Mammals of Africa Volume VI: Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer and Bovids 9781472926968, 9781408189955

Table of contents :
Mammals of Africa Volume VI
Cover
Copyright
Contents
Series Acknowledgements
Acknowledgements for Volume VI
Mammals of Africa: An Introduction and Guide
ORDER CETARTIODACTYLA Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer, Bovids
SUBORDER SUIFORMES Pigs
SUPERFAMILY SUOIDEA Pigs
FAMILY SUIDAE Pigs, Hogs
SUBFAMILY SUINAE Pigs, Hogs
TRIBE SUINI Old World Pigs
GENUS Sus Wild Boar
Sus scrofa Wild Boar (Eurasian Wild Pig)
GENUS Potamochoerus – Bushpig, Red River Hog
Potamochoerus larvatus Bushpig
Potamochoerus porcus Red River Hog
GENUS Hylochoerus Forest Hog
Hylochoerus meinertzhageni Forest Hog (Giant Forest Hog)
TRIBE PHACOCHOERINI Warthogs
GENUS Phacochoerus Warthogs
Phacochoerus aethiopicus Desert Warthog
Phacochoerus africanus Common Warthog
SUBORDER WHIPPOMORPHA Hippopotamuses, Cetaceans
INFRAORDER ANCODONTA Hippopotamuses
FAMILY HIPPOPOTAMIDAE Hippopotamuses
GENUS Hippopotamus Common Hippopotamus
Hippopotamus amphibius Common Hippopotamus
GENUS Choeropsis Pygmy Hippopotamus
Choeropsis liberiensis Pygmy Hippopotamus
SUBORDER RUMINANTIA Ruminants
INFRAORDER PECORA Horned Ruminants
SUPERFAMILY GIRAFFOIDEA Giraffe, Okapi
FAMILY GIRAFFIDAE Giraffe, Okapi
SUBFAMILY GIRAFFINAE Giraffe
GENUS Giraffa Giraffe
Giraffa camelopardalis Giraffe
SUBFAMILY OKAPINAE Okapi
GENUS Okapia Okapi
Okapia johnstoni Okapi
SUPERFAMILY CERVOIDEA Deer
FAMILY CERVIDAE Deer
SUBFAMILY CERVINAE Old World Deer
GENUS Cervus Red Deer
Cervus elaphus Red Deer (Barbary Red Deer)
SUPERFAMILY BOVOIDEA Bovoids
FAMILY BOVIDAE Bovines, Antilopines
SUBFAMILY BOVINAE African Buffalo, Spiral-horned Antelopes
TRIBE BOVINI African Buffalo
GENUS Syncerus African Buffalo
Syncerus caffer African Buffalo
TRIBE TRAGELAPHINI Spiral-horned Antelopes
GENUS Tragelaphus Spiral-horned Antelopes
Tragelaphus imberbis Lesser Kudu
Tragelaphus angasii Nyala
Tragelaphus strepsiceros Greater Kudu
Tragelaphus buxtoni Mountain Nyala (Gedemsa)
Tragelaphus scriptus Bushbuck
Tragelaphus spekii Sitatunga
Tragelaphus eurycerus Bongo
Tragelaphus derbianus Giant Eland (Lord Derby’s Eland)
Tragelaphus oryx Common Eland
SUBFAMILY ANTILOPINAE Antelopes, Sheep, Goats
TRIBE NEOTRAGINI Dwarf Antelopes
GENUS Neotragus Dwarf Antelopes
Neotragus batesi Bates’s Pygmy Antelope (Dwarf Antelope, Bates’s Dwarf Antelope)
Neotragus pygmaeus Royal Antelope
GENUS Nesotragus Suni
Nesotragus moschatus Suni
TRIBE CEPHALOPHINI Duikers
GENUS Philantomba Blue Duikers
Philantomba maxwelli Maxwell’s Duiker
Philantomba monticola Blue Duiker
GENUS Sylvicapra Common Duiker
Sylvicapra grimmia Common Duiker (Grey Duiker, Bush Duiker, Grimm’s Duiker)
GENUS Cephalophus Forest Duikers
Cephalophus zebra Zebra Duiker (Banded Duiker)
Cephalophus adersi Aders’s Duiker
Cephalophus rubidus Rwenzori Red Duiker
Cephalophus leucogaster White-bellied Duiker
Cephalophus natalensis Natal Red Duiker
Cephalophus harveyi Harvey’s Duiker
Cephalophus rufilatus Red-flanked Duiker
Cephalophus nigrifrons Black-fronted Duiker
Cephalophus ogilbyi Ogilby’s Duiker
Cephalophus weynsi Weyns’s Duiker
Cephalophus callipygus Peters’s Duiker
Cephalophus niger Black Duiker
Cephalophus spadix Abbott’s Duiker
Cephalophus silvicultor Yellow-backed Duiker
Cephalophus dorsalis Bay Duiker
Cephalophus jentinki Jentink’s Duiker
TRIBE RAPHICERINI Grysboks, Steenbok, Beira
GENUS Raphicerus Grysboks, Steenbok
Raphicerus melanotis Cape Grysbok
Raphicerus sharpei Sharpe’s Grysbok
Raphicerus campestris Steenbok (Steinbuck, Steinbok)
GENUS Dorcatragus Beira
Dorcatragus megalotis Beira
TRIBE MADOQUINI Dik-diks
GENUS Madoqua Dik-diks
Madoqua saltiana Salt’s Dik-dik
Madoqua piacentinii Silver Dik-dik (Piacentini’s Dik-dik)
Madoqua (kirkii) Kirk’s Dik-dik Species Group
Madoqua (kirkii) kirkii Kirk’s Dik-dik
Madoqua (kirkii) cavendishi Naivasha Dik-dik (Cavendish’s Dik-dik)
Madoqua (kirkii) thomasi Ugogo Dik-dik (Thomas’s Dik-dik)
Madoqua (kirkii) damarensis Damara Dik-dik
Madoqua guentheri Günther’s Dik-dik
TRIBE ANTILOPINI Gazelline Antelopes
GENUS Gazella Slender Gazelles
Gazella dorcas Dorcas Gazelle
Gazella spekei Speke’s Gazelle
Gazella cuvieri Cuvier’s Gazelle (Atlas Gazelle, Edmi Gazelle)
Gazella leptoceros Slender-horned Gazelle (Rhim Gazelle, Loder’s Gazelle)
GENUS Eudorcas Ring-horned Gazelles
Eudorcas rufifrons Red-fronted Gazelle
Eudorcas tilonura Heuglin’s Gazelle
Eudorcas thomsonii Thomson’s Gazelle
Eudorcas albonotata Mongalla Gazelle
GENUS Nanger Greater Gazelles
Nanger (granti) Grant’s Gazelle Species Group
Nanger (granti) granti Grant’s Gazelle
Nanger (granti) notata Bright’s Gazelle
Nanger (granti) petersii Peters’s Gazelle (Tana Gazelle)
Nanger soemmerringi Soemmerring’s Gazelle
Nanger dama Dama Gazelle (Addra Gazelle)
GENUS Ammodorcas Dibatag
Ammodorcas clarkei Dibatag (Clarke’s Gazelle)
GENUS Litocranius Gerenuk
Litocranius walleri Gerenuk (Waller’s Gazelle)
GENUS Antidorcas Springbok
Antidorcas marsupialis Springbok (Springbuck)
TRIBE OUREBIINI Oribi
GENUS Ourebia Oribi
Ourebia ourebi Oribi
TRIBE REDUNCINI Reduncines
GENUS Pelea Grey Rhebok
Pelea capreolus Grey Rhebok (Vaal Rhebok, Rhebok)
GENUS Redunca Reedbucks
Redunca fulvorufula Mountain Reedbuck
Redunca arundinum Southern Reedbuck
Redunca redunca Bohor Reedbuck (Common Reedbuck)
GENUS Kobus Kobs
Kobus kob Kob
Kobus vardonii Puku
Kobus leche Southern Lechwe
Kobus megaceros Nile Lechwe (Mrs Gray’s Lechwe)
Kobus ellipsiprymnus Waterbuck
TRIBE OREOTRAGINI Klipspringer
GENUS Oreotragus Klipspringer
Oreotragus oreotragus Klipspringer
TRIBE AEPYCEROTINI Impala
GENUS Aepyceros Impala
Aepyceros melampus Impala
TRIBE ALCELAPHINI Alcelaphines
GENUS Beatragus Hirola
Beatragus hunteri Hirola
GENUS Damaliscus Damalisks
Damaliscus pygargus Bontebok / Blesbok
Damaliscus lunatus Topi / Tsessebe / Tiang / Korrigum
GENUS Alcelaphus Hartebeest
Alcelaphus buselaphus Hartebeest
GENUS Connochaetes Wildebeest
Connochaetes gnou Black Wildebeest (White-tailed Gnu)
Connochaetes taurinus Common Wildebeest
TRIBE HIPPOTRAGINI Horse-like Antelopes
GENUS Hippotragus Roan and Sable Antelopes
Hippotragus equinus Roan Antelope
Hippotragus niger Sable Antelope
GENUS Addax Addax
Addax nasomaculatus Addax
GENUS Oryx Oryxes
Oryx gazella Gemsbok (Southern Orxy)
Oryx beisa Beisa Oryx (Fringe-eared Oryx)
Oryx dammah Scimitar-horned Oryx (Scimitar Oryx)
TRIBE CAPRINI Sheep, Goats
GENUS Ammotragus Aoudad
Ammotragus lervia Aoudad (Barbary Sheep, Arui)
GENUS Capra Ibexes
Capra nubiana Nubian Ibex
Capra walie Walia Ibex (Ethiopian Ibex)
INFRAORDER TRAGULINA Chevrotains
SUPERFAMILY TRAGULOIDEA Chevrotains
FAMILY TRAGULIDAE Chevrotains
GENUS Hyemoschus Water Chevrotain
Hyemoschus aquaticus Water Chevrotain
Glossary
Bibliography
Authors of Volume VI
Indexes
French names
German names
English names
Scientific names

Citation preview

mammals of africa volume VI

pigs, hippopotamuses, chevrotain, giraffes, deer and bovids

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Series Editors Jonathan Kingdon Department of Zoology, University of Oxford David C. D. Happold Research School of Biology, Australian National University Thomas M. Butynski Zoological Society of London/King KhalidWildlife Research Centre Michael Hoffmann International Union for Conservation of Nature – Species Survival Commission Meredith Happold Research School of Biology, Australian National University Jan Kalina Soita Nyiro Conservancy, Kenya

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mammals of africa volume VI

pigs, hippopotamuses, chevrotain, giraffes, deer and bovids edited by jonathan kingdon and michael hoffmann

Illustrated by Jonathan Kingdon

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First published in 2013 Copyright © 2013 by Bloomsbury Publishing Copyright © 2013 illustrations by Jonathan Kingdon All rights reserved. No part of this publication may be reproduced or used in any form or by any means –photographic, electronic or mechanical, including photocopying, recording, taping or information storage or retrieval systems – without permission of the publishers. Bloomsbury Publishing Plc, 50 Bedford Square, London WC1B 3DP Bloomsbury USA, 175 Fifth Avenue, New York, NY 10010 www.bloomsbury.com www.bloomsburyusa.com Bloomsbury Publishing, London, New Delhi, New York and Sydney A CIP catalogue record for this book is available from the British Library. Library of Congress Cataloging-in-Publication Data has been applied for. Commissioning editor: Nigel Redman Design and project management: D & N Publishing, Baydon, Wiltshire ISBN (print) 978-1-4081-2256-3 ISBN (epdf) 978-1-4081-8995-5 Printed in China by C&C Offset Printing Co., Ltd This book is produced using paper that is made from wood grown in managed sustainable forests. It is natural, renewable and recyclable. The logging and manufacturing processes conform to the environmental regulation of the country of origin. 10 9 8 7 6 5 4 3 2 1

Recommended citations: Series: Kingdon, J., Happold, D., Butynski, T., Hoffmann, M., Happold, M. & Kalina, J. (eds) 2013. Mammals of Africa (6 vols). Bloomsbury Publishing, London. Volume: Kingdon, J. & Hoffmann, M. (eds) 2013. Mammals of Africa.VolumeVI: Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer and Bovids. Bloomsbury Publishing, London. Chapter/species profile: e.g. Klingel, H. 2013. Hippopotamus amphibius Common Hippopotamus; pp 68–77 in Kingdon, J. & Hoffmann, M. (eds) 2013. Mammals of Africa:VolumeVI: Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer and Bovids. Bloomsbury Publishing, London.

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Donors and Patrons T. R. B. Davenport, D. De Luca and the Wildlife Conservation Society, Tanzania R. Dawkins R. Farrand & L. Snook R. Heyworth, S. Pullen and the Cotswold Wildlife Park G. Ohrstrom Viscount Ridley & M. Ridley L. Scott and the Smithsonian UK Charitable Trust M. & L. Ward R. & M. Ward

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Contents Series Acknowledgements10 Acknowledgements for Volume VI11 Mammals of Africa: An Introduction and Guide – David Happold, Michael Hoffmann, Thomas Butynski & Jonathan Kingdon

13

ORDER CETARTIODACTYLA Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer, Bovids – E. R. Seiffert & J. Kingdon

GENUS Hippopotamus Common Hippopotamus – S. K. Eltringham, J. Kingdon & J.-R. Boisserie Hippopotamus amphibius Common Hippopotamus – H. Klingel 

64 68

GENUS Choeropsis Pygmy Hippopotamus – J.-R. Boisserie & S. K. Eltringham Choeropsis liberiensis Pygmy Hippopotamus – P. T. Robinson

78 80

22

SUBORDER RUMINANTIA Ruminants – R. R. Hofmann & J. Kingdon

84

SUBORDER SUIFORMES Pigs – J. M. Harris

24

INFRAORDER TRAGULINA Chevrotains – J. Kingdon

86

SUPERFAMILY SUOIDEA Pigs – J. M. Harris

24

SUPERFAMILY TRAGULOIDEA Chevrotains 

87

FAMILY SUIDAE Pigs, Hogs – C. Groves & J. M. Harris

25

FAMILY TRAGULIDAE Chevrotains – A. Gentry

87

SUBFAMILY SUINAE Pigs, Hogs – J. M. Harris

25

GENUS Hyemoschus Water Chevrotain – P. Grubb Hyemoschus aquaticus Water Chevrotain – J. A. Hart

88 88

TRIBE SUINI Old World Pigs – J. M. Harris

27

GENUS Sus Wild Boar – J. M. Harris Sus scrofa Wild Boar (Eurasian Wild Pig) – F. Cuzin & E. Randi

28

INFRAORDER PECORA Horned Ruminants – C. Janis & J. Kingdon

93

SUPERFAMILY GIRAFFOIDEA Giraffe, Okapi – J. Kingdon & J. M. Harris

94

GENUS Potamochoerus – Bushpig, Red River Hog – C. Groves Potamochoerus larvatus Bushpig – A. H. W. Seydack Potamochoerus porcus Red River Hog – K. Leus & P. Vercammen

31 32

FAMILY GIRAFFIDAE Giraffe, Okapi – A. Gentry

95

37

SUBFAMILY GIRAFFINAE Giraffe

96

GENUS Hylochoerus Forest Hog – J. Kingdon Hylochoerus meinertzhageni Forest Hog (Giant Forest Hog) – J.-P. d’Huart & J. Kingdon

40

GENUS Giraffa Giraffe – R. Seymour Giraffa camelopardalis Giraffe – I. Ciofolo & Y. Le Pendu

96 98

TRIBE PHACOCHOERINI Warthogs – J. M. Harris

49

GENUS Phacochoerus Warthogs – P. Grubb Phacochoerus aethiopicus Desert Warthog – P. Grubb & J.-P. d’Huart Phacochoerus africanus Common Warthog – D. H. M. Cumming SUBORDER WHIPPOMORPHA Hippopotamuses, Cetaceans – E. R. Seiffert & J. Kingdon

28

42 SUBFAMILY OKAPINAE Okapi

110

50

GENUS Okapia Okapi – P. Grubb Okapia johnstoni Okapi – J. A. Hart

110 110

51

SUPERFAMILY CERVOIDEA Deer – J. Kingdon

115

54

FAMILY CERVIDAE Deer – C. Groves

116

SUBFAMILY CERVINAE Old World Deer

116

GENUS Cervus Red Deer – V. Geist Cervus elaphus Red Deer (Barbary Red Deer) – V. Geist

117 117

SUPERFAMILY BOVOIDEA Bovoids

120

61

INFRAORDER ANCODONTA Hippopotamuses – J. M. Harris

62

FAMILY HIPPOPOTAMIDAE Hippopotamuses – A. Gentry

63

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Contents

FAMILY BOVIDAE Bovines, Antilopines – C. Groves

120

SUBFAMILY BOVINAE African Buffalo, Spiral-horned Antelopes – J. Kingdon

122

TRIBE BOVINI African Buffalo – J. Kingdon

124

GENUS Syncerus African Buffalo – P. van Hooft & H. H. T. Prins Syncerus caffer African Buffalo – H. H. T. Prins & A. R. E. Sinclair TRIBE TRAGELAPHINI Spiral-horned Antelopes – J. Kingdon GENUS Tragelaphus Spiral-horned Antelopes – J. Kingdon Tragelaphus imberbis Lesser Kudu – W. Leuthold Tragelaphus angasii Nyala ­– J. Anderson Tragelaphus strepsiceros Greater Kudu – N. Owen-Smith Tragelaphus buxtoni Mountain Nyala (Gedemsa) – C. Sillero-Zubiri Tragelaphus scriptus Bushbuck – A. J. Plumptre & T. Wronski Tragelaphus spekii Sitatunga – J. May & R. Lindholm Tragelaphus eurycerus Bongo – P. W. Elkan & J. L. D. Smith Tragelaphus derbianus Giant Eland (Lord Derby’s Eland) – H. P. Planton & I. G. Michaux Tragelaphus oryx Common Eland – C. R. Thouless

124 125 137 138 142 148 152 159 163 172 179 186 191

SUBFAMILY ANTILOPINAE Antelopes, Sheep, Goats – J. Kingdon

199

TRIBE NEOTRAGINI Dwarf Antelopes – J. Kingdon

206

GENUS Neotragus Dwarf Antelopes – P. Grubb Neotragus batesi Bates’s Pygmy Antelope (Dwarf Antelope, Bates’s Dwarf Antelope) – F. Feer Neotragus pygmaeus Royal Antelope – J. Kingdon & M. Hoffmann

207

GENUS Nesotragus Suni – J. Kingdon Nesotragus moschatus Suni – J. Kingdon & M. Hoffmann

213 214

TRIBE CEPHALOPHINI Duikers – J. Kingdon & C. Groves

220

Cephalophus rubidus Rwenzori Red Duiker – J. Kingdon 253 Cephalophus leucogaster ­White-bellied Duiker – J. A. Hart 255 Cephalophus natalensis Natal Red Duiker – M. Hoffmann & A. E. Bowland 258 Cephalophus harveyi Harvey’s Duiker – J. Kingdon & F. Rovero 261 Cephalophus rufilatus Red-flanked Duiker – J. Kingdon & M. Hoffmann 265 Cephalophus nigrifrons Black-fronted Duiker – A. J. Plumptre 268 Cephalophus ogilbyi Ogilby’s Duiker – J. Kingdon 272 Cephalophus weynsi Weyns’s Duiker – J. A. Hart 275 Cephalophus callipygus Peters’s Duiker – F. Feer & M. Mockrin 279 Cephalophus niger Black Duiker – J. Kingdon & M. Hoffmann 281 Cephalophus spadix Abbott’s Duiker – F. Rovero, T. R. B. Davenport & T. Jones 285 Cephalophus silvicultor Yellow-backed Duiker – J. Kingdon & S. A. Lahm  288 Cephalophus dorsalis Bay Duiker – J. Kingdon & F. Feer 294 Cephalophus jentinki Jentink’s Duiker – B. Hoppe-Dominik 299 TRIBE RAPHICERINI Grysboks, Steenbok, Beira – J. Kingdon

302

GENUS Raphicerus Grysboks, Steenbok – P. Grubb Raphicerus melanotis Cape Grysbok – G. Castley & P. Lloyd Raphicerus sharpei Sharpe’s Grysbok – M. Hoffmann & V. J. Wilson Raphicerus campestris Steenbok (Steinbuck, Steinbok) – J. T. du Toit 

303 304

GENUS Dorcatragus Beira – P. Grubb Dorcatragus megalotis Beira – N. Giotto, A. Laurent & T. Künzel

315

TRIBE MADOQUINI Dik-diks – J. Kingdon

319

308 311

315

208 211

GENUS Philantomba Blue Duikers – C. Groves 223 Philantomba maxwelli Maxwell’s Duiker – D. Nett & H. Newing 224 Philantomba monticola Blue Duiker – J. A. Hart & J. Kingdon 228 GENUS Sylvicapra Common Duiker – C. Groves Sylvicapra grimmia Common Duiker (Grey Duiker, Bush Duiker, Grimm’s Duiker) – V. J. Wilson

235

GENUS Cephalophus Forest Duikers – J. Kingdon Cephalophus zebra Zebra Duiker (Banded Duiker) – B. Hoppe-Dominik Cephalophus adersi Aders’s Duiker – A. Williams

244

GENUS Madoqua Dik-diks – J. Kingdon 320 Madoqua saltiana Salt’s Dik-dik – D. W.Yalden 323 Madoqua piacentinii Silver Dik-dik (Piacentini’s Dik-dik) – A. M. Simonetta  325 Madoqua (kirkii) Kirk’s Dik-dik Species Group – P. N. M. Brotherton  327 Madoqua (kirkii) kirkii Kirk’s Dik-dik 328 Madoqua (kirkii) cavendishi Naivasha Dik-dik (Cavendish’s Dik-dik)329 Madoqua (kirkii) thomasi Ugogo Dik-dik (Thomas’s Dik-dik) 329 Madoqua (kirkii) damarensis Damara Dik-dik  329 Madoqua guentheri Günther’s Dik-dik – P. P. Hoppe & P. N. M. Brotherton 334

235

245 248

TRIBE ANTILOPINI Gazelline Antelopes – J. Kingdon

338

GENUS Gazella Slender Gazelles – C. Groves Gazella dorcas Dorcas Gazelle – P. Scholte & I. M. Hashim  Gazella spekei Speke’s Gazelle – T. L. Thurow

339 340 346 7

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Contents

Gazella cuvieri Cuvier’s Gazelle (Atlas Gazelle, Edmi Gazelle) – R. C. Beudels, P. Devillers & F. Cuzin Gazella leptoceros Slender-horned Gazelle (Rhim Gazelle, Loder’s Gazelle) – R. C. Beudels & P. Devillers 

349

Kobus megaceros Nile Lechwe (Mrs Gray’s Lechwe) – E. Falchetti & J. Kingdon Kobus ellipsiprymnus Waterbuck – C. A. Spinage

455 461

TRIBE OREOTRAGINI Klipspringer – J. Kingdon

469

GENUS Oreotragus Klipspringer – P. Grubb Oreotragus oreotragus Klipspringer – S. C. Roberts

469 470

TRIBE AEPYCEROTINI Impala – J. Kingdon

477

GENUS Aepyceros Impala – J. Kingdon Aepyceros melampus Impala – H. Fritz & M. Bourgarel

479 480

TRIBE ALCELAPHINI Alcelaphines – L. M. Gosling & J. Kingdon

488

GENUS Beatragus Hirola – T. M. Butynski & J. Kingdon Beatragus hunteri Hirola – T. M. Butynski

490 491

GENUS Damaliscus Damalisks – P. Duncan Damaliscus pygargus Bontebok / Blesbok – J. David & P. Lloyd Damaliscus lunatus Topi / Tsessebe / Tiang / Korrigum – P. Duncan

495

GENUS Alcelaphus Hartebeest – L. M. Gosling  Alcelaphus buselaphus Hartebeest – L. M. Gosling & I. Capellini

510

527

544

352

GENUS Eudorcas Ring-horned Gazelles – C. Groves Eudorcas rufifrons ­Red-fronted Gazelle – P. Scholte & I. M. Hashim Eudorcas tilonura Heuglin’s Gazelle – I. M. Hashim Eudorcas thomsonii Thomson’s Gazelle – C. FitzGibbon & J. Wilmshurst Eudorcas albonotata Mongalla Gazelle – I. M. Hashim & J. Kingdon

356

GENUS Nanger Greater Gazelles – C. Groves Nanger (granti) Grant’s Gazelle Species Group – H. R. Siegismund, E. D. Lorenzen & P. Arctander  Nanger (granti) granti Grant’s Gazelle Nanger (granti) notata Bright’s Gazelle Nanger (granti) petersii Peters’s Gazelle (Tana Gazelle) Nanger soemmerringi Soemmerring’s Gazelle – C. Schloeder & M. Jacobs Nanger dama Dama Gazelle (Addra Gazelle) – P. Scholte

372

GENUS Ammodorcas Dibatag – P. Grubb Ammodorcas clarkei Dibatag (Clarke’s Gazelle) – F. K. Wilhelmi

387

357 359 361 369

373 374 375 376 380 382

388

GENUS Litocranius Gerenuk – P. Grubb  390 Litocranius walleri Gerenuk (Waller’s Gazelle) – W. Leuthold 391 GENUS Antidorcas Springbok – J. D. Skinner Antidorcas marsupialis Springbok (Springbuck) – J. D. Skinner

