Boswell, J. and A. J. Beakbane, W. and N. Cooper, M.A. C. and E. Coverdale, G. R. Cunningham-van Someren, K. B. Newman, D. J. Pearson, M. Ross, T. Stevenson and D. A. Turner. He is especially indebted to the Trustees and staff of the British Museum (Natural History) Sub-department of Ornithology at Tring for the loan of specimens and for their help. Again it is our pleasure to thank C. Chappuis for preparing the acoustic references, Lois Urban for indexing the book, our wives Kathie Fry, Sallyann Keith and Lois Urban for their support and understanding, and lastly Andrew Richford of Academic Press (London) Ltd, himself an experienced ornithologist, who has done more to stitch the entire work together than its readers can ever know. September 1987

C. Hilary Fry Stuart Keith Emil K. Urban

VII

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CONTENTS ACKNOWLEDGEMENTS ..

V

LIST OF PLATES

Xl

INTRODUCTION

X111

ORDER PSITTACIFORMES Psittacidae, parrots, parakeets and lovebirds (the late C.W.Benson, F.M.Benson, S.N. Stuart and C.H.Fry)

1

ORDER MUSOPHAGIFORMES Musophagidae, turacos, go-away birds and plantain-eaters (A.Brosset and C.H.Fry)

26

ORDER CUCULIFORMES Cuculidae, cuckoos, malkohas and coucals (M.P.S.Irwin) ..

58

ORDER STRIGIFORMES Tytonidae, barn owls and bay owls (R.T.Wilson, M.P.Wilson and C.H. Fry) Strigidae, typical owls (A.C.Kemp) ..

105 110

ORDER CAPRIMULGIFORMES Caprimulgidae, nightjars (C.H.Fry and R.M.Harwin)

155

ORDER APODIFORMES Apodidae, spinetails and swifts (C.H.Fry)

197

ORDER COLIIFORMES Coliidae, mousebirds or colies (J.P.Decoux) ..

243

ORDER CORACIIFORMES Trogonidae, trogons (G.R.Cunningham-van Someren and C.H.Fry) Alcedinidae, kingfishers (C.H.Fry) Meropidae, bee-eaters (C.H.Fry) Coraciidae, rollers (C.H.Fry) .. Phoeniculidae, wood-hoopoes (D.Ligon and N.C.Davidson) Upupidae, Hoopoe (C.H.Fry) .. Bucerotidae, hornbills (A.C.Kemp) ..

255 263 302 338 356 371 375

ORDER PICIFORMES Capitonidae, barbets and tinkerbirds (L.L.Short and J. F.M.Horne) Indicatoridae, honeyguides (L.L.Short and J. F.M.Horne) Picidae, woodpeckers, piculets and wrynecks (L.L.Short) ..

413 486 512

BIBLIOGRAPHY 1. General and Regional References 2. References for Each Family . 3. Acoustic References ..

557 561 587

ERRATA, VOLUME III

591

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

593 605 609

ix

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1

Parrots..

32

17

Kingfishers

304

2

Parrots..

33

18

Bee-eaters

305

3

Turacos

48

19

Bee-eaters

320

4

Turacos

49

20

Rollers ..

321

5

Cuckoos

96

21

Wood-hoopoes and Hoopoe ..

352

6

Cuckoos

97

22

Hombills

353

7

Owls

112

23

Hombills

368

8

Owls

113

24

Hombills

369

9

Owls

160

25

Barbets..

400

Nightjars

161

26

Barbets..

401

Nightjars

176

27

Barbets..

416

12

Swifts

177

28

Barbets and wrynecks

417

13

Swifts

240

29

Honeyguides and honeybirds .

448

14

Colies

241

30

Piculets and woodpeckers

449

15

Trogons and kingfishers

256

31

Woodpeckers

464

16

Kingfishers

257

32

Woodpeckers

465

10

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INTRODUCTION This volume covers psittaciforms to piciforms, and completes the non-passerines. The literature has been thoroughly reviewed up to August 1986; for a great many species it has been possible to add material from sources as recent as May 1987. Style and layout are the same as in Volume II. Leslie Brown's death, and the consequent restructuring of editorship and authorship in that Volume, meant that the original 'Brown, Urban and Newman' credit se­ quence could not be maintained for the entire work. The three Editors therefore decided to rotate their names with each Volume, II to VII; but it must be emphasized again that the first-named Editor of each volume is simply primus inter pares and that the task of editing has been shared equally.

