The Birds of Africa [4] 0121373045, 9780121373047

Table of contents :
Title
CONTENTS
ACKNOWLEDGEMENTS
LIST OF PLATES
INTRODUCTION
ORDER PASSERIFORMES
Eurylaimidae, broadbills (C. H. Fry)
Pittidae, pittas (C. H. Fry)
Alaudidae, larks (W.R. J. Dean, C. H. Fry, S. Keith and P. Lack)
Hirundinidae, swallows and martins (C. H. Fry, E. K. Urban and S. Keith)
Motacillidae, wagtails, pipits and longclaws (C. H. Fry, D. J. Pearson and P. B. Taylor)
Campephagidae, cuckoo-shrikes (D. J. Pearson and S. Keith)
Pycnonotidae, bulbuls (S. Keith)
Bombycillidae, waxwings, Grey Hypocolius and silky flycatchers (C. H. Fry)
Cinclidae, dippers (C. H. Fry)
Troglodytidae, wrens (C. H. Fry)
Prunellidae, accentors (C. H. Fry)
Turdidae, thrushes (W. R. J. Dean, C. Erard, C. H. Fry, V. Haas, R. A. C. Jensen,S. Keith, P. Lack, T. B. Oatley, D. J. Pearson, S. N. Stuart and A. Tye)
BIBLIOGRAPHY
General and Regional References
Family
References
Acoustic References
ERRATA, VOLUMES I, II AND III
INDEX
Scientific Names
English Names
French Names

Citation preview

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Edited by

STUART KEITH

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

EMIL K. URBAN

Department of Biology, Augusta College, Augusta, Georgi,a, USA

C. HILARY FRY

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

Colour P !ates by Martin Woodcock Line Drawings by Ian Willis Acoustic References by Claude Chappuis

ACADEMIC PRESS

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Copyright © 1992 by ACADEMIC PRESS LIMITED

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.

British Library Cataloguing in Publication Data The Birds of Africa. Vol. IV, Broadbills to chats. 1. Africa. Birds I. Keith, Stuart, 1931II. Urban, Emil K., 1934III. Fry, C. Hilary, 1937ISBN 0-12-137304-S

Editorial and production services by Fisher Duncan Ltd, 10 Barley Mow Passage, London W4 4PH Colour plates printed in the UK by George Over Ltd, Rugby Printed and bound in the UK by Mackays of Chatham, PLC, Chatham, Kent

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 measure­ ments 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. MiJlward, 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. Serie, 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. 0. 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

Vl

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 !sack, 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 lateR. 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 Biol. Gabonica (1971) (Andropadus virens, Phyllastrephus icterinus, Criniger calurus, Chlorocichla simplex), and in Terre Vie (1974) (Bleda 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. Mcllleron in 'Complete Book of Southern African Birds' (1989) (Cercotrichas leucophrys); H. Morand (Pseudochelidon eurystomina); 'Newman's Birds of South­ ern 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) (Macronyx 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.); J. 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

CONTENTS v

ACKNOWLEDGEMENTS LIST OF PLATES

lX

INTRODUCTION

x

ORDER PASSERIFORMES ..

1

Eurylaimidae, broadbills (C.H. Fry) Pittidae, pittas (C.H. Fry)

1

Alaudidae, larks (W.R.J.Dean, C.H. Fry, S.Keith and P.Lack) Hirundinidae, swallows and martins (C.H.Fry, E.K.Urban and S.Keith) Motacillidae, wagtails, pipits and longclaws (C.H. Fry, D.J.Pearson and P.B.Taylor) Campephagidae, cuckoo-shrikes (D.J.Pearson and S.Keith) Pycnonotidae, bulbuls (S.Keith) Bombycillidae, waxwings, Grey Hypocolius and silky flycatchers (C.H. Fry) Cinclidae, dippers (C.H. Fry) .. Troglodytidae, wrens (C.H. Fry) Prunellidae, accentors (C.H. Fry) Turdidae, thrushes (W. R. J. Dean, C. Erard, C. H. Fry, V. Haas, R. A. C. Jensen, S. Keith, P.Lack, T.B.Oatley, D.J.Pearson, S.N.Stuart and A.Tye)

9 13 125 197 263 279 377 380 382 384 387

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

557 561 587

ERRATA, VOLUMES I, II AND III

590

INDEXES 1. Scientific Names 2. English Names

591 604 608

3. French Names

vu

Vlll

CONTENTS

Authors' Contributions to Shared Families

ALAUDIDAE Dean and Keith: Mirafra (except M. pulpa, M. hypennetra, 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

Plate

Facing Page

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, Bleda)

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

Wagtails . .