398 398

GENUS Connochaetes Wildebeest – R. D. Estes Connochaetes gnou Black Wildebeest (White-tailed Gnu) – S. Vrahimis Connochaetes taurinus Common Wildebeest – R. D. Estes

TRIBE OUREBIINI Oribi – J. Kingdon

404

TRIBE HIPPOTRAGINI Horse-like Antelopes – J. Kingdon

GENUS Ourebia Oribi – P. Grubb  Ourebia ourebi Oribi – J. S. Brashares & P. Arcese

405 406

TRIBE REDUNCINI Reduncines – J. Kingdon & A. Gentry

413

GENUS Hippotragus Roan and Sable Antelopes – R. D. Estes & J. Kingdon Hippotragus equinus Roan Antelope – P. Chardonnet & W. Crosmary Hippotragus niger Sable Antelope – R. D. Estes

GENUS Pelea Grey Rhebok – P. Grubb Pelea capreolus Grey Rhebok (Vaal Rhebok, Rhebok) – N. L. Avenant

416

GENUS Redunca Reedbucks – J. Kingdon Redunca fulvorufula Mountain Reedbuck – N. L. Avenant Redunca arundinum Southern Reedbuck – J. Kingdon & M. Hoffmann Redunca redunca Bohor Reedbuck (Common Reedbuck) – J. Kingdon & M. Hoffmann

421 422

GENUS Kobus Kobs – J. Kingdon Kobus kob Kob – F. Fischer Kobus vardonii Puku – R. Jenkins Kobus leche Southern Lechwe – R. Jeffery & R. Nefdt

437 439 445 449

417

426 431

496 502

511

528 533

547 548 556

GENUS Addax Addax – J. Kingdon Addax nasomaculatus Addax – J. Newby

566 566

GENUS Oryx Oryxes – J. Kingdon Oryx gazella Gemsbok (Southern Orxy) – M. Knight Oryx beisa Beisa Oryx (Fringe-eared Oryx) – T. Wacher & J. Kingdon Oryx dammah Scimitar-horned Oryx (Scimitar Oryx) – C. Morrow, R. Molcanova & T. Wacher 

571 572

TRIBE CAPRINI Sheep, Goats – M. Hoffmann & J. Kingdon

593

GENUS Ammotragus Aoudad – P. Grubb & M. Hoffmann

594

576 586

8

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Contents

Ammotragus lervia Aoudad (Barbary Sheep, Arui) – J. Cassinello

595

GENUS Capra Ibexes – P. Grubb & M. Hoffmann Capra nubiana Nubian Ibex – P. U. Alkon Capra walie Walia Ibex (Ethiopian Ibex) – B. Nievergelt

599 600 603

Glossary

607

Bibliography

621

Authors of Volume VI

696

Indexes French names German names English names Scientific names

702 702 703 704

9

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Series Acknowledgements Jonathan Kingdon, David Happold, Thomas Butynski, Michael Hoffmann, Meredith Happold and Jan Kalina

The editors wish to record their thanks to all the authors who have contributed to Mammals of Africa for their expert work and for their patience over the very protracted period that these volumes have taken to materialize. We also thank the numerous reviewers who have read and commented on earlier drafts of this work. We are also grateful for the generosity of our sponsoring patrons, whose names are recorded on our title pages, who have made the publication of these volumes possible. Special thanks are due to Andy Richford, the Publishing Editor at Academic Press, who initiated and supported our work on Mammals of Africa, from its inception up to the point where Bloomsbury Publishing assumed responsibility, and to Nigel Redman (Head of Natural History at Bloomsbury), David and Namrita PriceGoodfellow at D & N Publishing, and the whole production team who have brought this work to fruition. We also acknowledge, with thanks, Elaine Leek who copy-edited every volume. We are grateful to Chuck Crumly, formerly of Academic Press and now the University of California Press, for being our active advocate during difficult times.

above left:

We have benefited from the knowledge and assistance of scholars and staff at numerous museums, universities and other institutions all over the world. More detailed and personal acknowledgements follow from the editors of each volume. The editors are also grateful to the coordinating team of the Global Mammal Assessment, an initiative of the International Union for Conservation of Nature (IUCN), which organized a series of workshops to review the taxonomy and current distribution maps for many species of African mammals. These workshops were hosted by the Zoological Society of London, Disney’s Animal Kingdom, the Owston’s Palm Civet Conservation Programme, and the Wildlife Conservation Research Unit at the University of Oxford; additionally, IUCN conducted a review of the maps for the large mammals by the Specialist Groups of the Species Survival Commission. We owe a particular word of thanks to all the staff and personnel who made these workshops possible, and to the participants who attended and provided their time and expertise to this important initiative. We also thank IUCN for permission to use data from the IUCN Red List of Threatened Species.

photograph by Jan Kalina

Jan Kalina. above: from left to right: Jonathan Kingdon, Thomas Butynski, Meredith Happold, David Happold and Andrew Richford. left: Jonathan Kingdon (left) and Michael Hoffmann.

10

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Acknowledgements for Volume VI Jonathan Kingdon and Michael Hoffmann

The authors and editors of Volume VI would like to express their thanks and gratitude to the following people who kindly offered their time and expertise to help review species or higher-level profiles (or parts thereof), or who made available either published or unpublished information to authors or editors. The contributions, insights, advice and critiques of those listed below greatly improved the content and accuracy of individual profiles. We ask forgiveness from anyone whose name is inadvertently omitted or misspelled. In particular, we owe a significant word of thanks to the late Rod East, former Chair of the IUCN SSC Antelope Specialist Group, who provided immensely useful advice on authors, reviewed several species profiles and provided much useful information. We also remember fondly the late Peter Grubb, whose advice on ungulate classification and taxonomy was indispensable. Sadly, besides Peter, we also lost Keith Eltringham, John Skinner and Vivian Wilson before this volume went to press. We hope that the final volume of Mammals of Africa serves as a monument to their contributions and to those of the many biologists before them. Teresa Abaigar, Justin Akakpo, Philip Alkon, Jeremy Anderson, Markéta Antonínová, Cheri Asa, Nico Avenant, Robert Barnes, Carlo Belfiore, Jean-Renaud Boisserie, J. du P. Bothma, Thomas Breuer, James Brink, Martin Brooks, Tom Butynski, Jorge Cassinello, Philippe Chardonnet, Marcus Clauss, Tim CluttonBrock, Malcolm Coe, Nobby Cordeiro, Terrie Correll, Woody Cotterill, Sue Crocker, David Cumming, Meg Cumming, Fabrice Cuzin, Glyn Davies, Yvonne de Jong, Tom de Maar, Koenraad De Smet, Jean-Pierre d’Huart, Maurizio Dioli, Henk Dop, Werner Dörgeloh, Bob Dowsett, Johan du Toit, Gérard Dubost, Patrick Duncan, Kevin Dunham, Sarah Durant, the late Rod East, Sarah Elkan, Hermann Ellenberg, the late Keith Eltringham, Louise Emmons, Heiner Engel, Dieter Ernst, Lyndon Estes, Richard Estes, Don Farst, François Feer, Julian Fennessy, Frauke Fischer, Charles Foley, Adouma Fragonard, John Fryxell, Nicolas Gaidet, Gerard Galat, Anh Galat-Luong, Annie Gautier-Hion, Valerius Geist, Alan Gentry, Denis Geraads, Hubert Gillet, Morris Gosling, Colin Groves, the late Peter Grubb, Khushal Habibi, Elliot Handrus, David Happold, Rhidian Harrington, John Harris, John Hart, Terese Hart, Ibrahim Hashim, Alexandre Hassanin, Jens-Ove Heckel, Blair Hedges, Pavla Hejcmanová, Philipp Henschel, Dennis Herlocker, Chris Hillman, Reino Hofmann, Jeff Holland, Peter Hoppe, Bernd Hoppe-Dominik, Bill Houston, Reginal Hoyt, Mike Jacobs, Christine Janis, Trevor Jones, Jan Kalina, Erustus Kanga, Samuel Kasiki, Corinne Klaus-Hugi, Hans Klingel, Richard Kock, Orbinda Kok, Petr Komers, Lisa Korte (née Molloy), Karl Kranz, Sally Lahm, Francois Lamarque, Alain Laurent, David Lawson, Kristin

Leus, Barbara Leuthold, Walter Leuthold, Rebecca Lewison, Eline Lorenzen, Gideon Louw, Callie Lynch, Alastair Macdonald, Chris Magin, Florence Magliocca, James Malcolm, David Mallon, Athol Marchant, John Mason, Erik Meijaard, Graham Mitchell, Patricia Moehlman, Renata Molcanova, Steve Monfort, Yoshan Moodley, Michael Mooring, Dominic Moss, David Moyer, Wim Mullié, Souleye Ndiaye, John Newby, Helen Newing, Catherine Ngarachu, Bernhard Nievergelt, David Noble, Peter Novellie, William Oliver, Norman Owen-Smith, Ian Parker, Alexander Peal, Mike Perrin, Norbert Pfannenschmidt, Pierre Pfeffer, Djalle Pierre, Andy Plumptre, Gilfred Powys, Herbert Prins, Melvyn Quan, Wilhelm Räder, Befekadu Refera, Randy Reiches, Paul Reillo, Craig Roberts, Karen Ross, Francesco Rovero, Dave Rowe-Rowe, Ian Rushworth, Mostafa Saleh, Julia Salnicki, Dietrich Schaaf, Cathy Schloeder, Paul Scholte, Susanne Schultz, Erik Seiffert, Russell Seymour, David Shackleton, Steve Shurter, Kirstin Sicx, Claudio Sillero-Zubiri, Alberto Simonetta, Anthony R. E. Sinclair, the late John Skinner, Chris Smeenk, Jan Smielowski, Tommy Smith, Michael Somers, Clive Spinage, Mark Stanley Price, Emma Stokes, Tom Struhsaker, Chris Stuart, Tilde Stuart, Andrew Taylor, Ricky Taylor, the late Simon Thirgood, Kaci Thompson, Chris Thowless, Tom Thurow, Andrea Turkalo, Carlo Utzeri, Herman van Oeveren, Katja Vichl, Petri Viljoen, Fritz Volrath, Wolfgang von Richter, Tim Wacher, Alan Walker, Friedrich Wilhelmi, Stuart Williams, Doug Williamson, the late Vivian Wilson, Achim Winkler, Torsten Wronski, Piet Wit, Johannes Yagos, Derek Yalden, Alberto Zilli and Samuel Zschokke. A special word of thanks to Grant Hopcraft for permission to reproduce the map illustrating the Serengeti Wildebeest migration.

Through the first few formative years of the project, Mike Hoffmann was based in the University of Oxford’s Wildlife Conservation Research Unit. This stay was made possible thanks to the efforts of the Commissioning Editor at Academic Press at the time, Dr Andy Richford, and the Director of WildCRU, Dr David Macdonald. Andy and David made it both financially and academically possible for me to move to the UK to join Jonathan on the Mammals of Africa. WildCRU further afforded access to the wonderful Bodleian library and that of the Edward Grey Institute, as well as to many like-minded African mammal researchers, several of them Mammals of Africa authors (including Andrew Loveridge, Stuart Williams and Claudio Sillero). Claudio became a particularly close and trusted friend, although it is only now that I realize what a novel influence he was. Subsequent to leaving WildCRU, and facilitated by a connection made by Tom 11

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Acknowledgements for Volume VI

Butynski, I took up a position with Conservation International in Washington, DC., before transitioning into IUCN in 2006. I am particularly grateful to Tom Brooks, Russ Mittermeier and Simon Stuart, key supporters throughout my years at CI and IUCN, who made allowances for some of my time to be spent on MoA matters, and who cannot possibly know the profound inspiration that they have had on me personally as a conservationist. That I ever got involved in Mammals of Africa owes a great deal to the late John Skinner, my first mentor (with whom I jointly revised the 3rd edition of the Mammals

00 MOA v6 pp001-021.indd 12

of the Southern African Subregion), for putting in a good word; Dick Estes, for suggesting I get in touch with Jonathan in the first place, and, of course, Jonathan himself, for putting his trust in such a novice as I was then. I look back now on the day Jonathan and I first met on 4 August 2000, and realize how far indeed we both have come. How proud I am to have had the privilege to work so closely with this giant of a man – this legend – and to call him my friend. And, finally, to my parents, for giving me the room to spread my wings, and giving me the courage to do so. To them, I owe everything.

02/11/2012 15:40

Mammals of Africa: An Introduction and Guide David Happold, Michael Hoffmann, Thomas Butynski and Jonathan Kingdon

Mammals of Africa is a series of six volumes that describes, in detail, every extant species of African land mammal that was recognized at the time the profiles were written (Table 1). This is the first time that such an extensive coverage has been attempted; all previous books and field guides have either been regional in coverage, or have described a selection of mammal species – usually the larger species.These volumes demonstrate the diversity of Africa’s mammals, summarize what is known about the distribution, ecology, behaviour and conservation status of each species, and serve as a guide to identification. Africa has changed greatly in recent decades because of increases in human populations, exploitation of natural resources, agricultural development and urban expansion. Throughout the continent, extensive areas of forest have been destroyed and much of the forest that remains is degraded and fragmented. Savanna habitats have been altered by felling of trees and development for agriculture. Many of the drier areas are threatened with desertification. As a result, the abundance and geographic ranges of many species of mammals have declined – some marginally, some catastrophically, some to extinction. Hence, it seems appropriate that our knowledge of each

Table 1.  The mammals of Africa. Order Hyracoidea Proboscidea Sirenia Afrosoricida Macroscelidea Tubulidentata Primates Rodentia Lagomorpha Erinaceomorpha Soricomorpha Chiroptera Carnivora Pholidota Perissodactyla Cetartiodactyla 16 a

Number of families

Number of genera

Number of species

1 1 2 2 1 1 4 15 1 1 1 9 9 1 2 6 57

3 1 2 11 4 1 25 98 5 3 9 49 38 3 3 41 296

5 2 2 24 15 1 93 395a 13 6 150 224 83 4 6 93 1116b

Including five introduced species. b Species profiles in Mammals of Africa.

species is recorded now, on a pan-African basis, because the next few decades will see even more human-induced changes. How such changes will affect each mammalian species is uncertain, but this series of volumes will act as a baseline for assessing future change. The study of African mammals has taken several stages. During the era of European exploration and colonization, the scientific study of African mammals was largely descriptive. Specimens that were sent to museums were described and named. As more specimens became available, and from different parts of the Continent, there was increasing interest in distribution and abundance, and in the ecological and behavioural attributes of species and communities. At first, it was the largest and most easily observed species that were the focus of most studies, but as new methodologies and equipment became available, the smaller and more cryptic and secretive species became better known. Many species were studied because of their suspected role in diseases of humans and livestock, and because they were proven or potential ‘pests’ in agricultural systems. During the past decade or so, there has been greater emphasis on the genetic and molecular characteristics of species. All these studies have produced a wealth of information, especially during the past 40 years or so. These volumes are not only a distillation of the huge literature that now exists on African mammals, but also of much unpublished information. Readers will notice that there is a huge discrepancy among species in the amount of information available. Some species have been studied extensively for many years, especially the so-called ‘game species’, some species of primates and a few species that are widespread and/or easily observed. In contrast, other species are known only by one or a few specimens, and almost nothing is known about them. Likewise, some areas and countries have been well studied, while other areas and countries have been neglected. During the preparation of these volumes, the editors have often been surprised by the wealth of information about some species when little was anticipated, and by the paucity of information about others, some of which were assumed to be ‘well known’. In addition to presenting information that is based on sound scientific evidence, the aims of these volumes are to point out where there are gaps in knowledge and to correct inaccurate information that has become embedded in the literature. For most taxa, the detail provided in the species profiles allows accurate identification. Mammals of Africa comprises six volumes (Table 2). The volumes consist mainly of species profiles – each profile being a detailed 13

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An Introduction and Guide

Table 2.  The six volumes of Mammals of Africa. Volume

Contents

Number of species

Editors

I

Introductory chapters. Afrotheria (Hyraxes, Elephants, Dugong, Manatee, Otter-shrews, Golden-moles, Sengis and Aardvark)

49

II

Primates

93

III

Rodents, Hares and Rabbits Hedgehogs, Shrews and Bats

408

V

Carnivores, Pangolins, Equids and Rhinoceroses

93

VI

Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer and Bovids

93

Jonathan Kingdon, David C. D. Happold, Michael Hoffmann, Thomas M. Butynski, Meredith Happold and Jan Kalina Thomas M. Butynski, Jonathan Kingdon and Jan Kalina David C. D. Happold Meredith Happold and David C. D. Happold Jonathan Kingdon and Michael Hoffmann Jonathan Kingdon and Michael Hoffmann

IV

380

account of the species. They have been edited by six editors who distributed their work according to the orders with which they were most familiar. Each editor chose authors who had extensive knowledge of the species (or higher taxon) and, preferably, had experience with the species in the field. Each volume follows the same general format with respect to arrangement, subheadings and contents. Because Mammals of Africa has contributions from 356 authors (each with a different background and speciality), and each volume was edited by one or more editors (each with a different perspective), it has not been possible or even desirable to ensure exact consistency throughout. Species profiles are not intended to be exhaustive literature reviews, partly for reasons of space. None the less, they are written and edited to be as comprehensive as possible, and to lead the reader to the most important literature for each species. Inevitably, not all information available could be accommodated for the better-known species, and so such profiles are a précis of available knowledge. Extensive references in the text alert the reader to more detailed information. In addition to the species profiles, there are profiles for the higher taxa (genera, families, orders, and above). At the very least, there is a profile for each order, for each family within the order, for each genus within the family, and for each species within the genus. For some orders there are additional taxonomic levels, for example, subfamilies and tribes (Bovinae and Bovini, respectively). Species are presented according to phylogeny. The taxonomy used in these volumes mostly follows that presented in the third edition of Mammal Species of the World: A Geographic and Taxonomic Reference (Wilson & Reeder 2005), although authors have employed alternative taxonomies when there were good reasons for doing so. Volume I differs from the other volumes in that it contains a number of introductory chapters about Africa and its environment, and about African mammals in general.

The continent of Africa For the purposes of this work, ‘Africa’ is defined as the continent of Africa (bounded by the Mediterranean Sea, the Atlantic Ocean, the Indian Ocean, the Red Sea and the Suez Canal) and the islands on the continental shelf that, at some time in their history, have been joined to the African continent. The largest of the ‘continental islands’ are Zanzibar (Unguja), Mafia and Bioko (Fernando Po). All ‘oceanic islands’, e.g. São Tomé, Principe, Annobón (Pagulu), Madagascar, Comoros, Seychelles, Mauritius, Socotra, Canaries, Madeira and Cape Verde are excluded, with the exception of Pemba, which is included because of its close proximity (ca. 50 km) to the mainland. The names of the countries of Africa are taken from the Times Atlas (2005). The Republic of Congo is referred to as ‘Congo’ and the Democratic Republic of Congo (former Zaire) as ‘DR Congo’. Smaller geographical or administrative areas within countries are rarely referred to except for Provinces in South Africa, which are used extensively in the literature. A political map of Africa, and of the Provinces of South Africa, is given (Figure 1), as well as a list of the 47 countries together with their previous names that are used in the older literature on African mammals (Table 3). Africa is the second largest continent in the world (after Asia), but it differs from other continents (except Australia and Antarctica) in being essentially an island. At various times in the past, Africa has been joined to other continents – a situation that has had a strong influence on the fauna and flora of the continent. Africa is a vast continent (29,000,000 km², 11,200,000 mi²) that straddles the Equator, with about two-thirds of its area in the northern hemisphere and one-third in the southern hemisphere. As a result, Africa has many varied climates (with seasons in each hemisphere being 6 months out of phase), many habitats (including deserts, savannas, woodlands, swamps, rivers, lakes, moist forests, monsoon forests, mountains and glaciers), and altitudes ranging from 155 m (509 ft) below sea level at L. Assal, Djibouti, in the Danakil (Afar) Depression, to 5895 m (19,341 ft) on Mt Kilimanjaro, Tanzania. Africa is comprised of 47 countries, some of which are very large (e.g. Sudan [2,506,000 km²; 967,000 mi²], Algeria [2,382,000 km², 920,000 mi²] and Democratic Republic of Congo [2,345,000 km², 905,000 mi²]), and others that are relatively small (e.g. Djibouti [23,200 km², 9000 mi²], Swaziland [17,400 km², 6,700 mi²] and The Gambia [11,300 km², 4400 mi²]). The human population of each country also varies greatly, from about 346/km² in Rwanda to only about 2.5/km² in Namibia. With its great size and varied habitats, Africa supports a high biodiversity, including a large number of species of mammals. Likewise, most countries have a high diversity of mammals (especially when compared with temperate countries). Africa may also be categorized into Biotic Zones (Figure 2). A biotic zone is defined as an area within which there is a similar environment (primarily rainfall and temperature) and vegetation, and which differs in these respects from other Biotic Zones. Africa can be divided into 13 Biotic Zones, two of which may be divided into smaller categories. The Biotic Zones concept provides a general assessment of the environmental conditions in which a species lives, as well as providing an assessment of the geographic distribution of the species. The Rainforest Biotic Zone and the South-West Arid Biotic Zone may be divided into regions and sub-regions that reflect the different biogeographical distributions of species, each region/

14

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The continent of Africa

0°

10°

a

30°

M

c oro

10°

co

20°

Tunisia

30°

30°

Western Sahara

le Ni

Algeria Libya

20°

Egypt 40°

Mauritania

Niger

r Nige

Chad

Burkina Faso

Somaliland Ethiopia

ia

South Sudan

al

a

Cameroon Togo Benin Bioko (Equatorial 0° Guinea) Gabon 0° Rio Muni (Equatorial Guinea) 1000 miles Cabinda (Angola)

Uganda

Congo

Kenya

Co

ng

o

10°

Central African Republic

So

Liberia

10°

an

Côte d’Ivoire

Djibouti

Nigeria

Gh

GuineaGuinea Bissau Sierra Leone

500 1000 km

0°

Pemba Zanzibar

Tanzania

Mafia

10°

10°

Angola

10°

Malawi

Zambia

qu

e

i bez am

bi

Z

Figure 1. (a) Political map of Africa; (b) provinces of South Africa; (c) altitudes and major rivers of Africa. South Sudan and Somaliland are not identified as separate countries in the text.

Zimbabwe

20°

Namibia

am

500

Rwanda Burundi

50°

oz

0

Democratic Republic of Congo

M

0

50°

Eritrea

Sudan

m

Senegal The Gambia 10°

20°

Mali

Botswana

20° 40°

Swaziland

c

30°

30°

South Africa

Lesotho 30°

20°

le Ni Awa sh

W hite Nile

Tana

Za

Shire

e en un

e

Limpopo

Gauteng

North West

a um Ruv Lake Malawi

opo mp Li

Or

b

Free State Northern Cape

Eastern Cape Western Cape 0

ang e

KwaZulu– Natal

zi be m

Lake Kariba Okavango Delta

C

Mpumalanga

Rufiji

Lake Mweru Lake Bangweulu

o ng ba Cu

altitude (metres) 0 1–200 201–500 501–1000 1001–2000 2001–4000 above 4000

Lualaba

ili Kw o ang Kw

1000 miles

1000 km

i Lomam Sankuru Kasai

é

oou

500 500

a

a Og

0 0

o She bel Om u l Mbomo Lake Uele Albert Lake Turkana Congo Aruwimi-Ituri Mt Elgon Rwenzori Mtns Mt Kenya Lake Lake Tshuap a Edward Victoria Lukenie Mt Kilimanjaro Galana Lake Tanganyika Jub

Sangh

Cross

e nu Be Mt Cameroon aga San Ivindo

Lu an gw a

Lake Volta

Lake Tana

Ouban gui

Black Volta

olta ite V Wh

Lake Chad

ile eN Blu

gal

e Sen r Nige

0

300 miles 300 km

15

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An Introduction and Guide

Table 3.  The countries of Africa: names, areas and human population density. Country name Algeria Angola (includes Cabinda) Benin * [Dahomey] Botswana [Bechuanaland] Burkina Faso * [Upper Volta; Burkina] Burundi [part of Ruanda-Urundi (= part of Belgian Congo)] Cameroon [includes former French Cameroon, German Cameroon and part of Eastern Nigeria] Central African Republic # Chad [Tchad] Congo [Republic of Congo] Côte d’Ivoire * [Ivory Coast] Democratic Republic of Congo [Belgian Congo; Congo (Kinshasha); Zaire] Djibouti [French Somaliland] Egypt Equatorial Guinea # (includes Rio Muni [Spanish Guinea] and Bioko I. [Fernando Po]) Eritrea (formerly part of Ethiopia) Ethiopia [Abyssinia] Gabon # The Gambia Ghana [Gold Coast] Guinea * Guinea-Bissau [Portuguese Guinea] Kenya Lesotho [Basutoland] Liberia Libya Malawi [Nyasaland] Mali * Mauritania * Morocco [includes former Spanish Morocco and French Morocco]; (now also includes Western Sahara = former Spanish Sahara) Mozambique [Portuguese East Africa] Namibia [South-west Africa] Niger * Nigeria Rwanda [part of Ruanda-Urundi (= part of Belgian Congo)] Senegal * Sierra Leone Somalia ¥ [British Somaliland and Italian Somaliland; Somali Republic] South Africa Sudan § [Anglo-Egyptian Sudan] Swaziland Tanzania [German East Africa; Tanganyika] (now includes Zanzibar I., Mafia I. and Pemba I.) Togo [Togoland] Tunisia Uganda Zambia [Northern Rhodesia] Zimbabwe [Southern Rhodesia] Totals/mean density

Area (km2) ’000

Area (miles2) ’000

Human population ’000 (2006)

People per km2

2,382 1,247 113 582 274 27.8 475

920.0 481.0 43.0 225.0 106.0 10.7 184.0

33,500 15,800 8,700 1,800 13,600 7,800 17,300

14.1 12.7 77.0 3.1 49.6 280.5 36.2

623 1,284 342 322 2,345

241.0 496.0 132.0 125.0 905.0

4,300 10,000 3,700 19,700 62,700

6.9 5.8 10.8 61.2 26.7

23.2 1,001 28.1

9.0 387.0 10.8

800 75,400 500

34.5 75.3 17.8

94 1,128 268 11.3 239 246 36 580 30.4 111 1,760 118 1,240 1,030 447

36.0 436.0 103.0 4.4 92.0 95.0 13.9 224.0 11.7 43.0 679.0 46.0 479.0 412.0 172.0

4,600 74,800 1,400 1,500 22,600 9,800 1,400 34,700 1,800 3,400 5,900 12,800 13,900 3,200 32,100

48.9 66.3 5.2 132.7 94.6 39.8 38.9 59.8 59.2 30.6 3.6 108.5 11.2 3.1 71.8

802 825 1,267 924 26.3 197 71.7 638 1,220 2,506 17.4 945

309.0 318.0 489.0 357.0 10.2 76.0 27.7 246.0 471.0 967.0 6.7 365.0

19,900 2,100 14,400 134,500 9,100 11,900 5,700 8,900 47,300 41,200 1,100 37,900

24.8 2.5 11.3 145.6 346.0 60.4 79.5 13.9 38.7 16.4 63.2 40.1

56.8 164 236 753 391 29,448

21.9 63.0 91.0 291.0 151.0 11,383

6,300 10,100 27,700 11,900 13,100 902,600

110.9 61.6 117.4 15.8 33.5 56.8

Former names are listed in chronological order in square brackets, with the oldest name listed first. Obsolete names are listed because much of the older literature refers to past colonial entities. * = formerly part of French West Africa. # = formerly part of French Equatorial Africa. § At the time of going to press, the country of Sudan had been divided into two: the Republic of Sudan in the north, and the Republic of South Sudan in the south. ¥ The former British Somaliland is now a self-declared state under the name of the Republic of Somaliland, but remains internationally unrecognized.