Systematic Treatment The past few years have been an exciting time for evolutionary ornithologists. Many new ideas about relationships among the higher taxa of birds have emerged, as a consequence of recent studies in anatomy (Burton 1985), palaeontology (Cracraft in press, Olson 1985) and biochemistry (Brush and Witt 1983, Sibley and Ahlquist 1985, 1986, Stapel et al. 1984). Of these, the DNA-DNA hybridization researches of C. G. Sibley and J.E. Ahlquist have been the most iconoclast­ ic. Respecting birds in this volume, some novel pro­ posals are that: turacos are not cuckoos but are distantly related to owls and nightjars; colies have been isolated from their nearest relatives for 100 million years; trogons are coraciiform, and African and American trogons compose separate subfamilies; hornbills and hoopoes should be separated from Coraciiformes as Bucerotiformes; wood-hoopoes and scimitarbills form two separate families; and woodpeckers and honey­ guides are more closely related to each other and to toucans and American barbers, than any of those families is to Old World barbers. Some proposals affecting other African birds are even more startling, for instance that larks are related to finches, and that pipits and accentors belong with the weavers in the Ploceidae. Such conclusions, if accepted, will dictate a radically new and heterodox classification of birds; but some are open to criticism (e.g. Houde 1987), and broad consens­ us amongst systematists seems unlikely to be achieved soon. For their part the Editors share the sentiments expressed by K. H. Voous, in Campbell and Lack (1985): "In this situation of uncertainty and hopeful expectation it will be understood that it has not been possible to follow all of the recently proposed changes in the classification of birds. More time is needed for their evaluation, before they are accepted and m­ corporated into standard lists and handbooks".

At low taxonomic levels we have introduced a few modest changes. Whilst treatment of superspecies gen­ erally follows Snow ( 1978), new evidence brought to light during the preparation of this Volume necessitates recognition of several new superspecies, for instance amongst turacos, nightjars and swifts. Contrarily, a few pairs of species, the two Ground Hornbills Bucorvus abyssinicus and B. cafer for example, whose ranges are now known to overlap, have been 'demoted' from a superspecies to independent species. A problem with superspecies stems from differing specific rates of morphological evolution. Turacos and bee-eaters illustrate the point. The two grey turacos, Crinifer piscator and C. zonurus (Plate 3), have very similar plumages and parapatric ranges; they compose a superspecies. Analysis of their feather proteins suggests that they diverged from each other less than 1.4 million years ago (Brush and Witt 1983). But the two parapatric violet turacos, Musophaga violacea and M. rossae (Plate 3), which have about the same degree of plumage similarity as the grey ones, are nearly ten times older: they diverged from each other 9.5-ll.5 million years ago (Brush and Witt 1983, and see van Tuinen and Valentine 1986). The bee-eaters Merops bullocki and M. bullockoides, which closely resemble each other and are parapatric, form a classical, woodland superspecies. But some evidences suggest that the parapatric Somali Bee­ eater M. revoilii belongs to the same superspecies (Plate 19) and may be of the same age (Fry 1984). If the three birds did diverge from one common ancestor about the same time, the faster evolution of M. revoilii probably owes to its invading and having to adapt to a new, sub­ desert environment. Since systematists have barely yet the means for determining such relationships, even our best efforts to delineate superspecies must fall far short of perfection.

Maps We have followed all the conventions established in Volume II in preparing and presenting the range maps. A full explanation of these appears in the introduction to that volume, but for convenience we have repro­ duced the explanatory maps here. Figure 1 shows the shading conventions used to denote the various cate­ gories of occurrence. Figure 2 updates the political map featured in Vo!ume I. The inset map, drawn at the same size as the range maps in the text, provides an extra aid for interpreting distributions defined by geographical co-ordinates. Figure 3 shows the five climatic zones of East Africa referred to under Laying Dates. Xlll

XIV

INTRODUCTION

English Names Some of the principles which have guided us in arriving at a single preferred English name for each species were set out in Volume II. Decisions continue to be trying. For Musophaga rossae, for instance, we have had to evaluate the relative merits of five widely-used regional names: Ross's Turaco, Ross's Lourie, Ross's Violet Loerie, Lady Ross's Turaco and Lady Ross's Violet Plantain-eater. Nightjars presented even worse prob­ lems, with taxonomic problems in addition to regional name variation. Wherever possible we have chosen existing names, but for a few birds have been compelled to coin quite new ones, like Swamp Nightjar for Caprimulgus natalensis and Square-tailed Nightjar for C.fossii. Our hope is, of course, that these names may in future stick in regional as well as in continental usage.