176

27 Thrushes (scrub-robins and others)

416

12 Pipits, Sharpe's Longclaw

177

28 Thrushes (Phoenicurus, Saxicola)

417

13 Pipits

240

29 Wheatears

448

14 Longclaws

241

30 Wheatears

449

15 Cuckoo-Shrikes

256

31 Chats (Cercomela)

464

16 Waxwing, Hypocolius, Dipper, Wren, accentors, Pycnonotus bulbuls

257

32 Chats (Myrmecocichla)

465

11

ix

INTRODUCTION

species represents one further step in the process of differentiation from the megasubspecies of Amadon and Short (197 6), which refers to well-differentiated, con­ specific populations which 'approach the level of differ­ entiation of species ...but show by their interbreeding in contact or, if allopatric, provide some evidence ... that they are still conspecific, i.e. have not evolved effective reproductive isolation' (Short 1980). 'Super­ species' is a term used to designate a group of parapatric or essentially allopatric species (allospecies ) that have very recently evolved from a common ancestor, so re­ cently that they may be considered conspecific by some writers. When parapatric, allospecies sometimes hybridize in the zone of contact, but such hybridization reinforces the reproductive isolating mechanisms. Later, such species become fully reproductively isolated; on establishing contact they may either '(l) remain para­ patric, showing reproductive isolation but no ecological compatibility, hence competitive exclusion occurs; (2) overlap somewhat, showing reduced competition; or (3) overlap broadly with minimal interactions' (Short 1980). Such species comprise a 'species-group'. For allopatric populations, the taxonomist must still judge which stage they are in. Hall and Moreau (1970) emphasized the great value of the superspecies concept but did not apply it in the strict sense intended by Amadon (19 66). Their inter­ pretation tends to be broader, and while many of their groupings fall within the definition, others do not. As a result, some of their superspecies are being dismantled, either because allospecies have been found to be broadly sympatric without hybridization (e.g. Alethe polio­ cephala/A. poliophrys) or, in the case of allopatric species, because vocal and behavioural characters strongly sug­ gest lack of close relationship (e.g.Andropadus montanus/ A. masukuensis). It must be emphasized that the super­ species was never intended by its definers to be simply a convenient place to put species that show some degree of relationship. Some writers are becoming somewhat disenchanted with Hall and Moreau's superspecies (Vande weghe 1988, R. J. Dowsett, M. P. S. Irwin and L. L. Short, pers. comm.). Nevertheless, Pat Hall and the late Reg Moreau would probably be the first to admit that their groupings were tentative, and delighted at the controversies some are causing; that is how science progresses. One must, however, guard against the tendency to 'quote them as if they were holy writ' (M. P. S. Irwin, pers. comm.).

in Volume 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.

References Amadon, D. (1966). The superspecies concept. Syst. Zoo!. 15, 245-249. Amadon, D. and Short, L. L. (1976). Treatment of subspecies approaching species status. Syst. Zoo!. 25, 161-167. Campbell, B. and Lack, E. (1985). 'A Dictionary of Birds'. T. and A. D. Poyser, Calton. Hall, B. P. and Moreau, R. E. (1970). 'An Atlas of Speciation in African Passerine Birds'. British Museum (Natural His­ tory), London. Short, L. L. (1980). Speciation in African woodpeckers. Proc. IV Pan-Afr. Orn. Congr. pp. 1-8. Sibley, C. G. and Ahlquist, J.E. (1990). 'Phylogeny and Classi­ fication of Birds. A Study in Molecular Evolution'. Yale University Press, New Haven. Sibley, C. G. and Monroe, B. L. Jr. (1990). 'Distribution and Taxonomy of Birds of the World'. Yale University Press, New Haven. Vande weghe, J.-P. (1988). Problems in passerine speciation in Rwanda, Burundi and adjacent areas. Proc. XIX lnt. Orn. Congr. pp. 2547-2552.

D

Dense breeding

[l]Il]J] ? X ✓ /

Migration Dense non-breeding Sparse non-breeding Occurrence uncertain Extralimital record Points to isolated occurrence

□ Sparse breeding

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 reproduced the explanatory maps here. Figure 1 shows the shading conventions used to denote the various categories of occurrence. Figure 2 updates the political map featured

,,;J

Migration route

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

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

Xl

s --t-----+-cc::-t-----tt-----ffii------Jl----r-10°

0 0

200

400

500

600 800 1000 miles 1000

1500 km

TANZ ANIA

MOZAMBIQUE

Fig. 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).