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The Cetartiodactyla of Africa

The pigs, hippopotamuses, chevrotain, giraffes, deer and bovids of Africa

1

This volume,Volume VI, is devoted to the order Cetartiodactyla, a large order that unites the traditional ‘artiodactyls’ (even-toed ungulates) with the whales and dolphins. The order includes three main subdivisions: Suiformes (pigs); Whippomorpha (hippopotamuses, whales and dolphins); and Ruminantia (including the chevrotain, giraffe and okapi, deer and the bovids). In total, the order includes 98 species (treated in 93 species profiles), although the majority comprise the antelopes, which have radiated into a remarkable 76 species in 12 different tribes. A single species of African cetartiodactyl, the Bluebuck Hippotragus leucophaeus, is documented as having become extinct since 1500 (though many are teetering on the brink) and is discussed briefly under the genus account, but is otherwise not treated separately. Introduced species are mentioned in the higher-level profiles (where relevant), but are not otherwise profiled. At the time of going to press, a new species of duiker in the genus Philantomba had been described, Philantomba walteri (Colyn et al. 2010). Unfortunately, it was not possible to include an individual species profile, but we provide general characteristics that distinguish this new species under the genus profile. Details on the type locality description follow (see also the Philantomba genus profile).

2

3 6a 5 6 1 = Mediterranean Coastal Biotic Zone 2 = Sahara Arid Biotic Zone 3 = Sahel Savanna Biotic Zone 4 = Sudan Savanna Biotic Zone 5 = Guinea Savanna Biotic Zone 6 = Rainforest Biotic Zone    6a = Northern Rainforest–Savanna Mosaic    6b = Eastern Rainforest–Savanna Mosaic    6c = Southern Rainforest–Savanna Mosaic 7 = Afromontane–Afroalpine Biotic Zone (discontinuous, shaded brown) 8 = Somalia–Masai Bushland Biotic Zone 9 = Zambezian Woodland Biotic Zone 10 = Coastal Forest Mosaic Biotic Zone 11 = South-West Arid Biotic Zone    11a Kalahari Desert    11b Namib Desert    11c Karoo 12 = Highveld Biotic Zone 13 = South-West Cape Biotic Zone

4

7

5 6a

8 6

6b

6c 10 9

11a 11b

12 11c 13

Figure 2. The biotic zones of Africa. The numbers refer to the biotic zones as described in the text.

sub-region having a community of mammals and other animals which is different to any other. Details of the Biotic Zones of Africa, and the regions and sub-regions of the Rainforest Biotic Zone and SouthWest Arid Biotic Zone, are given in Volume I of Mammals of Africa.

crown nape

forehead

Walter’s Duiker Philantomba walteri Colyn, Hulselmans, Sonet, Oudé, de Winter, Natta, Nagy & Verheyen, 2010. Igbere (08° 59' N 01° 57' E), Forêt Protégée de Wari-Maro (328 m altitude), near the Ecological Center of Manigri, Benin. As Mammals of Africa was being finalized, a new work by Colin Groves and the late Peter Grubb, Ungulate Taxonomy, was published but it has not been possible to fully consider and evaluate the conclusions and classification presented in that work.

withers

rump

muzzle back

neck

tail base

nostrils lips

cheek chin

shoulder

throat

buttock

flanks hindquarter

belly

dewlap elbow

upper hindleg

upper foreleg hock knee lower hindleg

Figure 3. External features of a mammal: Common Eland Tragelaphus oryx.

pastern

lower foreleg fetlock

fetlock

pastern

hoof

17

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An Introduction and Guide

Figure 4. External features of a mammal: Genetta sp. (side-view and face frontal).

head

neck

body mid-dorsal line

tail

DORSAL

back (dorsal pelage)

external ear (pinna) crown

rump

forehead

face

flank buttock

neck

eye muzzle

cheek throat

nostrils lips

forehead

ventral surface ventral pelage

crown

basal end of tail thigh

VENTRAL

chin

tuft

chest shoulder

tail

POSTERIOR

distal end

tip of ear forelimb (or foreleg) (upper and lower) base of ear

temple

ear, pinna digits (1, 2, 3, 4, 5)

digit(s)

muzzle

hindlimb (or hindleg) (upper and lower)

forefoot

eye nostrils (nose) lips

pelage (= fur) hair (= single hair(s))

Species profiles Information about each species is given under a series of subheadings. The amount of information under each of these subheadings varies greatly between species; where no information is available, this is recorded as ‘No information available’ or words to this effect. The sequence of subheadings is as follows: Scientific name (genus and species)  The currently accepted name of the species. Vernacular names  English, French and German names are given, as available. The first given English name is the preferred vernacular name for the species; alternative names are given in parentheses for some species. Wilson & Cole (2000) list proposed vernacular names for all the world’s mammals; most of these names were also given in the third edition of Mammal Species of the World (Wilson & Reeder 2005). Although these works have been consulted, the names used have not always been adopted in Mammals of Africa. French and German names were usually provided by authors. Scientific Citation  This provides the full scientific name of the species, i.e. genus name, species name, authority name, and date of authority. Parentheses around the authority’s name and date indicate that the species was originally named in a different genus to its present generic allocation. The scientific name is followed by the publication where the species was described, and the location where the type specimen (or type series) was obtained. Most of this information is taken from Wilson & Reeder (2005).

Taxonomy  This section contains information on taxonomic problems, if any, associated with the species, and its relationship with other species in the genus. For some species, there is considerable information about these topics; for others, there may be nothing. A list of synonyms (without the taxonomic authority for each) and the number of subspecies (if any) is presented, mostly taken from Wilson & Reeder (2005). The chromosome number is given if available, and in some cases this is followed by other information relevant to the chromosomes. Description  This section, together with the illustrations, provides the reader with adequate information to identify the species. The section begins with a brief overall description of the species, including an indication of size. This is followed by a detailed description of the external features of the species’ head (and parts of the head), dorsal pelage, legs, feet, ventral pelage, and tail (in this order), as well as any special characteristics unique to the species. For some species, diagnostic characteristics of the skull and dentition are given, and for bovids the horns. This section may also present information on ageing criteria, where this has been investigated with some degree of rigour. The characters described in this section are common to all subspecies of this species (unless otherwise noted). The mammary formula, i.e. the number and arrangement of nipples in adult females, is noted wherever this feature varies between the taxa being discussed. Geographic Variation  Variation within the species may be of two sorts: (a) clinal variation without subspecies, or (b) subspecific variation. If (a), there is a description of the character(s) that alter clinally across the geographic range of the species. If (b), each of the subspecies is listed with its geographic range and, where available,

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Species profiles

the characters that distinguish it from other subspecies of the species. For some species, subspecies have been described that are no longer considered to be valid; in some cases, such names may be listed but without further comment. Similar Species  Species that are sympatric or parapatric with the species under consideration, and with which it may be confused, are listed along with diagnostic characteristics (additionally, readers may refer to profiles of the similar species in question). In some instances, species that are allopatric in distribution are also included. Distribution  The first sentence ‘Endemic to Africa’ informs the reader that this is an African species and does not occur on any other continent; if a species also occurs outside Africa, this is noted at the end of this section with a very brief synopsis of the extralimital range. The distributions of the cetartiodactyls are among the best studied and well documented of African mammals, and the monumental work of Rod East (1999), complemented by subsequent updates from the IUCN SSC Antelope Specialist Group, provides an unrivalled platform for detailing the ranges of most species. For widespread species that generally remain so today (e.g. Sylvicapra grimmia), the text provides a general idea of the range, highlighting only places or countries where the species may have been extirpated, recently newly recorded, or providing clarity on previously incorrectly attributed country records and range limits. In the case of widespread species that have undergone significant range contractions and declines (e.g. Addax nasomaculatus), the text generally differentiates between former and current ranges in an attempt to elucidate a clearer picture of where species do, or no longer, occur. Finally, for more rangerestricted species (e.g. Beatragus hunteri), the information provided may be quite precise, detailing even localized distribution within the confines of its small range. For species with recognized and well-differentiated subspecies, we have attempted to present the details of the distributions accordingly, and within the framework presented above. Although great effort has been expended in ensuring this section is current and provides a true picture of the current range of the species, many data gaps remain, especially for parts of North and West Africa, Sudan, Somalia, parts of DR Congo, Angola and Mozambique. A distribution map (see below) augments the information given here. Habitat  This section provides a description of the range of habitats where the species lives. Details of plant communities, plant species, vegetation structure, soil type and/or structure, and water availability, etc. (if available) are also recorded. Other information may include average annual rainfall, altitudinal limits, and seasonal variation in habitat characteristics. Abundance  A general indication of abundance in the habitat. This may be unquantified, such as abundant, common, uncommon, rare, or phrases such as ‘rarely seen but frequently heard’, etc. For better-known species, abundance may be expressed as estimates of density (e.g. number/ha or number/km2) as well as in terms of actual numbers of individuals for the species. Other information may include seasonal changes in density, frequency of observations, or the relative abundance of specimens in collections.

Adaptations  This section describes morphological, physiological and behavioural characteristics, which show how the species uniquely interacts with its environment, conspecifics and other animals. This section may also describe species-specific adaptations for feeding, locomotion, burrowing, mechanisms for orientation, production of sound, sensory mechanisms and activity patterns. In some instances comparison with related or convergent species allows the unique adaptations of the species under discussion to be detailed or highlighted. Foraging and Food  This section provides information on the diet and foraging habits of the species. The diet is described either by a list of the taxa of animals or plants consumed, or as a quantitative measure of the contents of the stomach or the faeces. This section can also include any of the following: location of food, foraging behaviour, times when foraging occurs and daily distance moved; hoarding; seasonal changes in diet and food availability; individual or cooperative behaviour used in foraging and hunting; sex and age differences in foraging and diet; and nomadic or migratory movements in relation to food availability. Social and Reproductive Behaviour  Topics in this section may include group structure (whether solitary, social, or colonial), group size and composition; agonistic and amicable behaviour, comfort behaviour, etc.; home-range (including quantitative data), territorial behaviour, courtship and mating behaviour, behaviour of young, parental–young interactions; presence of helpers, vocalizations, and interactions with other species (mammals, birds, etc.). Reproduction and Population Structure  This section begins with an assessment of reproductive strategy (if known) and the times/seasons of the year when individuals are reproductively active (pregnancy and lactation in females, active spermatogenesis in males). Other information may include length of gestation, times/ seasons of births, including peaks of births, litter-size, birth-weight and size, spacing of litters, growth and time to weaning, maturity, longevity, and mortality rates. Reproductive strategies, if known, are described with respect to locality, food availability and population density. Population structure (sex ratio, adult/young ratio, abundance of different cohorts in the population at different times of the year) may be described, and related to seasonal variations in reproduction and environmental variables. Predators, Parasites and Diseases  The known predators, parasites and diseases are listed. Information on parasites and diseases is not intended to be exhaustive, but simply to provide an entry point into the literature on the topic. In some cases, information on diseases from captive animals is presented. Additional information is given if the species is a host to diseases that affect humans and domestic stock. Conservation  The conservation status of the species is stated, as given by the IUCN Red List of Threatened Species (version 2011.2). The IUCN Red List categories follow the definitions and criteria given in the IUCN Red List Categories and Criteria Version 3.1 (Table 4). For those species classified as threatened (i.e. ‘Vulnerable’, 19

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An Introduction and Guide

Table 4.  IUCN Red List Categories (from IUCN – International Union for Conservation of Nature). Category

Description

Extinct (EX)

A taxon is Extinct when there is no reasonable doubt that the last individual has died. A taxon is presumed Extinct when exhaustive surveys in known and/or expected habitat, at appropriate times (diurnal, seasonal, annual), throughout its historic range have failed to record an individual. Surveys should be over a time frame appropriate to the taxon’s life-cycles and life-form. A taxon is Extinct in the Wild when it is known only to survive in cultivation, in captivity or as a naturalized population (or populations) well outside the past range. A taxon is presumed Extinct in the Wild when exhaustive surveys in known and/ or expected habitat, at appropriate times (diurnal, seasonal, annual), throughout its historic range have failed to record an individual. Surveys should be over a time frame appropriate to the taxon’s life-cycle and life-form. A taxon is Critically Endangered when the best available evidence indicates that it meets any of the criteria A to E for Critically Endangered, and it is therefore considered to be facing an extremely high risk of extinction in the wild. A taxon is Endangered when the best available evidence indicates that it meets any of the criteria A to E for Endangered, and it is therefore considered to be facing a very high risk of extinction in the wild. A taxon is Vulnerable when the best available evidence indicates that it meets any of the criteria A to E for Vulnerable, and it is therefore considered to be facing a high risk of extinction in the wild. A taxon is Near Threatened when it has been evaluated against the criteria but does not qualify for Critically Endangered, Endangered or Vulnerable now, but is close to qualifying for (or is likely to qualify for) a threatened category in the near future. A taxon is Least Concern when it has been evaluated against the criteria and does not qualify for the Critically Endangered, Endangered, Vulnerable or Near Threatened categories. Widespread and abundant taxa are included in this category. A taxon is Data Deficient when there is inadequate information to make a direct, or indirect, assessment of its risk of extinction based on its distribution and/or population status. Data Deficient is not a category of threat. Listing of taxa in this category indicates that more information is required and acknowledges the possibility that future research will show that a threatened classification is appropriate. A taxon is Not Evaluated when it has not yet been evaluated against the criteria.

Extinct in the Wild (EW)

Critically Endangered (CR) Endangered (EN) Vulnerable (VU) Near Threatened (NT) Least Concern (LC) Data Deficient (DD)

Not Evaluated (NE)

‘Endangered’, ‘Critically Endangered’), the criteria met are also indicated. Some species have changed status due to improved knowledge, taxonomic revision, or the impact of threatening processes or conservation actions. Readers can obtain detailed reasons for the past and present status of a species by going to the IUCN Red List website (www.iucnredlist.org). If a species was listed on Appendix I, II or III under CITES (Convention on International Trade in Endangered Species – www.cites.org – as of 22 December 2011) or Appendix I or II of CMS (Convention on Migratory Species – www.cms.int – as of 5 March 2009) this is also indicated. For some species, additional information is provided, such as presence in protected areas, major threats, and current or recommended conservation measures. Measurements  A series of morphological measurements is provided. For each species there is a standard set of measurements. The abbreviations for each measurement are given in the Glossary. A measurement is cited as the mean value (with minimum value to maximum value in parentheses), and sample size. For some, the standard deviation (mean ± 1 S.D.) is given instead of the range. Where possible, data for males and females are presented separately. In some cases, more than a single set of measurements is given; this is particularly the case for widespread species where geographic variation in size may be evident, and also for species with several well differentiated subspecies (in which case, we have endeavoured to present a set of measurements for each). Some species have additional stand-alone measurements presented beneath the primary series. Skull measurements generally are not provided; however, in the case of the antelopes, maximum recorded horn length (and the location of this record) based on the Rowland Ward (27th edition) measuring system is indicated. The majority of measurements also contain the location(s) where the specimens were obtained, and the source of the

data. Sources are either cited publications, or specimens in museums, or unpublished information from authors or others.The acronyms for museums where specimens were examined and measured are given in Table 5. Key References  A select list of references, which provides more general information on the species, or is generally considered as a key reference work on the species. Each reference is given in full in the Bibliography. Citations given in the text (but not cited in ‘Key References’) are also given in full in the Bibliography. In general, profiles account for all literature published up until the end of 2007. Authors and editors have endeavoured to keep the species profiles up-to-date throughout the long production schedule, and references published from 2008 onwards have been incorporated wherever possible. None the less, certain key recent papers will have been missed or omitted. Author  The name of the author, or authors, is given at the end of each profile. All profiles should be cited using the author name(s). Tables  With one or two exceptions, the use of tables to present data has been avoided in this volume of Mammals of Africa.

Table 5.  Museum acronyms. Acronym

Museum name

AMNH BMNH

American Museum of Natural History, New York, USA Natural History Museum, London, UK [formerly British Museum (Natural History)] Musée Royal de l’Afrique Centrale, Tervuren, Belgium National Museums of Kenya, Nairobi, Kenya Powell-Cotton Museum, Birchington,UK

MRAC NMK PCM

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Editors of Mammals of Africa

Higher taxon profiles The profiles for orders, families and genera are internationally much less structured than for the species profiles. Each profile usually begins with a listing of the taxa in the next lower taxon; for example, each family profile lists the genera in that family. An exception to this arrangement is where a taxon has only one lower taxon. Higher taxa profiles provide the characteristics common to all members of that taxon. Some of these characteristics (for example, number of nipples or dental formula) may not be repeated in lower taxon profiles (unless essential for identification), so readers are encouraged to consult also the higher-taxa profiles, e.g. the species profile for Raphicerus campestris should be consulted in association with the genus Raphicerus and the tribe Raphicerini profile.

Distribution maps Each species profile contains a pan-African map showing the geographic range of the species. In the case of the antelopes, the maps draw heavily on the publications of East (1988–1990, 1999), subsequent ASG updates and recent survey reports (e.g. Fay et al. 2007 for southern Sudan). The purpose of the maps is to show current known limits of distribution of the species within historical range, recognizing that within this mapped range a particular species’ distribution will not be homogeneous. In general, reintroductions within the former range of a species are included and mapped (unless otherwise indicated), but introductions outside of the former known range are not. Even applying these rules, mapping current distributions in some parts of the continent, and especially in southern Africa where the game ranching industry has resulted in major faunistic shufflings, has proved very complicated. After much consideration, it was decided not to map historical ranges for species, given the difficulty in mapping these with any level of accuracy for a number of species, differing concepts on what ‘historical’ means, and the availability of such maps in other sources (e.g. Sidney 1965, Du Plessis 1969). However, in the absence of more recent and reliable information for some countries (especially Angola, Mozambique, Somalia and so forth), the range maps may actually reflect historical distributions far better than they do current range. Subspecies are

only indicated in cases where the boundaries can be reasonably delineated. Each map shows the boundaries of the 47 countries of Africa, some of the major rivers (Nile, Niger–Benue, Congo [with the tributaries Ubangi, Lualaba and Lomani], Zambezi and Orange), and Lakes Chad, Tana, Turkana (formerly Rudolf), Albert, Edward, Victoria, Kyoga, Kivu, Tanganyika, Malawi, Mweru, Bangwuela and Kariba. The map projection is Transverse Mercator, with the following parameters: False Easting: 0; False Northing: 0; Central Meridian: 20; Linear Unit: metre; Datum: Clarke 1866. The geographic distribution of a species is indicated as: • red shading = current range. Different colour shading denotes subspecies, where appropriate. • × = isolated locations considered to be separate from the main geographic range(s); some locations indicated by × may include two or more closely spaced locations. • ? = uncertain, but possible, presence. • red arrow = recorded from the island indicated by the arrow.

Editors of Mammals of Africa Jonathan Kingdon, Department of Zoology, University of Oxford, WildCRU, Tubney House, Abingdon Road, Tubney OX13 5QL, UK. (Vols I, II, V & VI) David Happold, Research School of Biology, Australian National University, Canberra, ACT 0200, Australia (Vols I, III & IV) Thomas Butynski, Eastern Africa Primate Diversity and Conservation Program, PO Box 149, Nanyuki 10400, Kenya, and Zoological Society of London, King Khalid Wildlife Research Centre, Saudi Wildlife Authority, PO Box 61681, Riyadh 11575, Kingdom of Saudi Arabia (Vols I & II) Michael Hoffmann, International Union for Conservation of Nature – Species Survival Commission, 219c Huntingdon Road, Cambridge CB3 0DL, UK. (Vols I, V & VI) Meredith Happold, Research School of Biology, Australian National University, Canberra, ACT 0200, Australia (Vols I & IV) Jan Kalina, Soita Nyiro Conservancy, PO Box 149, Nanyuki 10400, Kenya (Vols I & II)

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Order CETARTIODACTYLA

Order CETARTIODACTYLA – Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer, Bovids Cetartiodactyla Montgelard, Catzeflis & Douzery, 1997. Mol. Biol. Evol. 14: 550. Suidae (4 genera, 6 species) Hippopotamidae (2 genera, 2 species) Tragulidae (1 genus, 1 species) Giraffidae (2 genera, 2 species) Cervidae (1 genus, 1 species) Bovidae Bovinae (2 genera, 10 species) Antilopinae (29 genera, 76 species*)

Pigs, Hogs Hippopotamuses Chevrotains Giraffe, Okapi Deer

p. 25 p. 63 p. 87 p. 95 p. 116

Bovines Antilopines

p. 122 p. 199

*including species in the Nanger (granti) and Madoqua (kirkii) species groups.