Fry, C. H. (1984). 'The Bee-eaters'. T. and A. D. Poyser, Calton. Houde, P. (1987). Critical evaluation of DNA hybridization studies in avian systematics. Auk 104, 17-32. Olson, S. L. (1985). The fossil record of birds. In 'Avian Biology' (Farner, D. S. and King, J. R. Eds) pp. 79-252. Academic Press, London and New York. Sibley, C. G. and Ahlquist, J.E. (1985). The relationships of some groups of African birds, based on comparisons of the genetic material, DNA. In 'African Vertebrates: Systematics, Phylogeny and Evolutionary Ecology' (Schuchmann, K.-L. Ed.) pp. 115-162. Selbstverlag, Bonn. Sibley, C. G. and Ahlquist, J.E. (1986). Reconstructing bird phylogeny by comparing DNAs. Sci. Amer. 254 (2), 82-92. Stapel, S. 0., Leunissen, J. A. M., Verspeeg, M., Watte), J. and de Jong, W.W. (1984). Ratites as oldest offshoot of avian stem-evidence from a-crystalline A sequences. Nature 311, 257-259. van Tuinen, P. and Valentine, M. (1986). Phylogenetic relationships of turacos (Musophagidae; Cuculiformes) based on comparative chromosome banding analysis. Ibis 128, 364-381.

Remarks by the Artist about the Plates I have paid careful attention to the design of the plates, to which there have been several constraints. First, I have shown birds in a variety of different postures so as to characterize the appearance in the field of each genus or family; second, I have tried to group birds according to affinity, resemblance, and geographical distribution (considerations which are sometimes conflicting); and third, I have attempted to use space as fully as possible so as to end up with good sized figures. Where plumage differences between species are slight and comparative structure is important, I have employed a more dia­ grammatic or 'field-guide' design, for example Plate 29 (in the forthcoming passerine volumes many plates will require that type of lay-out). Overall, I have aimed for plates which are aesthetically pleasing. All of these requirements, together with the need for all of the birds in a plate to be drawn to scale, have posed considerable design problems: designs of Plates 21, 22, 30 and 31 each took more than a week of drawing and re-drawing, before painting commenced!

References Brush, A. H. and Witt, H. H. (1983). Intraordinal relation­ ships of the Pelecaniformes and Cuculiformes: electro­ phoresis of feather keratins. Ibis 125, 181-199. Burton, P. J. K. (1985). Anatomy and evolution of the feeding apparatus in the avian orders Coraciiformes and Pici­ formes. Bull. Brit. Mus. ( Nat. Hist.) Zoo/. Ser. 47, 331-443. Campbell, B. and Lack,E. (1985). 'A Dictionary of Birds'. T. and A. D. Poyser, Calton. Cracraft, J. (in press). The clades of birds. In 'The Phylogeny of Tetrapods' (Benton, M. Ed.). Oxford University Press, Oxford.

D D

Dense breeding Sparse breeding lllllIIlll Migration Dense nan-breeding - Sparse nan-breeding ? Occurrence uncertain X Extralimital record Points to isolated occurrence

B

Migration route N.B. This tone - shows where breeding and non-breeding ranges overlap

Fig. 1. Shading, symbols and arrows used on maps.

Madeira-:

ocotra

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400 600 800 ,.lp00 miles

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INTRODUCTION

K E N Y A

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MOZAMBIQUE

Fi g. 3. Climatic zones of East Africa (Uganda, Kenya, Tanzania), referred to under specific Laying Dates (from Brown and Britton 1980 after F. Griffiths, 'Climatic zones of East Africa', 1958, 179-185).

2

Plate 2 (Opp. p. 33)

PSITTACIDAE

Psittacus erithacus Linnaeus. Grey Parrot. Perroquet gris. Psiuacus erichacus Linnaeus, 1758. Syst. Nat. (10th ed.), p. 99; Guinea. Range and Status. Endemic, mainly sedentary. Isolated populations in Guinea-Bissau (mainland and Bijagos Archipelago) and S Mali (Mandingo Mts and probably near Guinea border); otherwise widely dis­ tributed in Sierra Leone, S Guinea, Liberia, S Ivory Coast, Ghana, Togo, Benin, S Nigeria, S Cameroon, Equatorial Guinea (including Bioko), Principe (very abundant) and Sao Tome (probably introduced,sparse) (de Naurois 1983), Gabon,Congo, Zaire (except south), W Uganda, around L. Victoria, W Kenya (Kakamega, Nandi) and NW Tanzania (Bukoba). Mainly in low­ lands but up to 800 m in Cameroon, 2000 m in E Zai"re and 2200 m in E Africa. Common to abundant. Very popular as a cage bird, but no evidence of reduction in numbers due to trapping, except near towns. Traditional-site roosts of 5000-10 000, Gabon.

Psiltacus enlhacus

A have occasional scarlet feathers in grey plumage,especially on belly and thighs; rarely,birds almost entirely red. Albinos also reported. SIZE: (7 7