Topography of African Passerine Birds I

Uppertail-

Forecrown

Crown

Hindcrown Nape

.,,

Hindneck

Secondaries

Median coverts Greater primary

UPPERWING­ COVERTS

Primaries Upper belly--

UNDERWING-COVERTS Greater under primary

Axillarie:;; pg

Superciliary stripe Undertail­ coverts �

Eye-ring

ind toe

P8

P5

Topography of African Passerine Birds II Forecrown crest

Mouth spots Orl:iital ring (featherlets)

Orbital ring (feathers)

Recurved !oral crest

Eye wattle

T1

Ornamental plumes

Breast band or pectoral band

Bare skin Cutting edges

Central streamers (T1)

Eye stripe

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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 Eurylaimidae 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 ltombwe Mts, Zafre, where known at 5 localities (lbachilo, Miki, Luvumba, Karungu, Muusi) between 3° 4l'S, 28° 3l'E and 3° 4'S, 28° 48'E and from 1760 to 2480 m; common around Miki. 80% of c. 30 birds col­ lected were at 1940-2390 m. Known also from mountains west ofL. Kivu, Zafre (Mt Kahuzi: Nyawaronga, 2° 0'S, 28° 49'E) and Impenetrable Forest, Uganda (Bwindi, Ruhizha, 2100-2200 m). Threatened by forest clearance around villages; L. Kivu population probably safe­ guarded in Kahuzi-Biega Nat. Park; other 2 areas need protection (Collar and Stuart 1985).

Pseudocalyptomena graueri

Description. ADULT d: 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 narrower­ billed than others (Friedmann and Williams 1968). Sexes alike. SIZE: (3 cfcf, 2 22) wing, cf 74·2, 75·5, 76, 2 75, 76·5; tail, cf 39, 39, 39·5, 2 39, 39; bill, cf 11-8, 14, 15, 2 15, 15; bill breadth at gape, cf 8·5, 11·5, 2 10·6, 10·6; tarsus, d 19, 20, 2 20, 20. WEIGHT: cf (n = 1) 32·5, 2 (n = 4) 29·0-32·5 (31· l). IMMATURE: like adult but slightly duller; undertail-coverts green. NESTLING: unknown.

Field Characters. Readily identified, even at considerable height in trees, by uniquely green and pale blue plumage: green upperparts, pale blue throat and undertail-coverts, pale green belly. Colours somewhat bee-eater-like, but short-tailed silhouette and stocky, upright stance are more flycatcher-like. Bill, short eye-mask, and underside of tail black.

Voice. Not tape-recorded. A very feeble 'tsi-tsi' repeated

3-8 times at rate of c. 4 notes per s; sounds like voice of Oriole Finch Linurgus olivaceus. Flight call of breeding bird an extremely high-pitched and quite prolonged bell-like ringing (E. F. G. Smith, pers. comm.). General Habits. Inhabits primary forest, occurring especially in middle storey and canopy, at edge of forest, at 1900-2400 m; also isolated trees in cleared ground. Usually avoids dense interior of forest. Perches mainly at 7-20 m, in dense foliage, sometimes quite immobile, like small barber. 1 bird climbed along a bough, clung

S mithornis

underneath it woodpecker-like, and clung to an upright bough (Chapin 1978) - perhaps feeding; also said to feed by sallying outward from canopy (Prigogine 1971), although these 'sallies' may in fact be display flights (Friedmann 1970). Bird at Ruhizha (Uganda), Nov 1986, gleaned insects actively, constantly fluttering up to pick insect from leaf, then landing briefly again. Another, in Chrysophyllum gorungosanum forest there, Dec 1987, flew from dry M imulopsis on log lying on ground, to join mixed foraging party of White-headed Wood-Hoopoes Phoeniculus bollei, Black-backed Puff­ backs Dryoscopus cub/a and Masked Apalis Apa/is binotata 7 m up; later, flock of 4-5 African Green Broadbills foraged actively 3-30 m high in forest, each perching usually for less than 15 s, then flying for only a few m, always calling in flight (T. Butynski and J. Kalina, pers. comm.). In Uganda said to feed more in manner of waxbill Estrilda in Neoboutonia trees and 2-3 m high in forest understorey. Not particularly shy; sometimes forages near human habitation. Flies up to 30 m between trees; flight direct, not undulating, rather slow, with gliding. Occurs singly, in pairs, or parties of up to 10. Often joins mixed-species bird parties.