Cetartiodactyla, of which 98 terrestrial species occur in Africa, are commonly referred to as the ‘even-toed ungulates’ or Artiodactyla.This placental order contains pigs (suborder Suiformes), camelids (suborder Tylopoda; extralimital to Africa), ruminants (suborder Ruminantia) and the hippopotamuses (hereafter hippos), the latter of which now comprise the suborder Whippomorpha together with the whales and dolphins.The distinctness as well as the relatedness of each of these taxa is now well supported by molecular and genomic data. The etymology of this new taxon therefore combines the two traditional orders ‘Artiodactyla’ and ‘Cetacea’ into an enlarged order ‘Cetartiodactyla’. Traditional taxonomy, based on morphology, placed whales in their own order, Cetacea, but over the course of the last decade a steady accumulation of nucleotide sequence data and other genomic information has made it clear that whales are actually deeply nested within the ‘artiodactyl’ radiation as the sister-group of Hippopotamidae (e.g. Gatesy et al. 1996, Nikaido et al. 1999, Murphy et al. 2001, Amrine-Madsen et al. 2003, Arnason et al. 2004, Meredith et al. 2011, Hassanin et al. 2012). This hypothesis has since come to be supported by phylogenetic analyses of morphological data from living and extinct cetartiodactyls (Geisler & Uhen 2003, 2005), as well as combined molecular-morphological analyses of the order (Spaulding et al. 2009). It has been argued that the taxon Artiodactyla should be retained for this assemblage (Archibald 2003, Asher & Helgen 2010), but this practice threatens to engender considerable confusion among specialists, because some workers have used the taxa Cetartiodactyla and Artiodactyla in order to express their recognition of a sister-taxon relationship between a monophyletic Cetacea and a monophyletic Artiodactyla (e.g. Thewissen et al. 2001). Relationships among the major cetartiodactyl clades are a matter of ongoing debate, and estimates of cetartiodactyl phylogeny have generated an extensive literature (Gatesy et al. 1999, 2002, Nikaido et al. 1999, Matthee et al. 2001, Murphy et al. 2001, Arnason et al. 2004, Geisler & Uhen 2005, Spaulding et al. 2009, Meredith et al. 2011, Hassanin et al. 2012). One of the first responses to the molecular evidence supporting the placement of whales within Artiodactyla was to re-examine artiodactyls and cetaceans, both extant and fossil, to seek morphological evidence for or against this unexpected affinity (Luckett & Hong 1998). In spite of continued scepticism, living and extinct cetartiodactyls can now be recognized on the basis of at least three distinctive postcranial

Astragalus of extinct hippopotamus (from Kingdon 1982).

and dental features: (1) the head of the astragalus is trochlear, leading to the characteristic ‘double-pulley’ shape; (2) manus and pes are paraxonic, which means that the weight-bearing axis passes through digits III and IV (metapodials III and IV have become elongate and thickened, whereas metapodials II and V have been reduced in length and diameter, and digit I has been greatly reduced or lost entirely); and (3) the deciduous fourth premolar (p4) is distinctly trilobed, and the anterior lobe occludes between the outer (paracone and metacone) cusps of the upper deciduous third premolar (p3). The molariform p4 is later replaced by a premolariform p4. Until recently, the astragalar and pedal morphology of primitive cetaceans was not known; however, it is now clear that the archaeocetes Pakicetus, Ichthyolestes, Rodhocetus and Artioclavus all had double-pulley astragali (Gingerich et al. 2001, Thewissen et al. 2001), and Rodhocetus had a paraxonic arrangement of the pedal digits (Gingerich et al. 2001). Molecular estimates for the origin of crown Cetartiodactyla extend back to near the K–T (Cretaceous–Tertiary) boundary (Springer et al. 2003, Meredith et al. 2011), but the group is not known in the fossil record before the earliest Eocene, around 55 mya.The most generalized cetartiodactyl is arguably Diacodexis (sometimes called ‘Gujaratia’) pakistanensis, from the early Eocene of Pakistan (Thewissen & Hussain 1990, Geisler & Uhen 2005). Diacodexis appears to have migrated out of Asia into Europe and North America during a brief period of intense global warming at the Palaeocene–Eocene boundary, and thereafter primitive cetartiodactyls (many of which are now of unclear phylogenetic position) are known from all three northern continents, and with time become common members of Laurasian mammal faunas. Recognizable stem or crown members of the Ruminantia, Tylopoda and Suiformes do not appear until much later in the Palaeogene, but all are ultimately of Laurasian origin.The presumably semi-aquatic anthracotheriid artiodactyls first appear in Africa in the late Eocene (Holroyd et al. 1996); otherwise, fully terrestrial cetartiodactyls are not known in Africa before the early Miocene. Details of taxonomy and biology among the very diverse cetartiodactyls are discussed under appropriate taxonomic profiles,

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Order CETARTIODACTYLA

but it is difficult to overstate the revolution in biological thinking that has accompanied these molecular discoveries.While earlier scenarios had, for example, suggested that whale ancestors were terrestrial carnivores, more recent discoveries make larger, more omnivorous, 100

90

80

70

60

50

40

30

20

10

0 mya

Suiformes Camelidae Hippopotamidae Cetacea Tragulidae Giraffidae Cervidae Bovidae Perissodactyla

and perhaps superficically pig-like ancestors more likely (Thewissen et al. 2007). Reconstructing the earliest history of whale evolution now presents new challenges and an added impetus for scientists to find still earlier proto-whale fossils. Among the challenges will be the plausible reconstruction of sequences and successions of feeding styles between ancient terrestrial artiodactyls and today’s fish, squid and krill-eating whales. Interpolation of purely herbivorous hippopotami into the whales’ ancestral tree has brought many new dimensions to study of the Cetartiodactyla, but in spite of the challenges and new uncertainties, one thing remains unchanged, and Dawkins (1996) put it this way: ‘there is no doubt that whales and dugongs come with their dry-land history written all over them. If they had been deliberately created for the sea, they’d be very different, and a lot more like fish than they are. Animals that have their history written all over them are among the most graphic pieces of evidence we have that living things were not created for their present ways of life but evolved from very different ancestors’ (p. 133). Modern genetics has forced us to examine hippos with more open minds and eyes. When we get to re-examine other Cetartiodactyla with similar intensity we will find that antelopes, giraffes and others will also have their history written all over them, even if in subtler phrases.

Tentative phylogenetic tree for Cetartiodactyla (after Bininda-Emonds et al. 2007 and Hassanin et al. 2012).

Suidae

Camelidae

Hippopotamidae

Tragulidae

Erik R. Seiffert & Jonathan Kingdon

Giraffidae

Bovidae

Sacculation in cetartiodactyl stomachs (from Kingdon 1982).

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Suborder SUIFORMES

Suborder SUIFORMES – Pigs Suiformes Jaekel, 1911. Die wirbeltiere: eine ubersicht uber die fossilen und lebenden formen, Berlin. pp. 233.

Traditionally, the Suiformes have been viewed as a clade that included not only the African pigs and hippos, and their extant and extinct Eurasian relatives, but also forms with no known living counterparts, such as the Anoplotheriidae from Europe, the Raoellidae from Asia, the Choeropotamidae from Eurasia and the Oreodontoidea from North America, plus the Dichobunoidea, Anthracotheroidea and Entelodontoidea that are known from both Eurasia and North America (McKenna & Bell 1997). Recent advances in our understanding at the molecular level of artiodactyl evolution, together with new and complete Palaeogene whale fossils and reinterpretation of previously documented fossil evidence, have led to the recognition of the order Cetartiodactyla in which ancodonts (anthracotheres and hippos) are now grouped with cetaceans (whales and dolphins) in the suborder Whippomorpha, whereas pigs and peccaries are retained in the suborder Suiformes. The Suiformes are today represented in Africa by four genera of the family Suidae, which is in turn grouped with the extralimital peccaries (Tayassuidae) and extinct sanitheres (Sanitheriidae) into the superfamily Suoidea. The Suoidea are united by the uniquely derived features (autapomorphies) of: lack of a lingual cingulum on the upper molars; the similarity of the lower first and second incisors; a rootless lower male canine; paraconid fused to metaconid in the lower cheekteeth; tympanic process of squamosal dorsoventrally elongate and external auditory meatus opening dorsally; and by the presence of ossified tympanic bullae (Liu 2003).

The sister relationship between Suidae and Tayassuidae is well documented on both morphological (Gentry & Hooker 1988) and molecular grounds (Irwin & Arnason 1994, Randi et al. 1996). Pigs and peccaries occupy similar adaptive zones (sensu Simpson 1944) in the Old and New World, respectively. Both are primarily omnivorous and of the living artiodactyls display the most primitive traits – retaining four distinct digits, separated foot bones, absence of frontal appendages, a simple, non-ruminating stomach, and the less progressive forms having low-crowned cheekteeth with simple bunodont cusps. Sanithere cheekteeth are bunoselenodont, with a tendency to further complicate the premolars by polycuspy and polycristy accompanied by heavy wrinkling of occlusal enamel. In some dental and cranial features the sanitheres are convergent on selenodont artiodactyls, but the astragalus is characteristically suoid (Pickford 1984). The oldest suoid fossils now known are from the late Eocene of China (Tong & Zhao 1986, Liu 2001) and Thailand (Ducrocq 1994, Ducrocq et al. 1998). Although the evidence is still sparse and incomplete, it would appear that suoids originated in eastern Asia during the Eocene, subsequently dispersing into the New World (Tayassuidae) and elsewhere in the Old World (Suidae) during the Oligocene (Ducrocq 1994, Ducrocq et al. 1998, Liu 2001). John M. Harris

Superfamily SUOIDEA PIGS Suoidea Gray, 1821. London Med. Repos. 15: 306.

The superfamily Suoidea describes a taxon that subsequently bifurcates into the New World peccaries (family Tayassuidae) and the Old World pigs of the family Suidae. John M. Harris

Diversity of morphology and gait in contemporary suids. Comparison of gaits in two suid species: upper row: Bushpig Potamochoerus larvatus ‘gallop’. Lower row: Common Warthog Phacochoerus africanus ‘fast trot’ (from film, courtesy of D. Cumming).

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Family SUIDAE

Family SUIDAE PIGS, HOGS Suidae Gray, 1821. London Med. Repos. 15: 306. Sus (1 species) Potamochoerus (2 species) Hylochoerus (1 species) Phacochoerus (2 species)

Wild Boar Bushpig, Red River Hog Forest Hog Warthogs

p. 28 p. 31 p. 40 p. 50

Sus scrofa Potamochoerus porcus Potamochoerus larvatus Hylochoerus meinertzhageni Phacochoerus aethiopicus Phacochoerus africanus

Phylogenetic tree of extant African suidae (modified after White & Harris 1977 and Hassanin et al. 2012).

The family Suidae contains the pigs and their relatives. Suids are robust animals with relatively short legs and large heads. The modern forms have a short braincase and an exceedingly elongated facial skeleton forming a tubular snout. Peculiarly characteristic of suids among African ungulates is the terminal position of the nostrils, surrounded by a partly cartilaginous, mobile disc attached to a central prenasal bone; they have leathery, sometimes sparsely haired skin. All pigs have simple stomachs, lacking the fermentation chambers of ruminants. Suidae are separated from other suoids by the absence of the angular process of the mandible (Liu 2003). Suid molars are also characterized by the presence of three furrows or ‘furchen’ on each of the four main cusps (Hünermann 1968, Pickford 1986), a feature that is lacking in the bunodont peccaries and bunoselenodont sanitheres. This furrowed enamel is thrown into subsidiary cuspules surrounding the main cusps. Suids have developed a diastema between the canine ‘tusks’ and the masticatory toothrow with progressive reduction of the premolars in grass-eating species. This trend can be

correlated with improved milling of tough vegetation and finds its most extreme expression in the warthogs Phacochoerus spp. where mature individuals only retain M3. The mesial premolars are small and unicuspid, but the premolar and molar series gets progressively more complex distally, with the third molars capable of elongation with three or more pairs of cusps. The upper incisors are small and spaced, the lowers somewhat procumbent. The upper canines are small and pointed in the !, large and curved outward and upward in the ", and form potent weapons. The Suidae have been divided into several subfamilies. The Listriodontinae are characteristic of the early to middle Miocene (21–14 mya) in Eurasia and Africa; early forms had low-crowned bunodont teeth but more advanced representatives developed tapir-like lophodont molars. The Kubanochoerinae had a similar temporal and geographic range to the listriodontines; they retained low-crowned bunodont teeth, but some species developed horns on the frontal and above the orbits. Tetraconodontines were most abundant in the late Miocene and Pliocene (9–4 mya) of Africa and Asia and developed very large third and fourth premolars. The Namachoerinae were precociously lophodont, short-snouted suids from the early to mid-Miocene of Africa. Cainochoerines were pygmy forms from the late Miocene of eastern and southern Africa. The bunodont Palaeochoerus of Europe (Oligocene to early Miocene, 34–22 mya) was at one time interpreted as an ‘Old World peccary’, but Liu (2003) reinterpreted it as part of an unresolved stem group of the Suoidea. The lophodont Schizotheriinae of the African and Eurasian Miocene were also interpreted as ‘Old World peccaries’, but Liu (2003) included them in the Suidae. The extant suids are sometimes interpreted as representing three subfamilies: Suinae (widely distributed in the Old World), Phacochoerinae (sub-Saharan Africa) and Babyrousinae (eastern Asia), but their similarities to each other, and differences from extinct suid subfamilies, probably only warrants separation at the tribal level (which is the approach followed herein). Colin Groves & John M. Harris

Subfamily SUINAE – Pigs, Hogs Suinae Gray, 1821. London Med. Repos. 15: 306.

The first representatives of the Suinae appeared towards the end of the middle Miocene in Europe and Asia. Suinae crania are distinguished by an elongated flange – the prezygomatic shelf – that projects anteriorly from the zygomatic root. This flange separates the chewing musculature from the muscles that operate the snout. It was not present in the hyotheriines, kubanochoeres, listriodonts and tetraconodonts, where the anterior tendinal guides for the snout musculature in the vicinity of the canines were generally poorly developed. This suggested to Pickford (1993) that the characteristic

rooting habit of extant pigs is restricted to the Suinae. Another progressive feature of suines is the development of sagittal cusplets between the labial and lingual cusps of the upper fourth premolar. The Suinae radiated rapidly and are represented by some 15 genera during and after the late Miocene.The heterogeneous nature of suine molars could permit these pigs to be treated as a subfamily, which could be further subdivided into several component tribes, but there is currently little consensus of opinion as to tribal compositions. However, there is general recognition that the hypsodont African 25

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suines (Phacochoerus and the extinct Metridiochoerus) form a natural grouping. A major faunal turnover occurred toward the end of the Miocene (7–5.3 mya) in sub-Saharan Africa, reflecting the opening up of forested habitats and the spread of C4 grasses (Leakey et al. 1996, Cerling et al. 1997b). The listriodonts, kubanochoeres and sanitheres characteristic of the African early and middle Miocene were replaced by tetraconodonts in the late Miocene and these were in turn replaced by suines that arrived during the late Pliocene (3–1.8 mya). The contrast in size between the relatively small early and middle Miocene African suids and their larger late Miocene and Pliocene counterparts reflects the more open habitats available to the later forms. The Suinae are first documented in Africa at about 3.4 mya. Kolpochoerus, Potamochoerus and Metridiochoerus represent the ancestral stocks of the extant forest hogs (Hylochoerus), bushpigs (Potamochoerus) and warthogs (Phacochoerus), respectively (Cooke 1978a, Harris & White 1979). Potamochoerus remained virtually unchanged during the ensuing 3-plus million years, but some of the Kolpochoerus and Metridiochoerus species underwent increase in size and in complexity of the cheekteeth. Later members of both the Kolpochoerus and

0

10 km

Principal range of Phacochoerus africanus Principal range of Potamocherus larvatus

Metridiochoerus lineages achieved gigantic size but Hylochoerus was probably derived from the moderately sized K. heseloni, whereas Phacochoerus probably originated from the diminutive M. modestus (Harris & White 1979). Today, the African suids live in wooded or thicket country, from rainforest and montane forest to savannas and semi-arid scrub, which provides requisite shelter. In some rare localities the ranges of all three genera overlap, but their ecological preferences and spatial partitioning are clear, with bushpigs preferring forest and thicket formations, warthogs in wooded savannas and the Forest Hog in forest/grassland mosaics. All species are capable of expanding their ecological range in the absence of the other species.They typically live in groups of one or more !! (sows) with young; adult "" (boars) are predominantly solitary. Females in most species make nests, to which they constantly return with their young; warthogs occupy burrows. Litters typically number 2–8 and single births are rare. The very large size achieved by some extinct Pliocene and Pleistocene African suids was probably a predator-selected adaptation for living in more open habitats. John M. Harris

ABOVE: Old adult skulls of (above left) Bushpig Potamochoerus lavatus and (above right) Common Warthog Phacochoerus africanus compared with juveniles showing succession and differential migration of teeth during life (from Kingdon 1979). LEFT: Map of habitat use by three suid species in and around Budongo Forest, Uganda: Common Warthog Phacochoerus africanus; Bushpig Potamocherus larvatus; Forest Hog Hylochoerus meinertzhageni.

Principal range of Hylochoerus meinertzhageni

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Tribe SUINI Old World Pigs Suini Gray, 1821. London Med. Repos. 15: 306.

Heel

M3 Hylochoerus

Heel

M3 Potamochoerus

ABOVE: Diagrams of upper third molar in (left) Forest Hog Hylochoerus meinertzhageni and (right) Bushpig Potamochoerus larvatus. LEFT: Three suid skulls showing flexion of snout in relation to cranial capsule: a) Bushpig Potamochoerus larvatus; b) Common Warthog Phacochoerus africanus (down); c) Forest Hog Hylochoerus meinertzhageni (upward).

Africa is home to three extant suin genera. The Wild Boar Sus scrofa was at one time restricted to North Africa, but feral stock is now also known from western and southern parts of the continent. The Forest Hog Hylochoerus meinertzhageni is essentially limited to the equatorial forest belt, but the bushpigs (Potamochoerus spp.) are more widely distributed in the less arid parts of Africa. Bushpigs are water-dependent opportunistic feeders that will take advantage of any available food source; Warthogs, in contrast, feed exclusively on C3 plants and are less water-dependent (Harris & Cerling 2002). All three genera were originally bunodont browsers or omnivores, but derived Pleistocene species of the extinct African suin Kolpochoerus had high-crowned teeth and a grazing diet (Harris & Cerling 2002). In common with some other pigs, the braincases of Suini are enclosed in bony capsules that are separate from the skull’s outer casing. One benefit of this separation between the cranium, with its associated basicranial axis, and the skull’s outer shell is that the latter can be damaged (and often is during violent male fights) without compromising brain function. Another attribute is very ready modification of the outer shape of pig skulls during evolution.

The Suini evidently originated in western Asia (Pickford 1993), from whence they migrated to eastern Asia, Europe and Africa. Kolpochoerus deheinzelini, known by teeth from the latest Miocene of Chad (Brunet & White 2001), is the oldest suin yet documented in the African record. The cranium of the Pliocene K. afarensis was rather similar to that of bushpigs but with downwardly drooping zygomatic arches and less prominent canine flanges (Cooke 1978b); in "" of later species, the zygomatic arches were pneumatized. Harris & White (1979) postulated that Hylochoerus was derived from Kolpochoerus during the early Pleistocene and that the scarcity of fossil Hylochoerus specimens is an artefact of their preferred habitat. Although a few teeth attributable to Potamochoerus have been recovered from late Pliocene localities in East Africa (Harris & White 1979), fossil bushpigs are similarly elusive and for the same reason. Sus appears to have been a post-Pleistocene immigrant to North Africa. John M. Harris

(Left) ‘Snout-boxing’ in Bushpig Potamochoerus larvatus; (middle) ‘forehead-clashing’ in Forest Hog Hylochoerus meinertzhageni; (right) ‘tusk-clashing’ in Common Warthog Phacochoerus africanus.

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GENUS Sus Wild Boar Sus Linnaeus, 1758. Syst. Nat., 10th edn, 1: 49.

The genus Sus includes ten extant species (Grubb 2005), of which only one, the Wild Boar Sus scrofa, occurs in North Africa. However, this species has been introduced elsewhere on the continent. Of the extant African suins, the ecologically equivalent Wild Boar and Bushpig Potamochoerus larvatus are of similar size and share many characteristics, including small upper tusks, three pairs of upper incisors, and relatively unspecialized, low-crowned bunodont cheekteeth, but Sus has a full complement of premolars and its enamel is thinner. The skulls of both Sus and Potamochoerus are elongate and pointed (or wedge-shaped) with an almost straight profile from the tips of the nasals to the parietals. The braincase is gently domed behind the orbits with a narrow parietal constriction. The snout of Sus is long and narrow with parallel-sided nasals, whereas in Potamochoerus the nasals widen above the canine flanges and in "" there is an expanded rugose area on the nasals and the adjoining parts of the maxillae. Sus has only small maxillary flanges around the roots of the upper canines, even in "", whereas in Potamochoerus the canine flanges are strongly developed and rugose in their upper portions. The zygomatic arches of Sus are narrow and not expanded laterally whereas in Potamochoerus the maxillary root of the zygoma juts out sharply and may be quite inflated in "". Sus lacks the distinct small shelf on the back of the mandibular symphysis for

the attachment of the genioglossus and geniohyoideus muscles that are present in bushpigs. The earliest (mid-Miocene) representatives of the genus Sus conceivably derive from Asian Propotamochoerus stock. By the end of the Miocene the genus was widespread throughout Eurasia. In Europe, Sus was the dominant suid genus from the Pliocene onwards; the small S. arvenensis characterized the early Pliocene, but overlapped with the very large S. strozzi in the later Pliocene.The extant species S. scrofa appeared in the late early Pleistocene. In Asia, Sus competed with other suine genera during the Pliocene and later diversified into a number of different species whose extant representatives are collectively known as the warty pigs (Groves & Grubb 1993). Curiously, Sus was not part of the suine radiation that brought Potamochoerus, Kolpochoerus and Metridiochoerus to sub-Saharan Africa during the early Pliocene and the Wild Boar was a relatively late immigrant to the continent. Cooke & Wilkinson (1978) indicated that remains of S. scrofa were present at many later Pleistocene sites in North Africa and the species is also documented at the mid-Pleistocene localities of Ternifine (Pomel 1896) and Mansoura (Joleaud 1933) in Algeria. The genus was not found south of the Sahara until introduced there through human agency. John M. Harris

Sus scrofa WILD BOAR (EURASIAN WILD PIG) Fr. Sanglier; Ger. Wildschwein Sus scrofa Linnaeus, 1758. Syst. Nat., 10th edn, 1: 49. ‘Habitat in Europa australiore’; shown to be Germany, from where wild boar had been introduced to Sweden, Oeland (Thomas 1911: 140).

Taxonomy Wild Boar populations show considerable variation in body size, cranial morphology and coat colour. Groves (1981a) recognized 16 subspecies, which can be assembled into four regional

groups (Groves & Grubb 1993): the Western subspecies (S. s. scrofa and S. s. meridonalis, distributed in Europe; S. s. algira, in North Africa; S. s. lybicus, in south-west Asia, and extinct in the Nile Delta,

Wild Boar Sus scrofa.

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Wild Boar Sus scrofa. Sus scrofa

though the origin of this population is obscure; and S. s. attila and S. s. nigripes in Central Asia); the Indian subspecies (S. s. davidi, in the sub-Himalayan region; S. s. cristatus, in NC India, Myanmar and W Thailand; S. s. affinis and a new, undescribed subspecies, in S India and Sri Lanka); the Eastern subspecies (S. s. sibiricus, in Mongolia; S. s. ussuricus, in eastern Asia; S. s. leucomystax and riukiuanus, in Japan; S. s. taivanus, in Taiwan; S. s. moupinensis, in S China and Vietnam); and the Indonesian subspecies S. s. vittatus, which is distributed in the Malay Peninsula, Sumatra, Java, Bali). All extant North African Wild Boars belong to S. s. algira; barbarus, the name sometimes used for this subspecies, is a nomen nudum. In Tunisia, molecular data reveal a clear break between northern and southern populations, possibly due to an Algerian origin of the southern animals (Hajji & Zachos 2011). Synonyms: Grubb (2005) lists numerous synonyms, but only algira, barbarus and sahariensis pertain to Africa. Three variations of chromosome number have been recorded in wild populations, with 2n = 36, 37 and 38, due to Robertsonian translocations (Bosma 1976, Mauget et al. 1977, Popescu et al. 1980, Jotterand-Bellomo & Baetting 1981, Mayr et al. 1984, McFee et al. 1986, Arroyo-Nombela et al. 1990). The chromosome number of Wild Boars in North Africa is 2n = 38 (El Mastour et al. 1983), but this requires confirmation as the origin of the two animals analysed (from Tunis Zoo) is uncertain. Description A grey, brownish pig of intermediate body size, with relatively short muzzle and no facial warts. The head profile is triangular. Ears rather large, not tufted. Snout elongated, the tip of which is flat and forms a cartilaginous rhinarium, which is used in digging. Limbs relatively long. Hair colouration agouti, in general greyish, blackish or brown; there may be whitish hairs on the face, cheeks and throat. Hairs are often dense, and the bristles on the dorsal line are long, sometimes forming a crest or a mane. Piglets longitudinally striped brownish-red during the first five to six months. Tail has tufts. Females are smaller and about 28% lighter than "" (El Mastour et al. 1983, Abáigar 1990). Females have four to six pairs of nipples.

The dental formula is I 3/3, C 1/1, P 4/4, M 3/3 = 44. Large upper canines are present, the length of which depend on the age, with their alveoli directed outward, curved outwards, backwards and upwards. Geographic Variation As noted under Taxonomy above, there is considerable geographical variation in colour and morphology, and many subspecies have been recognized, which Corbet (1978) states have never been shown to have discrete boundaries. Similar Species The Wild Boar cannot be confused with any other suid species within its range in Africa. Distribution The Wild Boar is widely distributed in the Palaearctic region and shows the largest range among all the suiforms. On the African continent, the species occurs naturally only in North Africa, then extralimitally formerly in Europe (including the British Isles), through S Russia, the Middle East and South Asia to the Malay Peninsula and the Greater Sundas. They are also present on Japan and Taiwan. The Wild Boar was eradicated in the British Isles and also in southern Scandinavia and N Japan in the seventeenth century. Isolated populations still exist on Honshu, and on Sardinia and Corsica (the only Mediterranean islands to which they are native), but their range also has been significantly expanded through reintroductions and introductions. Local populations in western and central Europe have been intensively restocked with allocthonous or hybrid animals for hunting purposes. The species has been introduced in the USA, Australia, New Zealand, the Lesser Sundas,West Indies, and numerous oceanic islands, including the Galapagos and Hawaii (Ellerman & Morrison-Scott 1951, Corbet 1978, Oliver et al. 1993, Grubb 2005). On the African continent, domestic pigs were released into plantations in the Western Cape of South Africa in an attempt to control the Pine Tree Emperor Moth Nudaurelia cytherea (Thomas & Kolbe 1942), but their invasive potential has been minimal when 29

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(Cuzin 2003). In Algeria, they range up to 2300 m in Djurdjura (K. De Smet pers. comm.). Usually needs water for drinking, but may live without any water in summer, for example, in Djebel Guettar (Algerian hauts plateaux) (Heim de Balsac 1936). Abundance In Morocco, Wild Boars seem to attain their highest densities in dense forest, as they do in south-eastern Andalucia, Spain (Abáigar et al. 1994). In Algeria and Tunisia, numbers are rising everywhere and the species is recolonizing parts of its former range, as in the Bou Hedma (K. De Smet pers. comm.). Adaptations Mainly crepuscular and nocturnal, but active also during the day in undisturbed areas (Benhamza 1995). In C Morocco, there are two peaks of activity at night, between 21:00 and 01:00h and between 03:00 and 07:00h (El Mastour et al. 1983). Animals tend to stay in the densest available vegetation during the day. Adults wallow in mud, where available.

Lateral, palatal and dorsal views of skull of Wild Boar Sus scrofa.

compared with other parts of the world (Botha 1989); they also have been introduced to several other African countries (e.g. Gabon, Sudan, Burkina Faso; Vercammen et al. 1993). Historical Distribution Formerly in North Africa (north of 26° N) from Morocco’s lower Seguia El Hamra (Morales Agacino 1950) through N Algeria, Tunisia and N Libya to the Nile R. in N Egypt (Osborn & Helmy 1980). The origin of Egyptian animals is questionable, as they probably represent indefinite crossings between wild and domesticated animals (Manlius & Gautier 1999). Animals from Sudan are also probably of domesticated origin (Ansell 1972, Manlius & Gautier 1999), as are animals from the offshore islands of Mafia and Pemba (Haltenorth & Diller 1980). Fossil remains of the Wild Boar are common in Pleistocene and Holocene layers in north-west Africa, and the animal is also depicted on Roman mosaics in ancient Libya (Kowalski & Rzebik-Kowalska 1991). Current Distribution In North Africa, confined to Maghreb (north of 27° N) from Morocco (south-western limit south of the lower Draa R.) to Tunisia. Probably no longer occurs in Libya (where the last observation dates from 1883), and extinct in Egypt since about 1902 (Hufnagl 1972, Osborn & Helmy 1980). Habitat Inhabits a variety of well-vegetated environments, from forests to dense steppes, including marshes and riparian vegetation, in areas with over 150 mm annual rainfall.Wild Boars are able to inhabit very steep slopes, and usually are more abundant on northern rainy slopes and in gullies (Cuzin 2003). Under drier climate (between 50 and 150 mm annual rainfall), in northern Sahara, confined to humid environments (oases, marshes and riparian habitats, especially in Tamarix thickets). In Morocco, they occur from sea level up to 2750 m in the High Atlas, though more common between 1500 and 2250 m

Foraging and Food Much of what is known about diet in Africa originates from studies in C and N Morocco (El Mastour et al. 1983) and N Algeria (Klaa 1992). Wild Boars are mainly herbivorous, the major part of the diet consisting of aerial and subterranean parts of plants. Leaves, stalks and fruits (Quercus spp., Chamaerops humilis, Ceratonia siliqua, Rhus pentaphylla) are consumed. In SE Andalucia, during autumn, acorns of oaks Quercus ilex may constitute the main part of the diet (Abáigar 1993). Wild Boars intensively dig for roots and bulbs, especially those of Arisarum vulgare, when the soil is not too hard, mainly after the rains and under forest cover with well-developed or humid soils; this activity is very limited in dry areas, where soil is usually too hard (Cuzin 2003). A minor, but common, part of the diet consists of insects, annelids, myriapods, snails and other invertebrates, and, occasionally, some meat, eggs, small mammals (Panouse 1957) and larger animals, probably as carrion. Strong canines allow them to open carcasses very efficiently, thereby allowing other small scavengers to feed (K. De Smet pers. comm.). In SE Andalucia, young animals consume more animal matter (Abáigar 1993). There is some evidence of seasonal variation in diet, with leaves and stalks mainly consumed after rains, usually in winter and spring; fruits are consumed from autumn to spring, and digging activity is more frequent when the soil is not too hard. In Algeria, consumption of animal food seems more frequent in autumn. In the driest habitats, in the absence of rain, animals may move to more productive habitats, e.g. oases (F. Cuzin pers. obs.), where they eat a lot of dates (K. De Smet pers. comm.). After periods of high rainfall in the N Moroccan Sahara, animals moved up to 110 km from their native area, returning there when the habitat dried out (F. Cuzin pers. obs.). Wild Boars are attracted to agricultural fields, especially cereals, maize, wheat and barley. Social and Reproductive Behaviour Wild Boars are gregarious, occurring in groups of up to 17 animals (Benhamza 1995). In Europe, groups of generally related !! with piglets, and subadult !! are the most common social unit, while "" stay isolated or in small groups, joining female groups only during the rutting season; territorial behaviour is evident only for resting places (Gerard et al. 1991). There are no details of home-range size in North Africa. In S France, home-ranges are very variable according to the season:

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Maillard & Fournier (1994) recorded an average of 263 ha for groups and 1063 ha for lone !! during summer, and 497–15,440 ha for "" and 1292–3698 ha for !! during autumn. Dispersal is more common among !! than "". Wild Boars have a wide range of vocalizations, from rhythmic grunting of "" when leading groups to teeth-chattering when angry.

belief, Wild Boars are a pest in agricultural fields and are sometimes hunted by local people (Monteil 1951, F. Cuzin pers. obs.).

Measurements Sus scrofa HB (!!): 1471 (1300–1600) mm, n = 8 HB (""): 1310 (1200–1430) mm, n = 8 Reproduction and Population Structure Mating usually WT (!!): 94.5 (64.0–110.0) kg, n = 8 occurs from Oct until Jul (thereby avoiding the driest season), with WT (""): 63.8 (42.0–94.0) kg, n = 8 parturition from Feb until Oct, with a peak in Mar (El Mastour N and C Morocco (El Mastour et al. 1983); animals older than three et al. 1983). Gestation is 112–126 days in Europe (Gerard et al. years 1991). Mean litter-size is 5.7 (range 2–11; n = 13), according to foetal counts, and 5.1 (range 3–8) based on direct observations (El HB (!): 1200 mm, n = 1 Mastour et al. 1983). Time to weaning is not recorded for North HB (""): 1160, 1200 mm, n = 2 Africa, but in Europe, "" stay alone from the end of gestation T (!): 210 mm, n = 1 until 1–5 weeks after birth (Gerard et al. 1991). Young are mature T (""): 165, 190 mm, n = 2 at 7–12 months of age. Longevity and population structure is E (!): 110 mm, n = 1 unknown in North Africa. E (""): 105, 110 mm, n = 2 HF c.u. (""): 240, 250 mm, n = 2 Predators, Parasites and Diseases Leopards Panthera pardus N Morocco (Cabrera 1932) have been observed feeding on Wild Boar (P. C. Beaubrun & J. Godart pers. comm.), although this species is on the verge of extinction in Key References Cuzin 2003; El Mastour et al. 1983. North Africa (Cuzin 1996). There are also reports of Golden Jackals Canis aureus preying on young (Khidas 1986).There is no information Fabrice Cuzin & Ettore Randi available on their susceptibility to disease or parasites in North Africa. Conservation IUCN Category: Least Concern. CITES: Not listed. The primary threat to the Wild Boar in North Africa is forest loss and habitat degradation, coupled by a drying of humid habitats, especially on the northern fringes of the Sahara (Cuzin 2003). Nonetheless, Wild Boars are expanding their range in some parts (Cuzin 1996), and in Tunisia and Algeria they are gradually expanding southwards, especially in the wake of the Islamic uproar in the Tell Atlas, and a ban on hunting (Kowalski & Rezebik-Kowalska 1991, K. De Smet pers. comm.). Though considered as impure by Islamic Wild Boar Sus scrofa male with area of dermal shield indicated.

GENUS Potamochoerus Bushpig, Red River Hog Potamochoerus Gray, 1854. Proc. Zool. Soc. Lond. 1852: 129 [1854].

This genus is represented by two very distinct species (Grubb 1993b): the monotypic Red River Hog P. porcus, characteristic of the rainforest belt but extending north of it into savanna country; and the geographically varying Bushpig P. larvatus, of heavy cover in the East and southern African savanna zone. The Bushpig was introduced in precolonial times to Madagascar (also the type locality for the species), where it flourishes; the reasons for its introduction seem obscure, but presumably relate to meat supply. Potamochoerus is the most plesiomorphic of the three endemic Afrotropical genera of Suidae, and is presumed, on morphological grounds, to be the sister genus of Sus, with which it shares the following character states: the rostrum is extremely elongated; there is a deep preorbital fossa; and the temporal ridges are strongly

developed.The canines of the !, which are comparatively small, flare sideways and curve upward; those of the " are much smaller, but likewise protrude sideways. The root of the maxillary canine sits in a large bony flange, the canine apophysis, which protrudes from the anterior maxilla lateral to the rostral end of the preorbital fossa. Also in common with Sus is the presence of three upper incisors. The premolars, contrary to the two other endemic Afrotropical genera, are not only well developed, but are enlarged and somewhat molarized. The adult dental formula is: I 3/3, C 1/1, P 3–4/3–4, M 3/3 = 40–44. In contrast to Sus, however, pigs of this genus are characterized by several important differences: the braincase bulges outward below the temporal ridges; the occiput is lower; the auditory canal, instead of ascending at 45 degrees or more, is relatively horizontal; and the 31

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nasals are dorsally flattened, and laterally expanded, so that they overhang the lateral walls of the rostrum.The canine apophysis of the " is greatly enlarged and roughened, and rises up toward the shelflike lateral nasal edge. The anterior premolar is always absent in the mandible, commonly so in the maxilla (see dental formula above). The other premolars are strongly molarized, with flattened, cuspidate occlusal surfaces (in Sus, they have narrow blades). The mandibular ascending ramus slopes more posteriorly than in Sus, and the condyle is relatively lower. Ewer (1958) described the functional anatomy of

the facial musculature, relating it to the mobility of the snout, especially its terminal disc. Externally, this genus of mainly nocturnal omnivorous pigs is characterized by the long, pointed ears; and, among the Afrotropical genera, by the longitudinal ridges along the top of the snout and forehead; by the retention of a long, bristly pelage; and by the lack of prominent facial warts, but the presence of a flat wart on either side of the snout supported by the enlarged canine apophyses. Colin Groves

Potamochoerus larvatus BUSHPIG Fr. Potamochère; Ger. Buschschwein Potamochoerus larvatus (F. Cuvier, 1822). Mem. Mus. Hist. Nat. Paris 8: 447. Madagascar.

Taxonomy The Bushpig was originally considered conspecific with the Red River Hog Potamochoerus porcus, which is now regarded as a separate allopatric species of West and central Africa (Grubb 1993b, 2005; and see De Beaux 1924). Grubb (1993b) noted that too many subspecies have been recognized in the past, and commented that the primary systematic division within the species is between the white-faced animals of East Africa and populations in southern Africa. He tentatively recognized three subspecies on the African mainland (Potamochoerus larvatus hassama, P. l. somaliensis and P. l. koiropotamus) and an additional two on Madagascar and Comoros Is. (P. l. larvatus and P. l. hova). However, Vercammen et al. (1993) proposed that insufficient evidence exists for the recognition of P. l. somaliensis as a distinct subspecies (indeed, Grubb 1993b was not convinced of the validity of somaliensis either), and recognized only two from the continent: the White-faced Bushpig P. l. hassama (including P. l. somaliensis; East Africa) and the Southern Bushpig P. l. koiropotamus (from Angola and southern Africa). The Madagascan and Comoron subspecies of the Bushpig were almost certainly introduced during historic times. Synonyms: africanus, arrhenii, choeropotamus, congicus, cottoni, daemonis, edwardsi, hassama, hova, intermedius, johnstoni, keniae, koiropotamus, madagascariensis, mashona, nyasae, somaliensis. Chromosome number: not known, but presumably as for P. porcus. Hybrids with introduced feral pigs Sus scrofa have been recorded (Milstein 1971, Smithers 1983). Description Medium-sized, thickset suid. The head is characterized by an elongated muzzle, with naked disc-like snout, cheek beards and terminal ear tassels. The morphology of the head differs between the sexes and the slightly heavier male body mass is related to these differences. Males develop three pairs of wartlike facial structures: over the canine root flanges on the snout (preorbital); on the malar eminences (infraorbital); and on the jaw angles (gonial). Ears pointed with tufts of long hair at tips. Pelage bristly, extending from the head over the whole body, with a striking dorsal crest extending from the head to the base of the tail; hairs long (60 mm), but relatively sparse. Undercoat absent. General colour variable within and between subspecies and individuals, and changes with age. Neonatal pelage of pale, yellowish-buff stripes on a brown ground colour is replaced by a rufous-brown at about three months of age, gradually changing to the colouration of maturity. Adult

Bushpig Potamochoerus larvatus.

body colour is predominantly black to dark grey, but, particularly in north-eastern parts of the range, may be light red to brown. Males often have a distinct white facial mask. Tail with sagittally aligned (feathered) black terminal hairs. Males have tusk, orbital (Harderian), digital and preputial glands; orbital and digital glands are found in !!. Females have three pairs of nipples. The highest point of the skull lies near the back and then slopes forward to the nostrils. Rostrum narrow and elongated, hollowed out on either side to accommodate the powerful muscles and tendons that open and close the rhinarium during rooting and digging. Mandible long and massive, its actuation relying on the powerful masseters provided with firm attachment between the thick zygomatic arches and the angle of the lower jaw. Supraoccipital ridges pronounced.The back of the skull arches high from the occipital condyles and provides a firm base for the attachment of the neck muscles, which actuate the movement of the heavy head. Adult dentition is developed at about two years, M3 being the last to erupt. The incisors are heavily built and persist throughout the life of the individual. The upper canines project outside the mouth cavity but are never as long as in warthogs Phacochoerus spp.; the lower canines are sharp-pointed and sharpedged, lying at an angle sloping outwards from the mandible, which

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Bushpig Potamochoerus larvatus juvenile skeleton.

Lateral, palatal and dorsal views of skull of Bushpig Potamochoerus larvatus.

makes them formidable weapons. The molars are brachyodont and bunodont, and in old individuals still retain some ridging indicating a chopping action rather than grinding during mastication (Seydack 1983, Skinner & Chimimba 2005). Geographic Variation P. l. hassama (including P. l. somaliensis) (White-faced Bushpig): East Africa, including Ethiopia, Eritrea, S Sudan (both west and east of the Nile R.), E DR Congo, Rwanda, Burundi, Uganda, Kenya and N Tanzania. Characterized by the white colour of the head, usually black body in adult "" and smaller average size, as indicated by skull length (32.7–35.3 cm in !! and 34.1–37.7 cm in ""; Grubb 1993b). P. l. koiropotamus (Southern Bushpig). SE DR Congo, S Tanzania, Malawi, Mozambique, Zambia, W and C Angola, Zimbabwe, Botswana, South Africa and Swaziland. Largest member of the genus (skull length 36.7–41.5 cm in "" and 34.5–39.5 cm in !!), typically never with contrasting black-and-white colour pattern of P. l. hassama, but "" from an isolated population in the Western Cape and Eastern Cape also generally have white faces contrasting with darker bodies. It is also possible that darkerbodied Bushpig "" have whitish facial patterns, whilst lighterbodied ones have darker facial markings for contrast. Similar Species Potamochoerus porcus. Allopatric, though ranges are contiguous in places. Predominant coat colour bright russet-orange with a white dorsal line; pelage over much of body short and dense; terminal ear tassels particularly elongated; forehead black or dark brown, with prominent white ‘spectacles’ around eyes. Hylochoerus meinertzhageni. Sympatric in some parts of eastern Africa. Larger, more heavily built suid, with long black hair, and a dorsal crest of thick, erectile hair; " has enormous bare cheeks and a broad, flat muzzle.

Bushpig Potamochoerus larvatus myology of juvenile.

Bushpig Potamochoerus larvatus juvenile.

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Abundance Mentis (1970) reported 0.22 individuals/km2 in Hluhluwe-iMfolozi G. R., KwaZulu–Natal, South Africa. Population density estimates during a study in southern Cape forests, South Africa, range between 0.3 and 0.5/km2 (Seydack 1990, 1991).

Potamochoerus larvatus

Distribution Endemic to Africa, extending from S Sudan, Ethiopia and possibly Eritrea in the north-east of the continent, southwards through East Africa and E and S DR Congo to Zambia, Malawi and Mozambique. There are isolated populations in W and C Angola. In DR Congo, the vicinity of the upper reaches of northward-flowing tributaries of the Congo R. are occupied by this species, whereas the continuous lowland forest that surrounds more northerly reaches of the same rivers is inhabited by Red River Hogs. In southern Africa, they are absent from Namibia, though they may occur in the Caprivi Strip. In Botswana they occur only in the northern and north-eastern parts, while in Zimbabwe they occur nearly throughout. In South Africa they are found in the northern and eastern parts of the country (and neighbouring Swaziland) to S KwaZulu–Natal. There is a break in their distribution between S KwaZulu–Natal and an isolated population in the Eastern Cape and Western Cape Provinces (which is also the origin of the holotype). Recorded from Zanzibar (Pakenham 1984) and Mafia I. (Kingdon 1979, Kock & Stanley 2009), and introduced on Madagascar and the Comoro Is. (Mayotte) off the east coast of Africa (Vercammen et al. 1993). Habitat Bushpigs are associated with relatively dense vegetation types with available food, cover and water, occurring in forests and riverine or xeric scrub forests and thicket formations. In East Africa they commonly are found in high-lying areas, such as the Ethiopian Highlands to elevations of 3000 m, and perhaps 3400 m (they are common in the Harenna Forest;Yalden et al. 1996), the highlands of the Albertine Rift (where Red River Hogs have never been recorded), and on Mt Kilimanjaro (to 4000 m; Vercammen et al. 1993). They also maintain their occurrence in areas now given over to subsistence farming or to agricultural crops such as sugarcane, maize, peanuts and beans, and have long been a serious problem in such areas.

Adaptations Bushpigs obtain the largest proportion of their food from the litter layer through superficial rooting. The compact body and thick short neck allow for the necessary digging forces to be applied. The build of the skull with its long snout, forwardly sloping lower incisors, short canines and bony support for the rhinarium (discussed above) are all adaptations for searching for food by rooting and digging, while their bunodont dentition and monogastric digestive system make the Bushpig well suited to an omnivorous diet. Olfaction plays a prominent role in locating food items. The thermoneutral zone is between 13 and 30 °C for juveniles and between 8 and 25 °C for adults (Seydack 1990). Accordingly, Bushpigs avoid temperature extremes, being active preferentially between 18:00 and 22:00h in both summer and winter. In summer, Bushpigs rested mainly between 10:00 and 18:00h and in winter between 10:00 and 14:00h and 2:00 and 6:00h (Seydack 1990). Thus, although primarily nocturnal, more diurnal activity may occur during winter. Behavioural thermoregulation includes wallowing, lying in extended posture and exposure to air movement at high temperatures and resting after midnight in winter, sheltering, huddling and nest-building during cold and wet spells. Foraging and Food Bushpigs are omnivorous generalists, feeding opportunistically, but primarily on subterranean and above-ground herbaceous plant material. The diet of Bushpigs in the Knysna Forest of theWestern Cape and the valley bushveld regions of the Eastern Cape (South Africa) comprised 40% subterranean plant structures (of which 54.3% consisted of rhizomes, 38.9% tubers, 5.7% roots and 1.1% corms), 30% above-ground herbage (of which 46% consisted of monocotyledons, 38.7% fern fronds and 12.3% dicotyledons), 13% fruit, 9% animal matter and 8% fungi (Seydack 1990). The most important dietary items in the Cape were underground fungal bodies of Rhizopogon, bracken (Pteridium aquilinum) rhizomes and Blechnum punctulatum fronds. In the Eastern Cape, tubers of Vitaceae (Rhoicissus and Cyphostemma), vertebrate animal matter and foliage of Aloe ciliaris predominated (Seydack 1990). In Matobo N. P. in Zimbabwe, Jones (1984) recorded 29 different food items, including invertebrates (earthworms). During the wet season, 37% of identifiable food remains in faeces were fruit, 26% finely digested food and 19% herbaceous plant roots; during the cool dry season 49% was finely digested food and 41% fruit; and in the hot dry season 83% were woody plant roots. Both wild and cultivated fruits are eaten and Bushpigs may be particularly troublesome in maize fields, sugarcane and other planted crops (Skinner et al. 1976, Melton et al. 1989). Carrion is taken opportunistically (Skinner et al. 1976 and references therein, Jones 1984). Dietary items are procured from a feeding stratum of ± 0.5 m above and below ground surface through superficial rooting, deep rooting and grazing. During superficial rooting, which is the most commonly employed foraging technique, subterranean food items are exposed by lifting the snout forward and upward through the earth or litter layer. In moist or loose earth, the snout may be worked well into the soil in pursuit of deeper subterranean food items.

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Bushpig Potamochoerus larvatus.

Suitable dietary items for an omnivore are typically widely dispersed in space and time. This requires much searching and Bushpigs move continuously while foraging, stopping only when a clump of food is discovered. When foraging in a group they tend to be spaced out in a variably extended formation. Bushpigs are predominantly nocturnal (Breytenbach & Skinner 1982, Seydack 1990). The main phase of intensive foraging generally occurs from before dusk to midnight, with a secondary activity phase in the morning. Animals are active for an average of 13.7 hours per day (Seydack 1990). This represents a comparatively extended activity period and is presumably related to the search time required where food is dispersed. They visit habitat with poor vegetation cover only during the hours of darkness. Social and Reproductive Behaviour Bushpig sociality is characterized by the family group, usually consisting of a boar and a sow with one or two generations of offspring.There is typically only one adult sow per family group (sounder). Populations may be subdivided into four socio-spatial classes: territorial male–female pairs with or without progeny; solitary individuals, in the case of "" occupying non-exclusive home-ranges; and dispersing individuals that are mainly yearlings or subadults and nomadic adults. The average group size for the study populations in the Knysna Forest was 2.4 individuals, ranging between one and ten (Seydack 1990). Resource territoriality is maintained by spatially related high levels of aggression, notably between !!, and patrolling and marking behaviour. The area of the average home-range or territory

in the Knysna Forest was 7.2 km2 (3.8–10.1 km2; n = 8) (Seydack 1990). Home-ranges are traversed every one to four days, with an average daily ranging distance of about 3 km (0.48–5.84 km). The monogamous mating system is characterized by extended male–female pair bonds, lasting beyond mating and rearing periods, and by breeding of only the alpha-female within each territory. Breeding deferment of sexually mature, pre-dispersal daughter sows may be associated with alloparental care, consisting of grooming and remaining in close proximity to neonates. During rearing of the young, which remain with the parents long past weaning, substantial male parental care is involved. The offspring usually disperse from the family home-range when between 1.5 and 2.0 years old. Farrowing may take place in shallow ground hollows. During cold or wet conditions, farrowing nests may be prepared during parturition. Nests then consist of a shallow depression lined with grass upon which additional twig material and shrubbery is piled to form an insulating heap. Neonates follow the mother away from the birth nest or hollow within the first one to three days, initially for short distances only. Within the first three weeks postpartum the daily ranging of juveniles hardly exceeds 200 m. When compared with the average daily ranging distance, it is apparent that during these initial weeks the ranging of adults is constrained. One or both parents remain at the resting sites and close to the neonates, lying in body contact to protect the young against both cold and predators. Paternal care involves guarding of, and concern for the young, especially during the temporary absence of the mother on foraging bouts. Maternal behaviour conforms to the general passive suid pattern; in addition to suckling the young, there is some limited vocal communication, naso-nasal contact and defensive responses to neonate distress calls (Seydack 1990). Sounds made by the Bushpig include grunts, squeals, snarls and snorts as in other suids (Frädrich 1974). Primarily the dominant boar in a sounder may give alarm by uttering a long, drawn-out, resonant grunt. When foraging, individuals in sounders emit monosyllabic soft grunts, presumably to maintain contact. Male tusk pouch glands and digital glands in both sexes apparently play an important role in olfactory communication (Jones 1978). Marking primarily involves tusk gland marking by the " and ground scratch marking by both sexes. Male Bushpigs mark objects by wiping them with the opening of the pouch gland, usually up and down stems of small trees. Both sexes scratch the ground forcefully with the front feet, leaving clearly visible slashes. It is the dominant boar in the group that marks with his tusk gland, presumably conveying information about social status, whereas ground scratch marking appears to convey information regarding territorial tenure. Agonistic interactions include threat gestures, snout thrusting, pushing aside, squabbling, snap biting and chasing off. Display contests are apparently confined to "" and contribute to mutual assessment of dominance ranking. Actual fighting, as described by Skinner et al. (1976), is accordingly rare in male Bushpigs. In contrast, territorial !! are very aggressive in expelling intruders from the territory. When two adult sows fight they rapidly thrust and push with their snouts, striking blows sideways or slashing whenever the opportunity arises (‘nose fencing’). Forehead contact forms a crucial part of the contest and the snouts are held in the form of crossed swords against each other. The contestants push-butt each other with vigorous forward movements of the head, continually striving 35

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to regain close head contact. They may sustain severe wounds and wound sepsis is a significant cause of female mortality. Latrines consist of scattered accumulations of faeces that appear to be used intermittently over long periods of time. Faeces accumulations are sited predominantly where the ground surface is clear of vegetation, such as bare stretches of paths and roads or in canopy openings. They tend to be visually conspicuous, indicating a possible role in communication.

collected below Crowned Eagle Stephanoaetus coronatus nests (Seydack 1990). Bushpigs in NE KwaZulu–Natal have been recorded infested with eight ixodid tick species, of which Rhipicephalus maculatus was the most abundant, and one louse species Haematopinus latus (Horak et al. 1991a). Little is known about diseases in Bushpigs. Whether Potamochoerus spp. act as vectors of tick-borne diseases, such as trichinosis, African swine fever and trypanosomes, has yet to be verified (Vercammen et al. 1993).