Food. Seeds, flowers, buds, fruit, beetles, larvae and snails. Bird picked up 5-cm hairy caterpillar and dis­ carded it after 5 s. Breeding Habits. For possible display flight, see above. NEST: only 1 found was spherical, diam. c. 200-250, with round entrance c. 50 across, made of Spanish-moss­ like green lichen, sited c. 11 m up in 20 m tree in outer­ most branches and directly overhanging a stream; in valley floor with open shrubby vegetation and many 20 m high trees (Impenetrable Forest, Uganda: E. F. G. Smith, pers. comm.); 2 was in nest and cJ evidently fed her there. LAYING DATES: Zaire, 4 adults with somewhat enlarged gonads, late July; an immature bird collected at same time, so breeding season probably long. Uganda (pair at nest, Apr). References

Aspenlind, L. J. (1935). Collar, N. J. and Stuart, S. N. (1985). Friedmann, H. (1970). Prigogine, A. (1971).

Genus Smithornis Bonaparte Small, sexually dichromatic, streaky brown broadbills with white backs. 4-5 outer primaries straight and stiff, narrow­ based, with inner webs down-turned towards tip. Outermost primary covert elongated. Rictal bristles 8-10 mm long. Toes 3 and 4 united. Tongue thick. Short, circular display flight, unique among birds, accompanied by loud wing-rattling noise. Endemic; 3 species, 2 forming a superspecies (S. rufolateralis/S. capensis). Smithornis rufolateralis

1.

2.

S. rufolateralis S. capensis

superspecies

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4 Plate 1 (Opp. p. 32)

EURYLAIMIDAE

Smithornis sharpei Alexander. Grey-headed Broadbill. Eurylaime

a tete grise.

Smithomis sharpei Alexander, 1903. Bull. Br. Orn. Club 13, p. 34; Mt St Isabel, 4000 feet, Fernando Po. Range and Status. Endemic, resident. 2 populations: in

Smithornis sharpei

Bioko, Cameroon east to Yokadouma and N Gabon (Woleu-N'Tem region; 1 record Makokou); and in E Zaire from about Bondo Mabe, lturi, to Kanyaa, ltombwe. Frequent to common. Description. S. s. zenkeri (Reichenow): S Cameroon, N Gabon. ADULT d': forehead and crown dark grey, the feathers with blackish centres; forehead suffused with orange at sides and just above nostrils; nape, cheeks and sides of neck dark brown­ ish grey; mantle, back, rump and uppertail-coverts rufescent olive-brown, the mantle and back feathers with silky white bases, rufescent brown tips, and grey-black intervening areas (black and particularly white showing through irregularly); uppertail-coverts rich rufescent brown. Tail-feathers dark brown or blackish brown, with rufescent edges. Lores buff or orange buff; chin and throat white or creamy white in rnidline with a few diffuse dark grey streaks, and at sides (bordering cheeks) bright orange; breast bright rufous; belly silky white with fine or broad blackish streaks; upper belly, sides of belly and flanks strongly suffused tawny; undertail-coverts pale rufous. Primaries and secondaries blackish brown with rufous­ brown edges; scapulars and tertials rufescent brown; upperwing­ coverts dark brown with small rufous tips and narrow rufous edges. Underwing-coverts pale rufous; underside of rerniges and rectrices brown-grey. Upper mandible black, lower man­ dible pale greyish white; eye dark brown; legs and feet greenish grey or greyish olive. ADULT Cj): upperparts like d' but crown slightly greyer; underparts like d' but centre of chin and throat suffused buff and sides of chin and throat paler rufous; breast rufous-buff (rather than brick red), diffusely striped with grey; belly and flanks black-streaked white, washed buff, with only small area in rnidline free of buff wash. SIZE: (5 d'd', 6 Cj)Cj)) wing, d' 80-83 (81·6), Cj) 79-84 (81·4); tail, d' 54-58 (56·2), Cj) 50-54 (52·0); bill, d' 15-16 (15·4), Cj) 14-15 (14·7), breadth at bi!se, d'Cj) 13-14; tarsus, d'Cj) 16-17 (16·5). WEIGHT: un­ sexed (n = 6) 34-40·3 (37·5) (Bowden 1986). IMMATURE: differs from adult in having crown olive-brown, and white patch on back vestigial or absent; back feathers lack black marks; upperwing-coverts tipped orange-buff; breast streaked with black. NESTLING: unknown. S. s. sharpei Alexander: Bioko. Crown blue-grey (rather than sooty grey of zenkeri); black streaking on underparts less heavy. S. s. eurylaemus Neumann: E Zaire. d' with clearer grey crown and Cj) with sides of breast more olive-brown than in nominate race. Smaller: wing (d'Cj)) 75-78, bill (d'Cj)) 12-13.