Reproduction and Population Structure Males reach sexual maturity between 16 and 20 months of age, while !! may first conceive at 17–22 months. Depending on social position and nutritional status, successful breeding usually occurs much later. Only pair-bonded territorial sows rear young successfully. In southern Africa, the farrowing season lasts from spring to early autumn, with a pronounced spring peak (Sowls & Phelps 1968, Seydack 1990). The most favourable nutritional conditions for the lactating sow apparently occur during this period due to the abundance of new plant growth. Mean litter-size in the wild is 2.1 (n = 32; Seydack 1990), but often three to four piglets are born, the maximum recorded being eight (Phillips 1926, Milstein 1971). Gestation lasts 17 weeks, and neonates weigh between 600 and 1000 g (Sowls & Phelps 1968, Seydack 1990). Lactation lasts up to 5.5 months (Seydack 1990). Within all age classes, mortality is often due to starvation. In the Knysna Forests, significant mortality factors are inclement weather and predation for immature Bushpigs, and intra-specific strife for adults, especially in !!. Mortality estimates for one- and twoyear-old Bushpigs varied between 40 and 45% and were similar in both age classes. Adult mortality rates varied between 19 and 27% per annum. Sex ratio is usually 1 : 1. The oldest free-ranging Bushpig was estimated to be 18 years old, but only very few individuals reach this age (Seydack 1990). In a study involving both Western and Eastern Cape Bushpig populations, differences in life history traits were revealed. Generation length and life expectancy at birth were 6.6 and 2.8 years in the Western Cape (nutrient-poor environment) and 5.4 and 1.8 years in the Eastern Cape (nutrient-rich), respectively. Breeding !! in the Western Cape were fatter, while Eastern Cape !! were smaller and leaner but exhibited higher fecundity. These regional differences were interpreted as resulting from a higher dietary nutrient to carbon ratio and associated higher population turnover in the Eastern Cape in comparison with Western Cape populations (Seydack 1990, Seydack & Bigalke 1992).

Conservation IUCN Category: Least Concern. CITES: Not listed. Bushpigs still occur widely in suitable habitat, albeit at naturally low population densities, and are present in several well-managed protected areas across their range. There is little indication that the distribution of the species has been, or is at present, substantially altered by human activities. They may be subject to localized declines and range contractions in some areas due to large-scale habitat destruction or as a result of hunting for crop protection and local consumption (Vercammen et al. 1993). In southern Africa, they are also sporadically subject to official population control measures when they feed on crops. However, their preferred habitat, nocturnal habits and relatively high reproductive potential are such that it has proved generally difficult to eliminate Bushpigs from larger tracts of relatively densely vegetated habitat.

Predators, Parasites and Diseases Verified cases of Leopard Panthera pardus predation on subadult Bushpigs are recorded. Bushpigs were also commonly hunted by Robust Chimpanzees Pan troglodytes in Gombe N. P.,Tanzania (Wrangham & Bergmann Riss 1990). Bushpig mandibles belonging to individuals 2–3 months of age have been

Key References Seydack 1990, 1991; Seydack & Bigalke 1992; Skinner et al. 1976; Sowls & Phelps 1968; Vercammen et al. 1993.

Measurements Potamochoerus larvatus HB (""): 1256 (1100–1540) mm, n = 80 HB (!!): 1207 (1090–1410) mm, n = 74 T (""): 402 (335–432) mm, n = 6* T (!!): 361 (305–432) mm, n = 4* HF c.u. (""): 247 (225–290) mm, n = 59 HF c.u.(!!): 241 (217–270) mm, n = 53 E (""): 175 (151–203) mm, n = 6* E (!!): 178 (161–190) mm, n = 4* WT (""): 72.3 (55.0–93.0) kg, n = 84 WT (!!): 68.9 (54.0–85.0) kg, n = 104 Western and Eastern Cape, South Africa (Seydack 1983); *Zimbabwe, Smithers & Wilson 1979) Mean body weights of 52 "" and 61 !! from E Zambia are slightly heavier at 77.5 kg and 72.0 kg, respectively; the upper weight range recorded for a ! was 96.0 kg (Wilson 1968) Record tusk length is 30.17 cm for an animal from the Save R., Mozambique (Rowland Ward)

Armin H.W. Seydack

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Potamochoerus porcus

Potamochoerus porcus RED RIVER HOG Fr. Potamochère; Ger. Pinselohrschwein (Flusschschwein) Potamochoerus porcus (Linnaeus, 1758). Syst. Nat., 10th edn, 1: 50. ‘Habitat in Africa’ (West Africa); based on animals exported to Brazil (Simoons 1953).

Red River Hog Potamochoerus porcus.

Taxonomy The most recent review of the taxonomy of the Afrotropical suids by Grubb (1993b) concluded that the Red River Hog and Bushpig P. larvatus should be treated as two separate species based on their clearly distinct morphology, at least until further field studies take place to further investigate possible intergradation or hybridization (and see Lönnberg 1910, De Beaux 1924, Allen 1939). As geographical variation cannot be differentiated from individual variation, no subspecies are recognized (Grubb 1993b). Synonyms: albifrons, albinuchalis, mawambicus, penicillatus, pictus, ubangensis. Chromosome number: 2n = 34 (Melander & HansenMelander 1980). The chromosome complement consists of 12 pairs of (sub)metacentric autosomes and 4 pairs of acrocentric autosomes (Musilova et al. 2010). In Wonga-Wongue Presidential Hunting Reserve in Gabon, these animals are reported to have interbred with introduced Wild Boars Sus scrofa and to have produced wild offspring (East 1990).

Geographic Variation Vercammen et al. (1993) note that geographical variation is slight, though there is an east (largest) to west (smallest) cline in body size in specimens from eastern DR Congo and Cameroon, respectively. Grubb (1993b) was unable to differentiate any geographical variation from individual variation.

Description Smallest of the Afrotropical suids (Grubb 1993b), characterized by bright, russet-orange dorsal colouration, a narrow, pure white dorsal line starting behind the head, a striking facial mask and narrow, pointed, tufted ears. In boars there is an exostosis on each side of the snout extending laterally toward a hypertrophied apophysis on the canine sheath – each supports a single large external cutaneous wart. No infraorbital warts. Facial mask black with a whitish muzzle, white ‘spectacles’ around the eyes and white cheek whiskers. Ears black with a whitish upper rim and elongated tip, carrying a tuft of long white hairs. Hair on forehead and body relatively short and dense, flanks and belly carry some longer bristles.

Lateral view of skull of Red River Hog Potamochoerus porcus.

Ventral pelage light orange with long white and orange bristles. Pelage of young piglets brown with beige-yellow longitudinal stripes and some spots. Legs blackish in colour. Sparse tuft of hair at distal end of tail. It is likely that Red River Hogs have the same scent glands as Bushpigs (Seydack 1990): maxillary tusk glands and preputial glands in !!, milky secretions from the Hardarian glands located in the eye sockets, and digital glands in both sexes. De Boer (1980) also describes a chin gland. Females have three pairs of nipples. Males are larger than "". Skull similar in build to the Bushpig, with upper canines projecting outside the mouth cavity.

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Similar Species Potamochoerus larvatus. As far as is known, the two species are allopatric, although their ranges are contiguous in some places (Grubb 1993b). Larger; pelage dominated by longer, coarse bristles, rather than thinner, shorter hair; much more variable and darker shaded body colouration (reddish-brown, grey and black) and more variable facial pattern (head never masked); ear tufts not so long. Phacochoerus africanus. Sympatric over the south-west and southcentral area of its range (Sahel until S Chad and possibly S Sudan in the east). Occurs on treeless open plains and lightly wooded savannas, but avoids densely wooded vegetation. Grey-brown skin, sparsely haired except for distinct black, brown or reddish mane of long thick hairs on neck and shoulders. Especially young animals often have a fringe of white hairs on cheeks. Ascending process of the mandible elongated; maxilla deepened, zygomatic arch much deepened and orbits displaced posterodorsally so that they rise above the shortened braincase. Tusks very long and curved. Prominent genal and rostral warts. Distribution Endemic to Africa. Widely, but now patchily, distributed through the West and central African rainforest belt, from Senegal in the west, throughout the Guinea–Congo forest to at least west of the Albertine Rift. Further east and south-east replaced by the Bushpig, although the precise borders between the ranges of the two species remain unclear; for example, along the lower Congo R., below Kinshasa in DR Congo, the two species are separated only by this river – the smaller P. porcus occurring on the right bank and the large P. l. koiropotamus on the left (Angolan) side (Lönnberg 1910, Schouteden 1947, Grubb 1993b, Vercammen et al. 1993). Ghiglieri et al. (1982) observed pigs that they considered as intermediates between P. l. hassama and P. porcus, but Grubb (1993b), while noting that it was possible that the latter species may have made a narrow penetration of Uganda and hybridized with P. larvatus in the Kibale Forest, found there was insufficient evidence to support this. There are no confirmed records

Potamochoerus porcus

from Sudan or Chad, though Red River Hogs may occur in extreme SW Sudan. There are also as yet no reliable records from Gambia, which is just outside their natural range (Grubb et al. 1998). Grubb (1993b) mentions a record from Bioko I. based on teeth picked up in the field (now in the Tervuren Museum), but notes this has yet to be evaluated. Habitat Typically associated with rainforest and gallery forest, but also found in dry forest, savanna woodland and cultivated areas, although usually in close proximity to the rainforest and in regions with limited seasonality in terms of availability of water, food and cover (Oduro 1989, Vercammen et al. 1993). Also frequents hydromorphic forest clearings (Magliocca 2000) and mountains with steep slopes (Malbrant & Maclatchy 1949). Like the Bushpig, Red River Hogs are highly adaptable and may even benefit by the opening up of former forested areas by the creation of secondary habitats, the provision of cultivated foods and reductions in the numbers of their natural predators. Abundance Densities (individuals/km²) reported for Red River Hogs vary greatly and include: 3.1 (Fa & Purvis 1997); 1.3–5.6 (White 1994: forested part of Lopé N. P., Gabon); 18.4 (Tutin et al. 1997: galleries and bosquets in savanna ecotone of Lopé N. P., Gabon); 3.1 (Fa et al. 1995: Equatorial Guinea); and 2.0 (Hart, J. A. et al. 1996: Ituri Forest, DR Congo). Periodic aggregations on ephemeral resources might explain the higher estimates (see Adaptations). Adaptations The bright, highly structured colouration and strong pattern of the Red River Hog imply a high premium on intraspecific visual communication; likewise, body stance and positioning of the head and ears are also likely to play roles in visual and social communication. The facial warts of pigs are believed to have evolved in association with tusk length and shape, skull shape and combat patterns between "". Species with no, or relatively small, warts and short tusks tend to engage in lateral combat postures, whereas species with large warts, large tusks, broad heads and thick skulls tend to use frontal combat postures. Both anatomically and behaviourally, the Red River Hog occupies a somewhat intermediate position between the two types (Frädrich 1974). Red River Hogs sometimes aggregate into large groups and it is suspected that they migrate long distances (Malbrant & Maclatchy 1949, Abernethy & White 1999). It is at present not certain if such aggregations and migrations are a response to mast fruit crops in the rainforest, as is the case for White-lipped Peccaries Tayassu pecari and Bearded Pigs Sus barbatus. Observations in the Ituri Forest suggest this may be so: densities in a monodominant Gilbertiodendron dewevrei forest near Epulu varied with a factor of 80 (0.1–8.0 animals/km²), with the highest densities occurring during periods when there was a mast crop of G. dewevrei seeds. During years with no mast crop, fallen fruit was scarce and during the dry season even almost absent. Densities in a mixed forest type, where variations in fruit abundance were smaller, only varied by a factor of 5 (1.0–5.8 animals/km²) (Hart 2001). Foraging and Food Omnivores, eating fruits, nuts, seeds, roots and tubers, leaves, fungi, insects, earthworms, small vertebrates and carrion (Carpaneto & Germi 1989, Hart 2001); they also uproot seedlings and saplings. Food items are gathered by walking along and continuously probing the leaf litter and soil with the nose. When visiting a forest clearing at Odzala N. P. in Congo, they spent 53% of

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Potamochoerus porcus

Red River Hog Potamochoerus porcus frontal head signals: aggressive (above); submissive (below).

their time feeding, 17% resting and 20% moving about; consumption of roots and herbaceous vegetation (mainly Cyperaceae) occupied 67% of feeding time (Magliocca 2000). Seeds are a major component of the diet in the Ituri Forest (Hart 2001). They are able to crack open very hard nuts and hard-coated drupes such as Irvingia spp. and some Sapotaceae (J. A. Hart pers. comm.). The first sign of their presence is often the sound of nuts and seeds ‘exploding’ as they are cracked in their jaws, a sound that can be heard up to 200 m away (Abernethy & White 1999, J. A. Hart pers. comm.). Red River Hogs like to root in elephant dung for undigested seeds (Abernethy & White 1999), an activity that occupied 33% of feeding time at a forest clearing in Odzala N. P. (Magliocca 2000). However, the dung is also attractive later in the decay process, when young seedlings come up (J. A. Hart pers. comm.). In Odzala N. P., Red River Hogs mainly foraged for nuts of Panda oleosa (Pandaceae) and Strombosia sp. (Olacaceae) and for seeds of Strychnos sp. (Loganiaceae) (Magliocca et al. 2002). Red River Hogs are, therefore, mainly seed and sapling predators rather than seed dispersers; however, seeds of Uapaca trees pass through whole and are often found germinating in the dung (Abernethy & White 1999). In agricultural regions, Red River Hogs will raid crops such as manioc, peanuts and potatoes. Crop raiding becomes more frequent during the main dry season when forest fruit is scarce (Malbrant & Maclatchy 1949). Social and Reproductive Behaviour Red River Hogs live in family groups, containing "", !! and youngsters. Malbrant &

Maclatchy (1949) reported that bands of about 20 were common in Gabon and that large troops could reach 40 individuals; solitary animals are very rare. Elsewhere, Oduro (1989) recorded a mean group size of 10.6 (range 1–15) in Nigeria, while White (1994) mentions a mean group size of 33 in the forests of the Lopé Reserve, and Tutin et al. (1997) a mean of 29 in the gallery forests and savanna bosquets of the same park. In Odzala N. P., where both hunting and logging are absent, three groups that visited a clearing during the day were composed of two, seven and three (later joined by a fourth) individuals. During the night a troop of about 40 individuals was observed twice (Magliocca 2000). In Lopé N. P., they sometimes aggregate into groups of more than 100 individuals (Abernethy & White 1999). Groups of 30–60 have been reported from Guinea and the eastern DR Congo (L. Macky & J.A. Hart pers. comm.). In areas with high logging and hunting pressure, Red River Hogs tend to occur in smaller, silent groups active only at night (Abernethy & White 1999). In non-disturbed areas they are also active during the day. The degree of territoriality, mating system, family relationships and duration of association of adult "" with !! are not known. Observations in captivity show that "" regularly inspect the urine and the anogenital region of the !! to detect oestrus. The " will follow an oestrous ! as she walks around and will regularly nuzzle her vulva, flanks and belly. The ! licks the anal and inguinal region of the boar. De Boer (1980) also describes a playful head to head pushing as an element of courtship behaviour. A ! that is ready for mating shows the typical immobilization reflex prior to and during mounting. Females leave the group to give birth and lie-up in an elaborate nest, 50–60 cm high and constructed of leaves and twigs (in the forest) and grasses (in savanna regions) (Malbrant & Maclatchy 1949, Abernethy & White 1999). They later re-join the group with their piglets. Red River Hogs are in almost continuous acoustic contact with one another. However, their rich vocal repertoire has never been studied. Reproduction and Population Structure Analysis of faecal steroid metabolites in captive animals has revealed that reproductive cycles in the Red River Hog start by the end of Dec and last into the summer (Berger et al. 2006). In the wild, reproduction indeed appears to be seasonal with most piglets being born at the end of the dry season/onset of the wet season. In Gabon, Malbrant & Maclatchy (1949) found one ! with three foetuses in Mar and one with four in Dec. Oestrous cycle length is 34–37 days (Berger et al. 2006). An average of 3.4 piglets (range 1–6) is born after a gestation length of 120 days (Macdonald 2000). Piglets weigh 650–900 g at birth and are brown with beige-yellow longitudinal stripes and some spots. The piglets remain tightly huddled in the nest to limit heat loss for the first few days after birth. They soon start to follow the mother while she forages and start to nibble on food items she unearths or that fall from her mouth (Abernethy & White 1999). Population structure, birth and mortality rates have not been published for wild populations, although Oduro (1989) recorded an immature to adult age ratio of 2 : 1 in groups in Nigeria. In captivity, the oldest zoo-born individual reached an age of 18 years. One wildcaught animal lived at Frankfurt Zoo for 22 years before it died (Winkler 2000, Weigl 2005). However, most individuals probably do not live longer than 15 years (Winkler 2000). 39

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Predators, Parasites and Diseases In rainforest habitat, Leopards Panthera pardus prey on adult individuals (Hart, J. A. et al. 1996, Abernethy & White 1999), and Red River Hogs made up 20% of the biomass consumed by Leopards in Lopé N. P. (Henschel et al. 2005) and up to 50% in Ivindo N. P., Gabon (Henschel et al. 2011). Other potential predators include Lions and Spotted Hyaenas (Abernethy & White 1999, Breuer 2005), while Robust Chimpanzees Pan troglodytes (Alp 1993, Abernethy & White 1999), large pythons and large raptors prey on piglets. Male Red River Hogs fiercely protect themselves and their group members from carnivores and can win a fight with a Leopard (A. Gautier-Hion pers. comm.). Like Bushpigs (Anderson et al. 1998), Red River Hogs are most likely vectors for African swine fever and allegedly also trypanosomiasis, and are an intermediate host for trichinosis (Vercammen et al. 1993). Ntiamoa-Baidu et al. (2005) recorded the ixodid ticks Ixodes cumulatimpunctatus, Rhipicephalus cuspidatus, R. lunulatus, R. simpsoni and R. ziemanni on animals in Ghana. Conservation IUCN Category: Least Concern. CITES: Not listed. Although generally widespread and abundant, increased hunting pressure (for subsistence purposes, as an agricultural pest or because it is a vector of livestock diseases, and also for the commercial bushmeat trade) has made the species rare outside protected areas in a number of countries, particularly in the west and north of its range (Vercammen et al. 1993, Abernethy & White 1999). At the bushmeat market of Basankusu, DR Congo, 38% of the 13,831 carcasses recorded in 276 counting days were artiodactyls and of these 21% were Red River Hogs (J. Dupain pers. comm.). Seventyeight per cent of hunters interviewed in Gabon by Lahm (1991) cited the sale of Red River Hog meat as among their most important sources of revenue; only about one-third of their gained bushmeat

was retained for domestic consumption. Consequently, hunting has led to direct declines in abundance in, for example, the forests of S Gabon (Laurance et al. 2006); however, Red River Hogs seem to stand up to hunting pressure relatively well in other regions (e.g. DR Congo; J.A. Hart pers. comm.), although this requires further study. Measurements Potamochoerus porcus HB (""): 1113 (1030–1190) mm, n = 6 HB (!!): 1070 (940–1170) mm, n = 11 T (""): 433 (405–455) mm, n = 6 T (!!): 418 (395–435) mm, n = 8 HF c.u. (""): 242, 260 mm, n = 2* HF c.u. (!!): 235 mm, n = 1* E (""): 190, 205 mm, n = 2* E (!): 180 cm, n = 1** WT (""): 63.0 (51.0–80.0) kg, n = 6 WT (!!): 43.5 (33.0–56.0) kg, n = 11 Gabon (Malbrant & Maclatchy 1949) *AMNH (D. Lunde pers. comm.) **MRAC (W. Van Neer pers. comm.) Record tusk length is 279 mm for an animal from Gabon (Rowland Ward) Key References Abernethy & White 1999; Grubb 1993b; Hart 2001; Magliocca 2000; Malbrant & Maclatchy 1949; Oduro 1989; Vercammen et al. 1993. Kristin Leus & Paul Vercammen

GENUS Hylochoerus Forest Hog Hylochoerus Thomas, 1904. Nature 70: 577.

Hylochoerus is a monospecific genus, represented only by the Forest Hog H. meinertzhageni that has been, from the first, recognized as generically different from any other type of pig. The teeth and skull, in particular, differ in many ways from those of other, rootling suini

such as Wild Boars Sus scrofa, or bushpigs of the genus Potamochoerus (Ewer 1958, Harris & Cerling 2002). None the less, the functional significance of these peculiarities was not well understood at first (Thomas 1904). In the century since the Forest Hog was first described, many suid fossils have been discovered and some of these, notably fossil skulls of the polymorphic Kolpochoerus heseloni, have helped illustrate the evolutionary divergence, within Africa, of this genus from a more

Forest Hog Hylochoerus meinertzhageni head of female (left) and male (right).

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Forest Hog Hylochoerus meinertzhageni head: above, superficial myology; below, cross-section showing digastric muscle. ABOVE: Diagram of upward flexion in Forest Hog Hylochoerus meinertzhageni skull from less elevated ancestral condition (from Kingdon 1979). LEFT:

typically suid ancestry. White & Harris (1977) have suggested that Hylochoerus derived during the Pleistocene from the Plio-Pleistocene fossil pig lineage Kolpochoerus. When correlated with known behaviour, ontogenetic growth patterns and with evolutionary changes manifest in the skulls of fossil pigs, almost all the unique features of Hylochoerus cranial anatomy can be attributed to a relatively recent change in diet (Ewer 1970, Kingdon 1979). Unlike their closest, rootling relatives, Hylochoerus are surfacecropping animals, harvesting herbs and grasses with their capacious mouths and broad, sharp-edged lips. The teeth have also been modified for such cropping but all these changes are likely to have been relatively late, Pleistocene, developments (Harris & White 1979) and manifest quite clumsy improvisations in the service of a relatively new mode of mastication superimposed upon an older suine pattern. It can be shown that alterations in the balance of masticatory muscle forces and muscle attachments have led to expansions, contractions, tilts and bucklings in the skulls of Hylochoerus. Typically, suine molar teeth have thick enamel cusps that are well suited to crushing large fibrous material, such as roots, and cement is virtually absent in such hard, compact teeth. Rootling pigs have six teeth aligned down a long, relatively narrow toothrow that champs up and down in a predominantly vertical action. This exerts little or no lateral pressure on the teeth. By contrast, Hylochoerus have higher crowns with thin dentine and enamel spaced out within a thick matrix of cement (Ewer 1970). Wear on such teeth produces sharp ridges and this creates the sort of milling surfaces typical of leaf-eating teeth. However, instead of retaining a complete toothrow of six chewing teeth, Hylochoerus juveniles begin with a row of five teeth, the anterior ones of which are soon shed. This attrition of the premolars (and eventually of the first molar, too) is due to Hylochoerus masticating its food with a significant sideways action. This lateral action exerts more movement at the front end of the toothrow, less at the back. It is clear that the small premolars are ill-suited to withstand the pressure and lateral chewing effectively wears away and gouges out the premolars at a relatively early age.

The main power for this novel type of jaw movement comes from the pterygoid muscles that attach the lower mandible to the roof of the mouth (see Kingdon 1979). To operate effectively these muscles (which are exceptionally well developed in Hylochoerus) have several requirements: (1) an increase in the amplitude of their partly lateral action; (2) enlarged muscle size; and (3) an increase in their areas of attachment to the mandible and pterygoid. In addition to these changes, the abandonment of rootling in favour of cropping has greatly diminished the need for powerful leverage at the occiput, which is broad, but very short in Hylochoerus. The most obvious results of these alterations have been to broaden the back of the skull and bulk out the region immediately behind the toothrows. These changes, in turn, have had the phylogenetic effect of lifting the entire tube of the muzzle in relation to the basicranial axis, making the Forest Hog skull, relative to that of a rootling pig, more of a horizontal tube than a diagonal wedge. These functional changes in core regions of the skull’s interior have remoulded the plastic, malleable surfaces of the skull’s outer shell. Thus, the area between and immediately anterior to the eyes has been forced to ‘buckle’ outwards as the muzzle lifts. Shrinkage of the occiput has caused the forehead region to cave in and create a deep concavity above the cranium. The skull of Hylochoerus is, therefore, an outstanding example of how changes in a single parameter, in this case diet, have far-reaching effects on cranial structure. The likelihood that these innovations are quite recent suggests that some characteristics, such as early loss of premolar teeth, merely represent incomplete accommodation, by an earlier format, to entirely new challenges. Those challenges would seem to have included an increase in the extent and predictability of formerly closed forests as they became more fragmented, perforated or degraded by a variety of forces, notably drier, colder climates, the actions of elephants, humans and falling trees, and the spread of low-level herbage and grasses in lacunae within these moist habitats. Hylochoerus is not, therefore, perfectly adapted to its niche but can, in some respects, be described as the cobbled-together descendant of a lineage that abandoned successful earlier adaptations in response to changing circumstances. Jonathan Kingdon 41

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Hylochoerus meinertzhageni FOREST HOG (GIANT FOREST HOG) Fr. Hylochère; Ger. Riesenwaldschwein Hylochoerus meinertzhageni Thomas, 1904. Nature 70: 577. Kenya, ‘Nandi Forest, near the Victoria Nyanza, at an altitude of 7000 feet’; near Kaimosi [2134 m].