Field Characters. The largest African broadbill, length

15 cm; a somewhat undistinguished bird with large grey head, buffy lores, white or whitish chin and throat, olive­ brown upperparts, bright rufous breast, whitish belly and streaked flanks. Rufous breast distinguishes it from African Broadbill S. capensis, and grey head from Rufous­ sided Broadbill S. rufolateralis. Upper mandible black, lower mandible white (like congeners); bill flat and broad. Back feathers have white bases, not always vis­ ible. Juvenile more uniformly olive-brown; breast duller rufous. Mechanical flight 'song', like congeners. Voice. Tape-recorded (78). Non-vocal sound made in

display flight is very loud, rapid, staccato trill of 0.5-

0.6 s duration, starting and finishing abruptly. Just per­ ceptibly, trill slows and then accelerates. Like trill of African Broadbill S. capensis but shorter and deeper­ pitched. General Habits. Inhabits lower and middle storeys of

primary lowland rain forest and heavily wooded mountain­ sides, from sea level (Cameroon), but mainly at 10001500 m on Bioko and in Itombwe. In pairs. Sluggish, sitting quietly for long periods. In evenings and at times during day, makes short display flight in tight circle at perch accompanied by loud, mechanical trill. Food. Unknown.

Breeding Habits

NEST: 4 described; large pocket-shaped mass of fresh green moss, suspended from twig 1-2 m above forest floor, with lining of dry leaves, fibres and stems. 1 nest built entirely of fine black fungal fibres and dried leaves. Height 22 cm, diam. 9-10 cm, entrance hole 6 x 3·5 cm, on upper side, once with porch. EGGS: 1-2. Long, pointed ovals; pure white. SIZE: (n = 7) 22-25 X 16-17·5 (23·3 X 16·9). LAYING DATES: Cameroon, Jan, Mar, and brood­ patch Feb; Zaire, Apr.

Smithornis rufolateralis

Smithornis rufolateralis Gray. Rufous-sided Broadbill; Red-sided Broadbill. Eurylaime

a flancs roux.

Smithornis rufolateralis Gray, 1864. Proc. Zool. Soc. Lond., p. 143; Gold Coast. Forms a superspecies with S. capensis. Range and Status. Endemic resident, lowland forests from Liberia to W Uganda, south to Kananga (Lulua­ burg) in Zai:re. Frequent to common; probably more widespread than shown on map. Density of 12 pairs per km2 (M'Passa, Gabon).

Smithornis rufolateralis

Description. S. r. rufolateralis Gray: Liberia to about 20° E in

Zaire. ADULT d': forehead, crown, nape, hindneck, sides of neck and cheeks black, sometimes with small line of white below eye; mantle black in centre, brown at sides, back and rump black; black feathers of mantle to rump have white bases which sometimes show irregularly through. Rump ochreous dark brown, tail dull black, the tail-feathers narrowly edged brown. Lores buff. Long, soft, black rictal bristles. Chin and throat creamy white, occasionally with irregular dark streaks; sides of throat and breast creamy white, heavily streaked black; large area at sides of breast orange; belly cream-white, heavily streaked black except in midline and around vent. Flanks cream-white heavily streaked black, or uniformly brown where concealed below wing. Remiges dull blackish, edged with brown. Scapulars and tertials dark ochreous brown; lesser covens dark ochreous brown; remaining upperwing-coverts black or blackish; median and greater primary covens with triangular or tear-shaped white tips, forming 2 rows of white spots on closed wing. Underwing-coverts creamy white; under­ side of remiges and rectrices grey. Upper mandible black, lower mandible white or horn, tinged pink; eye dark brown; legs and feet olive green. ADULT 2: forehead, crown and nape greyish or blackish brown; sides of neck and cheeks warm ochreous brown, mantle, back, rump, uppertail-coverts, scapu­ lars and tertials warm ochreous brown; mantle and back with white feather bases showing through irregularly; mantle feathers white proximally, brown distally, with black intervening mark. Rest of plumage like adult d', but sides of breast orange-brown rather than bright orange, greater and median primary covens brown-black with smaller, occasionally huffier, white tips, and tail browner. SIZE: (41 d'd', 13 2 Cjl) wing, cJ 60-72 (66·7), 2 60-65 (63· l); tail, d' 40-43 (41·8), Cjl 38-40 (39·5); bill, cJ 13-17 (15·7), 2 13-16 (14·9), breadth at base, d' 13, 2 12; tarsus, unsexed 13-15 (14·0). WEIGHT: (Liberia) d' (n = 12) 20·5 ± 2·7, 2 (n = 6) 10-23·1 (17·7). IMMATURE: like adult Cjl but somewhat duller. NESTLING: unknown. S. r. budongoensis van Someren: central Zai:re to W Uganda. Dark stripes on underside fewer and shorter; crown of 2 greyer than that of nominate 2. Smaller. WEIGHT: (Uganda) 2 (n = 8) 19-29 (20·9).