Forest Hog Hylochoerus meinertzhageni.

Taxonomy Despite its large size, the Forest Hog was the last wild pig – and one of the last large mammals – to be discovered in Africa. Between 1904 and 1930 seven subspecies were described, but recent taxonomic reviews (Ansell 1972, Grubb 1993b, 2005) only recognize three subspecies.The validity of one subspecies, H. m. schulzi, has been refuted by Mohr (1942), and shown to be a synonym of Potamochoerus larvatus hassama (Grimshaw 1998, Kock & Howell 1999). Pigs of this genus therefore belong to a single extant, polytypic species. Originally named Forest Pig, Hylochoerus was later qualified as ‘Giant’ Forest Hog, in reference to its great size in comparison with other wild pigs. However, it has been shown that only the nominate subspecies, H. m. meinertzhageni, truly deserves this epithet. All living Hylochoerus populations are substantially smaller than their prehistoric ancestral stock. Synonyms: gigliolii, ituricus, ituriensis, ivoriensis, rimator. Chromosome number: 2n = 32 (Melander & Hansen-Melander 1980). Description A large, heavily built pig covered in long black hair. Hair thickness, length and colour vary with age and location: the dorsal and lateral pelage is thick and dark brownish-grey to coalblack; ‘whiskers’ on jaw-line callosity white to yellowish; ventral pelage sparse and greyish to black. There is a dorsal crest of long, thick erectile hair. Hairs thick, oval in cross-section, and often split.

Lateral, palatal and dorsal views of skull of Forest Hog Hylochoerus meinertzhageni

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Forest Hog Hylochoerus meinertzhageni (female) skeleton.

Forest Hog Hylochoerus meinertzhageni (female) myology.

Broad, pointed ears fringed with black hair. Small orange-brown eyes, but very pale irises also occur, possibly reflecting nutritional or social stress (J. Kingdon pers. obs.). Adult boars significantly larger than !!, with a more bulky body and massive head; a dish-like depression over the forehead is surrounded by a circle of bony crests covered with bare skin. Zygoma are thickened and pneumatized, supporting inflated and naked infraorbital swellings. In boars these swellings commonly become stained with exudates from the preorbital glands which drain from a deep tear duct. The broad, flat snout ends in a very wide (10–15 cm), oval and swollen rhinarium with nostrils set wide apart. Legs robust (but can look spindly by comparison with the body mass); hooves large and rounded. Tail slender, flattened at its end, with long sparse bristles implanted laterally. Piglets are plain coloured, although dark-brown individuals with light-brown stripes have been reported (Grubb 1993b). Females have two pairs of nipples. The skull is broad when compared with that of Potamochoerus. Structure of the skull and facial musculature adapted to a folivorous rather than an omnivorous diet (Ewer 1970; for dissections and diagrams see Kingdon 1979). The dental formula is I 1/3, C 1/1, P 2/1, M 3/3 = 30; the single pair of upper incisors is often shed in adults. Tusks flare out horizontally and curve backwards in the same plane as the face. Dentition is hypsodont and much more specialized than in Potamochoerus, with molar cusps of the crown spaced out within a thick matrix of cement and sharp transverse ridges between enamel pillars. The anterior cheekteeth tend to be lost with age.

H. m. rimator (includes ituriensis): SE Nigeria to E DR Congo. Specimens from the western part of the range are on average smaller (length of skull 34.1–38.8 cm in "", and 33.0–37.7 cm in !!; Grubb 1993b) than those from further east (length of skull 36.6–49.5 cm in "", and 31.4–43.9 cm in !!; J.P. d’Huart, pers. obs.). Distinctly smaller and lighter than H. m. meinertzhageni, with more brown and yellowish hair in pelage. H. m. meinertzhageni: scattered populations from Albertine Rift Highlands of E DR Congo to Kenyan Rift Valley. Resembles rimator in shape of skull, but nominate race differs in its jet black pelage, larger size (length of skull 41.0–46.1 cm in "" and 38.1–42.7 cm in !!; Grubb 1993b) and size of the tusks. The cheekteeth are also different in that the cementum is developed at the expense of the enamel pillars, which are more widely separated (Grubb 1993b). Intergradation between the lowland forest rimator and this large highland race presumably occurs along the foothills of the Albertine Rift (Grubb 1993b).

Geographic Variation Three subspecies are provisionally retained, but most specimens appear to represent variants in a polymorphic species. Smaller specimens predominate in the western and central lowland forests, heavier ones in highland forests, east of the Rift Valley. H. m. ivoriensis: isolated populations from Guinea to S Ghana. Does not differ significantly from H. m. rimator in dimensions or features (length of skull in "" is 35.5–39.7 cm, and 33.3–37.2 cm in !!; Grubb 1993b), but the shape of the skull is distinctive and unlike that of other subspecies.

It is probable that Ethiopian Forest Hogs are derived from a common stock with meinertzhageni; however, given a wide separation of these two populations by a well-established eco-climatic boundary, d’Huart (1978) and Yalden et al. (1984) have suggested that it is possible that the Ethiopian population comprises a different, as yet undescribed, subspecies. Similar Species Potamochoerus larvatus. Sympatric in some localities in eastern Africa. Smaller and more compact, with pale pelage on face and nuchal/ dorsal crest; snout less swollen. Distribution Endemic to Africa. Unevenly distributed in scattered populations throughout undisturbed tracts of lowland rainforest in West Africa and on the right bank of the Congo R. Also present in transition gallery forests in the Guinean savanna zone, in highland mixed forests of Albertine Rift, and in isolated montane forests in Kenya and Ethiopia.Their range resembles that of the Bongo Tragelaphus eurycerus, which also exploits unstable forest-edge mosaics. In West Africa (S Guinea, Liberia, S Côte d’Ivoire, S Ghana), the range of H. m. ivoriensis now roughly corresponds with the least 43

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Rwanda since the late 1980s (B. Dowsett pers. comm.), but they may possibly survive in Nyungwe Forest. In the absence of reference specimens or photographic records, Forest Hogs are no longer considered to occur in Tanzania (Grimshaw 1998, Kock & Howell 1999). Some connecting corridors still exist within the complex of protected areas from the Rwenzori Mts in the north to the Virunga Mts in the south, but most of the remaining populations in this region are now isolated. A similar situation occurs in SC and WC Ethiopia (d’Huart & Yohannes 1995), where fragmented subpopulations are well isolated from the nearest neighbouring populations of Sudan, Uganda and Kenya.

Hylochoerus meinertzhageni

disturbed remnants of the lowland humid forest. The species may occur in other parts of Guinea, Sierra Leone and Côte d’Ivoire in montane forests (e.g. Mamou Mts, Foutah Djallon, Loma Mts) and in gallery forests (Outamba N. P., Comoé N. P.), but many records remain unsubstantiated (e.g. see Grubb et al. 1998). Forest Hogs have been extirpated from many intervening sites that connected primary forest and Guinean savanna. Their present distribution has become increasingly fragmented, though the full extent and pace of this process is difficult to document because this species is a premium target for the rapidly expanding bushmeat trade. Reports from Senegambia, Guinea-Bissau and Togo are unsubstantiated (Grubb 1993b, Grubb et al. 1998). Western populations of H. m. rimator (those between the Rivers Niger/Bénoué and Congo/Oubangui) still occur in five countries from the Nigeria/Cameroon border to N Gabon, N Congo and SW Central African Republic. The species is absent from Bioko I. and is believed to be extinct in the vicinity of Rio Muni in Equatorial Guinea. Forest Hogs have also disappeared from many areas in W Gabon and S Congo. Further east, this subspecies remains almost continuously distributed along the right bank of the Congo R. to the southernmost limits of the equatorial forest in E DR Congo in the south, to extreme N and NE DR Congo, E Central African Republic and SW Sudan in the north. It occurs in all types of primary, secondary and gallery forests. The northern limit of its range is in the vicinity of Massif des Bongo, south of Manovo–Gounda–St Floris N. P. in Central African Republic, where the progressive fragmentation of the forest galleries is likely to isolate this population in the future. This subspecies has disappeared from several sites in N Central African Republic and NW DR Congo. In the easternmost limits of the Forest Hog’s range (along the west branch of the Rift in E DR Congo, W Uganda and Rwanda, and along the east branch in E Uganda,W and C Kenya), the ‘giant’ form, H. m. meinertzhageni, now survives only in isolated montane forests, from 900 to 3800 m. There have been no records of Hylochoerus from

Habitat In common with the Red River Hog Potamochoerus porcus, the Forest Hog is more dependent on forest than other African suids. Throughout its range, it inhabits a wide variety of forest types, ranging from sub-alpine areas and bamboo groves through montane to lowland and swamp forests, galleries, wooded savannas and postcultivation thickets.Within these habitats, it occurs preferably where there is a convenient and permanent water source, thick understorey cover in some parts of the home-range, and a diversity of vegetation types. In E DR Congo and W Uganda, the Forest Hog is an ecotone species, preferring intermediate habitat zones where the edge effect is maximized and where resources from different vegetation types can be exploited within a limited area. This was well illustrated in Budongo, Uganda, where Kingdon (1979) mapped a 1600 km² block of mixed wooded grasslands, forest, forest/woodland mosaics and cultivation where he established the local ecological separation of Hylochoerus, Potamochoerus and Phacochoerus. The former shared habitat with one or other, or both species in areas where uniform forest and grasslands abutted, but Hylochoerus became dominant only in areas of extensive forest/grassland mosaic. In the Aberdare Mts in Kenya, the Forest Hog was extremely abundant during the 1970s and, over a period of five years (1970–1974), a total of many hundreds of sightings were recorded visiting Yathabara glade (a major waterhole/salt-lick in the Aberdare Mts of C Kenya). By contrast, there were only two sightings of Bushpigs P. larvatus and it is possible that competition for secure shelters (within a montane forest that was freckled with numerous small, grassy glades) might

Forest Hog Hylochoerus meinertzhageni.

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have been the primary cause of this imbalance between species (Kingdon 1979). In the central plain of Virunga N. P., Forest Hogs inhabit forest galleries (dominated by species of the genera Croton, Pterygota and Rauwolfia), dry forests (Euphorbia, Olea) and bush thickets (Capparis). These three habitats constitute a sequence in the natural succession from forest to savanna, and the hogs move between them, making full use of their resources. During the 1980s and early 1990s, Forest Hogs from forested areas of the central part of Queen Elizabeth N. P. (Uganda) increased and extended their range into grassland/ thicket mosaic areas, including the Mweya peninsula (Viehl 2003, Klingel & Klingel 2004a, b). In dense forest areas, Forest Hogs tend to concentrate in inselberg mosaics of savanna and forest, around clearings, or in mixed forest patches rather than in monodominant forest (Hart 2001). On a continental scale, the variety of forest habitats occupied implies a high degree of adaptability to local climatic conditions. Forest Hogs live in cold uplands (where night temperatures may fall to 0° C) as well as hot lowlands, but do not tolerate low humidity or prolonged solar radiation. Extreme mean annual rainfalls in Hylochoerus habitat vary from 800 mm in Masai Mara National Reserve to 3200 mm in W Liberia. Abundance A 1990 IUCN questionnaire survey suggested that this species had a restricted distribution at low density over most of its range, but was ‘widespread at low density and locally abundant’ in Guinea and Côte d’Ivoire, and ‘widespread at high density’ in Congo. The value of such assessments is obviously limited without further detail and validation, and, in any event, hog abundance has been widely reported to fluctuate over time (due to rinderpest outbreaks, and increases or declines in the number of predators). Until recently, the only explicit population density estimates came from DR Congo and these ranged from 0.4/km2 in Garamba N. P. to 2.6/km2 in the central plain of Virunga N. P. (d’Huart 1978). Local population densities may be much higher though, since these estimates were based on spot counts over large areas. Between Sep and Oct 1975 Kingdon (1979) estimated about 70 individual forest hogs coming into Yathabara glade from an estimated catchment area of about 70 km² (a density of about 10/km²). Since that time (when the density of this species was thought to have been the highest ever known in the area), Forest Hogs are known to have declined throughout the Aberdare Mts, primarily due to an influx of Lions Panthera leo (Klingel & Klingel 2004b); the neighbouring hog population on Lion-free Mt Kenya has seen no such decline. A 1998 count of known individuals on the very localized Mweya Peninsula, Queen Elizabeth N. P., gave a density of 9–13/km2 (H. Klingel pers. comm.). However, Klingel & Klingel (2004a) subsequently recorded a 30% fall in the density of Forest Hogs on the Mweya Peninsula; in two other study areas, the decline was 70–80%. The main cause of decline was predation by Lions, Spotted Hyaenas Crocuta crocuta and Leopards Panthera pardus, with competition from other herbivores, nutritional stress, diminished reproductive recruitment, emigration and road kills given as further causes. A 1999 sample transect census of Jibat Controlled Hunting Area near Ambo, Ethiopia, suggested a figure of 8.25/km² (F. Kebede pers. comm.). Comparable densities emerged from the frequency of observations and recorded group sizes in areas of suitable Forest Hog

Forest Hog Hylochoerus meinertzhageni tooth succession from left: juvenile, adult and aged adult.

habitat in Virunga N. P. and Queen Elizabeth N. P. (K. Viehl pers. comm.). Seasonal mass movements have been noted in forest clearings in N Congo (R. Ruggiero pers. comm.), and also in the Epulu Forest (Hart 2001). These have been attributed to masting cycles or similar fluctuations in the availability of grasses and other foods. Other explanations have been proffered, but documentation has been poor. Adaptations Features of fossil ancestors of Hylochoerus show that grazing is a relatively late development in this species and that some members of its ancestral lineage may have been substantially larger than the living species. Grass-eating has involved the need for powerful lateral chewing, and shaped the molars in a unique design (Bouet & Neuville 1930). These molar adaptations are very distinct from, and somewhat intermediate between, those of Phacochoerus and Potamochoerus. In the course of evolution the muscles and bony anchorage needed to support this has caused massive realignments in the skull (Ewer 1970, Kingdon 1979). The most significant of these has been the role of greatly enlarged pterygoid muscles (which serve side to side mastication). This enlargement appears to have acted as an ‘evolutionary wedge’ opening up the base of the skull, lifting the muzzle and buckling the external bones of the forehead. Thus, the external modelling of the Forest Hog’s forehead can be attributed to evolutionary changes in diet and masticatory function (see diagrams in Kingdon 1979).

Forest Hog Hylochoerus meinertzhageni underside of lower jaw.

The Forest Hog’s huge bulbous rhinarium is rarely used for rooting; unlike Potamochoerus, its prenasal bone is progressively losing its mobility by fusing with the intermaxillary septum, making the massive snout quite unsuited for excavating food from hard soil. By discarding use of the rhinarium for digging in all but the softest soils, the Forest Hog’s mouth has adapted more fully to grazing, involving a reshaping of 45

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the lips, the tusks and the nasal disc. Skull features are well adapted to competitive and fighting techniques by involving two different, often sequential, forms of frontal combat (Kingdon 1979, Estes 1991a). In the ‘snout-ramming/knocking’ phase adult and subadult"" join their broad nasal discs with heads horizontal and then test their weight and strength by pushing; during sparring matches between young "", both snouts are often pushed upwards, but contestants also deliver sideward blows and try to push the opponent’s snout to the side. This form of contest is facilitated by the slightly bulbous, flat-fronted nasal disc and the reinforced median wall of the snout. In the ‘forehead butting’ phase mature "" depress their heads and press their massively reinforced foreheads together and push until one withdraws. Sometimes contestants charge each other from a distance, in which case a very loud cracking noise can be produced should their foreheads clap together in exact opposition and thus compress air between their cupped foreheads. Thick protective tissues on the cheeks, heavy hollowed ridges round the forehead and zygomatic arch, a ‘doublehulled’ cranium (see Kingdon 1979) and the wide curving angle of the tusks can absorb or deflect the impact of these powerful clashes, but skull fractures, and occasionally deaths, can and do occur. The comparatively thick skin and coarse pelage with sparse hair (intermediate between the dense pelage of Potamochoerus and the bare skin of Phacochoerus) are adaptations that allow Forest Hogs to live in extreme environmental conditions both in closed and open habitats. Their activity patterns and feeding behaviour are adapted to both open and forested areas. In the grassland/ bushland/dry forests mosaic of Virunga N. P., Forest Hogs spend on average 25% of their daily time moving and foraging in savanna, 21% moving, foraging and wallowing in thickets or forested areas, and 54% resting in a sleeping site in the same habitat (d’Huart 1978). In lowland forest (notably in the Maya clearing of Odzala N. P., Congo), Forest Hogs spend 44% of the time feeding, 14% moving and 31% resting (F. Magliocca pers. comm.). In Queen Elizabeth N. P., during the wet season, the hogs feed during the day (as well as wallowing) and sleep at night, whereas they spend much of the day and part of the night feeding during the dry season, particularly if fresh growth is scarce, and may retreat to the lakeside when drought becomes extreme (Klingel & Klingel 1999). In the montane forest of Aberdares N. P., Kenya, the frequency of daylight sightings in Yathabara glade (from 5% to 27% in 1974) is inversely correlated with temperature; daylight foraging was commonest between Apr and Nov and minimal during the hot season, between Dec and Mar (Kingdon 1979). In the salines of Odzala-Kokoua, Nganongo (2000) observed the hogs wallowing in mudpools during the heat of the day, confirming this species’ intolerance of prolonged direct exposure to the sun. Social activity in Forest Hogs is strongly influenced by climate and season. Klingel & Klingel (2004b) found all members of family groups joined up in the wet season, stayed close together and synchronized activity when they rested, wallowed or walked, whereas they could split up, sometimes for several days, during the dry season. Foraging and Food Mainly a grass-eater and folivore. Field observations of feeding habits clearly document that Forest Hogs are by no means pure grazers. Depending on the seasonal content, stage of growth, and quantity of plant resources available in the various portions of their territory, they seem to display a great versatility

in the selection of their food. Both in montane areas and forest/ grassland mosaics, many species of grasses, sedges and herbs are cropped. Forest Hogs rootle very much less than other wild pigs and dig only in very soft or muddy soils. In Queen Elizabeth N. P., on the Mweya Peninsula, Giant Forest Hogs eat 46 of the 115 plant species that have been recorded there (Klingel & Klingel 2004a). In a savanna area of the same park, Forest Hogs were found to select more than 100 food plants and show a high flexibility in choice (Viehl 2003). In central and East Africa, Forest Hogs prefer to browse herbaceous growth or graze on mats of relatively short green grass. Among the monocotyledons, several species of Commelinaceae and Cyperaceae are grazed in addition to a large number of Gramineae. The selected dicotyledons include, among others, numerous species in the families Acanthaceae, Amaranthaceae, Compositeae, Convolvulaceae, Euphorbiaceae, Melostomaceae, Portulacaceae, Urticaceae and Salvadoraceae (Rahm & Christiaensen 1963, d’Huart 1978, Kingdon 1979). Studies on grass preferences in Virunga N. P. show that five major savanna species (Cynodon dactylon, Sporobolus pyramidalis, Panicum repens, Cenchrus ciliaris and Chloris gayana) were selected on a yearly average during 94% of grazing time (d’Huart 1978). Seasonal preferences by Hylochoerus for these grasses is less marked than those shown by Phacochoerus in contiguous Queen Elizabeth N. P., but the same grasses are selected by both pigs during the same periods (Field 1970b, d’Huart 1978). However, preferences for the same grasses are not repeated during each of both dry and wet seasons. In Virunga N. P., an order of preference (the same for both dry and wet seasons) was established by correlating the frequency of selection time with the protein and carbohydrate content of the grasses; these data showed that hogs select grasses when their nutrient and energetic content is highest (d’Huart 1978). A study on carbon isotopic composition of Hylochoerus hair from the Mweya Peninsula has confirmed this observation, estimating that Forest Hog diet actually comprised more than 95% of C3 dicotyledons over much of the year, but that the proportion of C4 grasses rose up to 25% in the wet season (Cerling & Viehl 2004). Young piglets are known to feed on grasses well before weaning. In dense forest both Forest Hogs and Bongos feed on mast seed in monodominant Gilbertiodendron stands in N Congo, but not in E DR Congo (Hart 2001). Of the time hogs spent (mainly in water-logged areas) in a forest clearing in Odzala N. P., 44% was spent feeding on herbaceous plants from hydromorphic areas; these plants, Enhydra fluctuans, Paspalum conjugatum, Ludwigia stolonifera and Rhynchospora corymbosa, were significantly richer in calcium and nitrogen than those growing in non-preferred ones and it was concluded that mineral-rich forage was the essential reason for hogs to visit the natural clearings (bais) (actual drinking was never observed; F. Magliocca pers. comm.). Examination of adult hog dung by various authors has revealed a variety of different materials, including poorly masticated grass stems and leaves, fragments of Iulus sp., shields of ticks (Amblyomma spp., Rhipicephalus spp.), earth, Forest Hog bristles and so forth. The animals seek salty earth, which they excavate with their tusks and their lower incisors; salt-licks may include termite mounds, shallow caves, dry river banks, or even the embankment of a deserted road. Occasionally, they eat meat and bones of carrion, eggs and larvae (Kingdon 1979). Coprophagy is not common, but piglets are fond of fresh elephant dung when available.