Voice. Tape-recorded (78). Voice, used in courtship, is short, weak, high-pitched whistle 'whee, whee . . . .' or 'huiiii'. Much more frequent noise is mechanical flight display (see above), a trill exactly like that of African Broadbill (q.v.) but higher-pitched.

Field Characters. A flycatcher-like bird of thick under­ growth. d distinguished by: black head and cheeks, sharply demarcated from buff lores and white chin and throat; brown upperparts with back irregularly white (hard to see at rest, can be very conspicuous in flight) and 2 white wing-bars; and white underparts, with dark­ streaked breast and flanks and large bright orange patch on side of breast. Bill flat and broad, upper mandible black, lower whitish (like congeners). S? similar, duller, less white in back, wing-bars huffy, and head brown where d is black. Commonly makes display flight ac­ companied by loud, mechanical rattle (see Voice); rattle

General Habits. Inhabits thick undergrowth in lower storey and at edges of lowland forests from sea level to 1300 m (1500 m on Mt Nyombe, E Zai:re); often near water. Restricted to primary and old secondary forests, so locally range complements that of African Broadbill, q.v. Occurs solitarily or in pairs, d and S? foraging some m apart; sits quietly on thin horizontal branch or creeper 3-15 m above ground in thick vegetation with long periods of immobility. Gives rapid chase to passing insect, through open spaces and among foliage, up to 25 m above ground; also picks insect from leaf or trunk in flight. Sometimes, as bird twists and turns in pursuit of prey, wings make toad-like noise of flight display.

higher-pitched than that of African Broadbill S. capensis and much higher-pitched than rattle of Grey-headed Broadbill S. sharpei. Flycatching behaviour like wattle­ eye Platysteira. Orange sides of breast distinguish it from African Broadbill; black (not grey) head and white wing­ bars distinguish d from d Grey-headed Broadbill; S? told from Grey-headed Broadbill by small size (length 12 cm) and 2 huffy wing-bars. All 3 species are partly sympatric.

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Plate 1 (Opp. p. 32)

6

EURYLAIMIDAE

Flight display performed solitarily, by both sexes but mainly by cf', at all times of day and year, but particu­ larly at dawn and dusk in all but drier months. Fre­ quency of 1 flight per min for 1 h, flights performed rather regularly, but 28 flights in 14 min by bird re­ sponding to sound of displaying neighbours (flights are also made regularly in response to playback of tape­ recording). Flights much less frequent in middle of day. Flight display performed from horizontal branch or creeper, 5-15 m above ground. Bird makes 1-2 abrupt jumps into air (sometimes turning about as it does so), then flies in ellipse of 1 m greatest diam., making burst of sound audible 1 SO m away, returning to same point on perch. Sound is made by vibration of stiff, twisted outer primaries. During flight, white back feathers puffed out. Plane of ellipse varies from horizontal to inclination of 45° the angle of inclination modifying timbre of sound. Successive display flights are made from same perch, bird sometimes giving weak 'huiiii' call between flights; or (usually) from sequence of dozens of perches as bird travels up to 1500 m in 15 min. Displaying bird usually solitary, and flight serves territorial function (Brosset and Erard 1986); display flights also sometimes given during pair-formation and before copulation. Flight frequency increases when rival heard or seen near territory border, but birds do not fight. Bird responding to another at border of territory puffs out plumage, orange patches being conspicuous, wags closed tail slowly up and down through wide arc, trembles wing­ tips, gives thin, stifled calls, and makes several abrupt jumps into air. Food. Arthropods. Small cicadas and small, hairless caterpillars commonest; beetles, grasshoppers, and an earwig, spider and millipede also recorded.

Monogamous, territorial. Courting cf' approaches