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Hylocherus meinertzhageni

Social and Reproductive Behaviour Forest Hogs live in relatively stable, non-territorial family groups (or ‘sounders’) consisting of one to three (occasionally four) !! and their young, and a dominant adult ". In Queen Elizabeth N. P., group sizes range from 2 to 19 individuals (very occasionally up to 24), with an average of 11.2 (Klingel & Klingel 2003, 2004b, H. Klingel pers. comm.). In contiguous Virunga N. P., average group size was 13.9 (d’Huart 1978). During the early 1970s in the Aberdare Mts, groups numbered 2–18 (average = 6.6); however, it was rare for more than four or five adult !! to associate for any length of time. In the Aberdares population some individual boars maintained a stable relationship with a group of !!, but some were effectively ousted during amalgamation episodes that were probably brief and may have been the direct product of convergence on a common resource, namely the artificial mineral lick in Yathabara glade (Kingdon 1979). These short-term displacements might give some support to the theory that dominance hierarchies may exist among "" observed within the same ‘group’. Such ‘groups’, with up to four adult "" and at least eight adult !!, have been recorded, but field observers agree that a single dominant boar does not accompany a sounder with more than four sows for any durable span of time. This may be primarily due to a weakening, as group size increases, of the familial bonds that tie sows to one another. Large groups consistently split into smaller groups and into small family units with a single boar predominating. Generally, only alpha-males have access to the !!, but copulations with peripheral individuals do occur outside the group (d’Huart 1978). In Virunga N. P., male tenures were recorded to last up to four years (d’Huart 1978), but in neighbouring Queen Elizabeth N. P., Klingel & Klingel (2004b) found the longest male tenure reached seven years. These researchers found infanticide was common immediately after a new " joined a group, in spite of all !! attacking the boar when he pursued any of their offspring (Klingel et al. 2001). Paradoxically, such risk to their young must strongly favour females’ maintenance of a long-term, stable relationship with one ". That this must be so is strongly implied by both sexes taking part in group defence, and by group cohesion being independent even of the alpha-male’s physical presence (Klingel & Klingel 1999). Depending on local group dynamics or the overall density of hogs (particularly the relative numbers of adult ""), the durability of male domination over specific groups of !! might vary or fluctuate from one locality or period of time to another, but the prevalence of infanticide, as revealed by the Klingels’ studies, must strongly favour stable family groups (Fimpel 2002). In the Aberdares, between 1970 and 1974, Kingdon (1979) thought that, in general, groups of !! were not strongly attached to a single boar, but the apparent turnover of "" attending female groups might have been a local and temporary by-product of a very rapid hog population increase and equally rapid decline over the period of observation. All studies have concurred that the core group of up to four adult !! is always stable, even in the absence of a dominant "; furthermore, with the exception of the alpha-male, all members of the group are directly related to each other. Indeed, younger !! patiently wait for aged sows and respond to the latters’ ‘lost’ calls (Klingel & Klingel 2004a). While young !! stay all their life in the group, subadult "" leave it voluntarily at the age of 2–3 years and form small temporary bachelor groups until they can compete for

Forest Hog Hylochoerus meinertzhageni.

the possession of !! and form their own sounder (d’Huart 1978, Klingel & Klingel 2004a). There are considerable fluctuations in the proportion of solitary animals, with a range of between 15% and 48% of all groups (in which singletons scored as a ‘group’) recorded at Yathabara glade. The number of Yathabara singletons, mostly boars, consistently peaked during periods of high temperature and declined, often very rapidly, when temperatures dropped (Kingdon 1979). A yearly average estimate of singletons among all hog sightings in Virunga N. P. was 17% (d’Huart 1978). Occasionally, several groups gather and stay together for a few hours forming aggregations of up to 40 or 50 individuals (d’Huart 1978, Kingdon 1979). It is possible that under such circumstances the bonds between certain individuals and their original group break down and new links are formed. These transfers would effectively scramble the individual composition of groups, but Klingel & Klingel (1999, 2004a) have established that female groups are consistently made up of close relatives. Fights between "" for possession of groups give the impression of being unritualized frontal combats between bachelors or alpha-males from other groups (but see discussion under ‘Adaptations’). These fights can result in severe injuries, including skull fractures (see Kingdon 1979) and male– male wounding can lead to significant mortality (d’Huart 1978). Contacts between groups were rarely observed in Virunga N. P., but when they did occur interactions between individuals from neighbouring groups were usually peaceful and dominated by reciprocal curiosity (d’Huart 1978). In Queen Elizabeth N. P., antagonism between groups is not uncommon in the dry season and takes the form of female-on-female running chases followed by retreat (Klingel & Klingel 2004a). Such fights notwithstanding, neighbouring groups usually actively avoid contact. Young "", instead, are prone to frequent fights and fights between rival bachelors can go on for weeks; Klingel & Klingel (2004b) recorded as many as 40 head-on collision-fights in just 10 minutes! Forest Hogs are not territorial as Mohr (1960) once suggested. Home-ranges were estimated to cover an area ranging from 2 to 47

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5 km2 in Virunga N. P. (d’Huart 1978). In Queen Elizabeth N. P., Klingel & Klingel (2004a) found that home-ranges covered several square kilometres with very extensive areas of overlap. At Yathabara glade the relative frequency with which particular groups were seen at the waterhole/salt-lick implied a graduated series of multiple, overlapping ranges, with those living nearest the glade being the most frequent visitors and the most distant ones the least frequent (Kingdon 1979). Rahm & Christiaensen (1963) estimated ranges of 7.5 km2 in Kahuzi-Biega N. P., DR Congo. Although home-ranges are not marked or defended as territories, and neighbouring groups share at different times the use of the same elements (tracks, wallow, salt-licks, latrines), hogs do defend core areas during periods when food is scarce, as it often is during the dry season; adult !! are particularly defensive (Klingel & Klingel 2004b). Dung is seldom dropped at random; favoured dung-sites are generally located at the base of a tree, under a thicket, or around a termite mound. The largest and most frequently used dung deposits are communal ‘latrines’ situated along access paths around the sleeping sites. The size and number of latrines near sleeping sites depend on the frequency with which the place is used and the size of the contributing group. For the positioning of latrines around typical sleeping sites see sketch-maps in Kingdon (1979). Courtship is rough and noisy; once the boar has identified an oestrous ! he keeps close to her, grunting loudly, urinating frequently and butting her hindquarters and flanks with snout or forehead until she finally stands. Copulation lasts from 1 to 10 min. Females isolate themselves from the group for 2–5 days to give birth, and piglets are born in a dry nest under a dense bush or a stack of branches or hay. They stay close to their mother and suckle standing up (sometimes, and momentarily, while walking) and the piglets respect a teat order (K. Viehl pers. comm.). Lying down and allosuckling are uncommon. Female hogs have been reported to be rather careless mothers, and piglets are frequently kicked, pushed or stepped on (Schneider & Viehl 2000). Forest Hogs have a wide range of vocalizations, including snorts, grunts and squeals; Fimpel (2002) and Klingel & Klingel (2004a) have recorded 18 different types of call, implying a highly differentiated auditory communication repertoire. Occasionally, Forest Hogs display aggressive and bold behaviour and family groups are known to chase away Leopards, Common Hippos Hippopotamus amphibius, Bongos and African Buffalo Syncerus caffer. However, they also graze peacefully next to African Buffalo, Savanna Elephants Loxodonta africana and various antelopes at Yathabara glade. In central Africa they commonly accompany Forest Elephants Loxodonta cyclotis, Western Lowland Gorillas Gorilla gorilla gorilla and African Buffalo on marshy bais (J. M. Froment pers. comm.), as well as Sitatungas Tragelaphus spekii and Red River Hogs (F. Magliocca pers. comm.). Reproduction and Population Structure In Virunga N. P. and in neighbouring areas, mating takes place all year round, with a peak in Feb–Mar and Aug–Sep, at the start of the wet seasons. Births occur mainly in Jul–Aug and Jan–Feb when optimal grazing is available for lactating !! and growing piglets. Weaning occurs after 8–10 weeks. Litters of up to six piglets (average = 2.5 in Virunga N. P.) are born after a gestation period estimated at 18 (Asdell 1946, K. Viehl pers. comm.) to 21.5 weeks (d’Huart 1978). Litters of less

than four piglets account for 96% of field records, but as many as 11 offspring are known (Prickett, in Kingdon 1979). Body growth is relatively rapid; adult size as well as sexual maturity are reached by both "" and !! at 18 months. No variation in female reproductive potential has been detected, and a yearly proportion of 46% of !! produce young. In the Virunga N. P. population, a reproductive potential of 113.6 young for 100 adult !! has been calculated. Net reproduction ratio suggests that each ! is replaced by 1.3 !!, showing an increasing population in that area during the period of observation (d’Huart 1978). In Queen Elizabeth N. P., the killing of piglets by newly arrived dominant "" seems to be sufficiently common to represent a major constraint on group dynamics, with up to 100% mortality in the litters of !! living with a newly established " (Klingel et al. 2001). Although such incidents can be reliably inferred, it is rare for them to be observed directly so that the overall fertility of !! at parturition has probably been substantially underestimated. Recorded sex ratios fluctuate between 1 : 1 and 1 : 2. The age ratio of the Virunga N. P. population is 67% adult (>18 months), 15% subadult (6–18 months) and 18% young. Seasonal changes in population structure are detected after the peaks of birth seasons. A mortality rate of 50–60% of piglets is estimated during the first year. It is mainly attributed to predation, but can also be due to weaning stress or to crushing by the mother. Additional mortality factors at adult age include fighting (""), poaching, predation, parasitism and disease. Life tables suggest an average life expectancy of 3.5 years and an average life-span of five years, with a maximum of 18 years (d’Huart 1978). Predators, Parasites and Diseases Major predators include Spotted Hyaenas, Leopards and Lions. Large eagles prey on young piglets in savanna habitats. The high density of Lions in Aberdares N. P. would seem to have been the principal cause of a drastic Forest Hog population decrease since the late 1980s (Klingel & Klingel 2004b, R. Kock pers. comm.). Inversely, the expansion of the Forest Hog population in Queen Elizabeth N. P. is attributed to a temporary collapse of the Lion population combined with several years of high rainfall (H. Klingel pers. comm.). Forest Hogs are hosts to a large number of ectoparasites, and a list of 11 species of ixodid ticks has been reported (d’Huart 1973). They are reportedly susceptible to rinderpest and populations have been greatly depleted by waves of rinderpest epidemics. Forest Hogs are also known to be a reservoir for African swine fever and trypanosomiasis, but their impact on the species is currently unknown. Periodic toxicity in food plants (notably Mimulopsis) has been invoked for die-offs of Forest Hogs in the Mau forest, Kenya (Simon 1962). Conservation IUCN Category: Least Concern. CITES: Not Listed. The status of all West African Forest Hog populations is cause for concern, particularly given the high rates of forest loss and fragmentation in the region. Elsewhere, Forest Hogs are very vulnerable because widespread deforestation and illegal hunting for local consumption and a rapidly expanding bushmeat trade has led to declining numbers, and rapid fragmentation of subpopulations. In some areas of the Congo Basin, Forest Hogs are avoided by shotgun hunters because their flesh is considered to have an unpleasant taste, but this is by no means a widespread aversion nor is the taint widely recognized.

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Hogs fall victim to snares and the meat is smoked to conceal its origin and sent to urban markets (R. Ruggiero pers. comm.). Protected areas known to have important populations of Forest Hogs include: Sapo N. P. (Liberia); National Park of Upper Niger (Guinea); Taï N. P. (Côte d’Ivoire); Bia N. P. (Ghana); Odzala N. P. (Congo); Minkebe N. P. (Gabon); the Sangha Tri-National complex (Central African Republic, Congo, Cameroon), where large groups graze the bais; Maiko N. P., Virunga N. P. and Kahuzi-Biega N. P. (DR Congo); Rwenzori Mountains N. P. and Queen Elizabeth N. P. (Uganda); Aberdares N. P. and Mt Kenya N. P. (Kenya); and Bale Mountains N. P. (Ethiopia). During the 1940s, an exceptionally high concentration of Forest Hogs was reported to have developed in the area of Rutshuru (near Virunga N. P., DR Congo). A special hunting operation around Rutshuru was organized in 1945/1946 to reduce the populations of wild pigs, which were judged responsible for crop damage; some 77 Common Warthogs, 329 Bushpigs and 619 Forest Hogs were destroyed (Hoier 1952). Both conservation (through captive breeding) and an obvious potential for domestication have been constrained by veterinary regulations designed to protect domestic pigs from any possible source of swine fever.This has led to extensive, but not always wholly rational, international bans on the import and export of wild pig species. The potential contribution of this grazing pig to protein

needs in Africa, under total or partial domestication, continues to be ignored in spite of increasing international calls to alleviate poverty. Measurements Hylochoerus meinertzhageni HB (""): 1374 (870–1900) mm, n = 7 HB (!!): 1172 (588–1780) mm, n = 6 T (""): 318 (230–360) mm, n = 6 T (!!): 282 (105–380) mm, n = 5 HF c.u.(""): 905 (790–1100) mm, n = 12 HF c.u.(!!): 828 (762–1000) mm, n = 10 E: n.d. WT (""): 177.4 (139.0–220.0) kg, n = 9 WT (!!): 150.6 (130.0–204.0) kg, n = 3 d’Huart (1978) The longest tusk on record, measured from the base of the extracted tusk along the outer curve to the tip, from Semliki Forest, Uganda, measured 394 mm (Rowland Ward) Key References d’Huart 1978, 1993; Grubb 1993b; Kingdon 1979; Klingel & Klingel 2004a, b. Jean-Pierre d’Huart & Jonathan Kingdon

Tribe PHACOCHOERINI Warthogs Phacochoerini Gray, 1868. Proc. Zool. Soc. Lond. 1868: 47.

The warthogs may be readily separated from the rest of the extant suids on the basis of their complex, elongated and very high-crowned third molars – often the only cheektooth retained in mature and elderly individuals. Hypsodonty has been selected as an adaptive response to the consumption of more fibrous or abrasive plants in progressively more open and arid-adapted types of vegetation. Hypsodont grazing suoids were thought to be absent in western Eurasia, but Pickford (1988) recognized a suine tribe Hippohyini to accommodate the hypsodont Miocene genera Hippohyus and Sivahyus from the Siwalik Hills of the Indian sub-continent. In Africa, three lineages of suids – the tetraconodont Sivachoerus– Notochoerus lineage plus species of the suines Metridiochoerus and Kolpochoerus – all displayed increase in size, height and complexity of the third molars through time, making them useful for biostratigraphic correlation (Cooke & Maglio 1972, White & Harris 1977, Cooke 1978a, Harris & White 1979). Kullmer (1999) compared the molar morphology of Plio-Pleistocene African suids with that of their extant counterparts and correlated increases in hypsodonty, length and complexity with adaptation to more open habitat and acquisition of a grazing diet. Harris & Cerling (2002) found, on the basis of stable carbon isotopes, that such a transition occurred during the Pliocene in the Sivachoerus–Notochoerus lineage, but that the earliest African representatives of the suines Kolpochoerus and Metridiochoerus were already well-established grazers.The diminutive Metridiochoerus modestus had a cranium that was very similar to that of Phacochoerus, but its teeth still retained the Y-shaped pillars that distinguish Metridiochoerus teeth from those of Phacochoerus.

Sectioned Common Warthog Phacochoerus africanus skull (old male) illustrating final position of third molars in maturity.

Contemporary warthogs are relatively small animals that probably derive from a still smaller immediate ancestor. However, some fossil Phacochoerini from the Plio-Pliestocene were as large as small rhinoceroses, an enlargement possibly selected for as these pigs moved out into more exposed habitats. John M. Harris 49

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GENUS Phacochoerus Warthogs Phacochoerus F. Cuvier, 1826. Dict. Sci. Nat. 39: 383.

Frontal views of: left, Common Warthog Phacochoerus africanus and right, Desert Warthog Phacochoerus aethiopicus.

Juvenile Common Warthog Phacochoerus africanus showing succession and migration of teeth indicated by arrows (from Kingdon 1979).

Phacochoerus includes two extant species, the Common Warthog Phacochoerus africanus and Desert Warthog, P. aethiopicus. Two species of warthog have long been recognized by palaeontologists (Van Hoepen & Van Hoepen 1932, Ewer 1956, 1957, 1958, Cooke & Wilkinson 1978); Cuvier (1817) also considered them different species, as did Cumming (1970). However, Lydekker (1915) placed all warthogs in one species and, subsequently, during most of the twentieth century, the Common Warthog has been incorrectly referred to as P. aethiopicus. The confusion has only recently been clarified with the discovery of an extant subspecies of the extinct Cape Warthog in Somalia, namely, the Desert Warthog (P. aethiopicus somaliensis) (Grubb 1993b, d’Huart & Grubb 2001). Divergence in mitochondrial and single-copy nuclear DNA sequences suggests differentiation as far back as the late Pliocene (Randi et al. 2002).The two species are distributed throughout wooded or open country of sub-Saharan Africa, and were present in North Africa during the upper Pleistocene.They are now absent from much of southern Africa, where their former extent of range is not well known. A comprehensive review of warthog systematics is provided by Grubb & d’Huart (2010). Both species are diurnal graminivorous pigs, relatively hairless, with a sparse pelage of coarse bristles not concealing the body outline. ‘Whiskers’ of long white hair are present along the lower jaw, white hairs in the ears, and there is a long, thick dorsal mane starting between the ears and ending at the base of the tail. The legs are densely haired except for the carpal callus, and there is a terminal tail tuft. The characteristic physiognomy includes a broad snout, enormous canine tusks curving upward as well as laterally, facial warts, and orbits elevated and placed far back; hence, the eyes are at the top of the head providing a clearer and more extensive field of view especially when feeding. There are distinctive warts on the cheeks and just below the eyes, which are better developed in "", and contribute to absorbing blows and cushioning the skull during combat. Legs are slender, relatively longer than in other genera; hooves relatively narrow.

The skull in warthogs is transformed by adaptations for graminivory, and so differs strongly from that of members of the genera Sus or Potamochoerus. The lateral outline is wedge-shaped, slanted backward due to the downward flexure of the facial part in relation to the neurocranium. Zygomatic arches are very broad. The snout is much shortened in front of the tusks, not elongated and vertical-sided as in Potamochoerus because digging is of reduced importance. The condyle (on the long ascending ramus of the dentary) and glenoid lie well above the level of the toothrow, facilitating crushing or shearing of forage. The ascending ramus of the dentary slopes backward. The superficial masseter closes the jaw and pulls it laterally, producing a shearing action. Raising of the jaw joint is accompanied by a deepening of the maxilla, accommodating the hypsodont cheekteeth. The extreme anteroposterior compression of the postorbital region reflects the reduction of the temporalis muscle, a rapid masticatory action not being required for plucking grass. Paraoccipital processes are lengthened allowing the digastric muscle to act as a jaw opener, even though the neurocranium is raised above the dental arcade. The adult dental formula is I 0–1/0–3, C 1/1, P 3/2, M 3/3 = 26–34. Only the first upper incisors alone are (variably) retained. Lower incisors lie in a shelf-like projection of the dentary, or are impacted, not erupting above the gum. The tubular upper tusk sockets are directed forward and outward; the upper tusks are large, curling outward and backward above the level of the snout, and are not abraded at the tips by wear with the lower tusks, which are compressed, wearing against the anterior face of the uppers. The tusks of !! are relatively large compared with other genera. Premolars are reduced and together with the anterior molars, commonly shed in adulthood. Molars are hypsodont, and formed of well-cemented, closely packed, littlefolded, oval to sub-triangular enamel columns; the third molar is considerably elongated by a large talon (heel). Peter Grubb

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Phacochoerus aethiopicus

Phacochoerus aethiopicus DESERT WARTHOG Fr. Phacochère du désert; Ger. Wüstenwarzenschwein Phacochoerus aethiopicus (Pallas, 1766). Misc. Zool. p. 16. ‘Promontoria Bona Spei advectus’; between Kaffraria and Great Namaqualand (South Africa, Eastern Cape Prov.), two hundred leagues from the Cape of Good Hope according to Vosmaer (1766).

Desert Warthog Phacochoerus aethiopicus. Phacochoerus spp. Outlines of rhinarium (snout) and underlying nasals and mandibular occlusion in: above, Common Warthog P. africanus; below, Desert Warthog P. aethiopicus.

ABOVE: RIGHT:

Taxonomy The Desert Warthog was the first species in the genus to be scientifically described and illustrated. Knowledge of this species was eventually eclipsed by the volume of information gleaned from the related Common Warthog Phacochoerus africanus in terms of numbers of museum specimens collected, extent of field studies, and published photographs, so diagnostic characters have been neglected (Grubb & d’Huart 2010). Polytypic, with two nominal subspecies, one extinct since about 1871. An East African population (delamerei) was named by Lönnberg (1909), who recognized its resemblance to the South African animal. Lydekker (1915) placed all warthogs in one species and has been followed by most authors in the twentieth century, though palaeontologists were aware that there were two species (Van Hoepen &Van Hoepen 1932, Ewer 1957, Cooke &Wilkinson 1978). Reappraisal of the status of P. aethiopicus (Grubb 1993b) has been followed by a study of its distribution in relation to P. africanus in the Horn of Africa (d’Huart & Grubb 2001), discovery of considerable genetic divergence from the latter species (Randi et al. 2002), and morphological differences between both species (d’Huart & Grubb 2005). See Grubb & d’Huart (2010) for a detailed review of the ‘rediscovery’ of the species. Synonyms: angalla, delamerei, edentatus, pallasii, typicus. Chromosome number: not known, but probably 2n = 34 as in P. africanus.

Grubb 2001). Lower incisors are absent or reduced to four or less, very small, hardly protruding from alveoli and probably always concealed by gums. Lower and upper canines are less curved than in P. africanus, the wear facet on the lower canine is differently placed, and the lower canines are less compressed (Ewer 1957). The third molars are also different: when all the enamel columns have come into wear, no roots have yet formed, unlike the condition in P. africanus. At this stage, all the columns are of about the same length and are able to continue growing, extending the life of the tooth (Van Hoepen & Van Hoepen 1932, Ewer 1957, Cooke & Wilkinson 1978). Sphenoidal pits in the neurocranium floor on each side of the vomer are enormously enlarged and opened out, occupying the whole area between internal nares and basioccipital and basisphenoid, deepening the vomerine ridge; they are separated from the auditory bullae only by a thin wall of bone. Zygomatic arches are robust with large sinuses forming a spherical inflation of the jugal (malar) just in front of its articulation with the squamosal (temporal) (De Beaux 1922, Ewer 1957). A fine ridge runs down the front of the skull from the orbital rim across the lachrymal and maxilla, and while not necessarily diagnostic, characterizes many skulls of this species (d’Huart & Grubb 2001). In the few available samples, skull lengths for the two sexes do not overlap, suggesting perhaps greater sexual dimorphism than in the Common Warthog.

Description Very poorly known species in spite of being relatively abundant in East Africa. South African animals are known from two illustrations indicating a small-eared animal with dark face and snout over a light cheek and lower jaw, and a swelling round the eye, below which is a drooping blunt wart. Skull similar in proportions to that of the Common Warthog, but diagnostically distinguished by other cranial and dental characters. The upper incisors are always absent (d’Huart &

Geographic Variation P. a. aethiopicus (Cape Warthog): formerly southern part of the species’ distribution. Known from two illustrations (eighteenth and nineteenth century). Aart Schouman’s painting of the lectotype (Tuijn & Van der Feen 1969) shows a stout animal with relatively small ears, dependent, blunt suborbital warts, and contrasting colouration between a dark face and snout and light cheek and 51

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Lateral and frontal views of skull of Desert Warthog Phacochoerus aethiopicus.

lower jaw. External differences between a boar in the menagerie of the Zoological Society of London and a Common Warthog from Eritrea were described as ‘very obvious’ (Sclater 1869): flanks less naked, hairs on the back and nape of the head less thick and shorter; ears shorter, less pointed and less naked, densely coated with hair; and whiskers less developed.The illustration of the head shows a swelling round the eye and a broad, drooping suborbital wart with a blunt end. P. a. delamerei (Lord Delamere’s or Somali Warthog): northern part of the species’ distribution. Suborbital warts in adult "" of a characteristic shape – curled or bent down at the tip. Skulls from N Somalia are smaller than those from S Somalia and Kenya, but sample sizes are very small. Recorded as having a blond dorsal mane in Somalia. Characters of canines and third molars recorded in P. a. aethiopicus not yet studied in this subspecies. Other dental and cranial characters identical with those of the nominate subspecies. Genal warts in adult "" unlike those known from P. a. aethiopicus, characteristically shaped, curled or bent down at the tip though with much variation in volume, form and orientation. Tips of ears bent backwards, appearing to have rounded or blunt tips and an angular contour. Suborbital region swollen, forming pouches that often extend to the base of the genal warts, resembling those known from P. a. aethiopicus. The comparatively broader skull and its shorter basioccipital region give the impression that the head is more egg-shaped than diabolo-shaped (d’Huart & Grubb 2005). Similar Species Phacochoerus africanus. Largely allopatric, but recorded in sympatry in Tsavo West and Tsavo East N. P. (Culverwell et al. 2008). There is also a population occupying an enclave within the range of P. aethiopicus in N Somalia (d’Huart & Grubb 2001). Attain larger size, at least in some populations, with skull length up to 466 mm in "" and up to 409 mm in !!; they possess functional incisors and very small sphenoidal pits; lack thickening of zygoma; canines

Phacochoerus aethiopicus

reported to be more curved and third molars less specialized; genal warts cone-shaped and usually not with bent or curled tips; ears leaf-shaped, with pointed tips and a sinuous contour; suborbital areas without a pronounced swelling; head looks more diabolo-shaped (differences illustrated in d’Huart & Grubb 2005). Occupy a much wider range of habitats. Distribution Endemic to Africa. A polytypic species occurring in the Somali–Masai Bushland BZ and formerly in part of the SouthWest Arid BZ, but not in intervening areas. Historical Distribution Formerly South Africa, in the south-eastern parts of former Cape Province and apparently adjacent parts of KwaZulu– Natal. Early records of wild pigs do not usually discriminate between Bushpig Potamochoerus larvatus and warthogs and the two species of warthog were not distinguished in the field. There are no records of warthogs from most of the Western Cape and Northern Cape. The Desert Warthog is known by reliably identified specimens from only the following localities: the type locality; between the Sondags and Boesmans Rivers in Eastern Cape; the upper Orange R., Northern Cape; and KwaZulu–Natal (Grubb & d’Huart 2010). It is not known whether the distributions of the two species overlapped or whether they were strictly allopatric, although the Desert Warthog ranged north into the former Transvaal in the late Pleistocene (Ewer 1957). Current Distribution Distribution in East Africa does not seem to have changed significantly even in recent times, and they remain known only from E Ethiopia and Somalia, and in Kenya mostly north of the Ewaso Ngiro R. The range has been found to extend southwards to Tsavo West and East National Parks (west of Athi River and south of the Galana R.; Culverwell et al. 2008, de Jong et al. 2009). Habitat The Desert Warthog is a species of open arid regions. The South African population occurred in the Karoo but there is no

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information on habitat preferences and habitat use. Its distribution may have been limited by winter temperatures. The range of the northern population lies within the Somalia–Masai Bushland BZ, with vegetation of ‘Somalia–Masai Acacia–Commiphora deciduous bushland and thicket’ and ‘Somalia–Masai semi-desert grassland and shrubland’. These vegetation types range from xerophylous bush and open woodland to sub-desert steppe. Dominant grasses are Chrysopogon species in more arid areas and Chloris species in semi-arid habitats. Desert Warthogs prefer plains on predominantly sandy soils, and avoid hilly terrain. Most records are from lowland areas below 200 m, a few are at higher altitudes, but none is above 1400 m. Desert Warthogs are dependent on occurrence of water and shade and occupy regions with rainfall of 100–600 mm per annum. Areas with higher rainfall, which correspond to the zone above the 1000 m contour, are avoided, as are the driest, hottest and most desertic regions with rainfall less than 100 mm per annum, corresponding with the hot coastal zone of Somalia between 12 and 9° N, and extending to the eastern tip of Ogaden.

Predators, Parasites and Diseases Likely predators include Lions Panthera leo, Leopards P. pardus and hyaenas. Extinction of the South African population may have been hastened by rinderpest.

Abundance The Desert Warthog was one of the most widespread and common game animals in Somalia, at least in the 1960s. Ground and aerial surveys in the Nugaal valley, Somalia, as well as an aerial survey between Garowe and Ras Hafun in 1989, confirmed that warthogs are locally abundant (P. Moehlman pers. comm.). Similarly, field observers reported that warthogs were very abundant in 1996 in Mahadday and Jowhar Districts of Middle Shebelle (A. Massarelli pers. comm.). In Ethiopia, they are common in the whole Ogaden region and can be observed both in family sounders in bushy areas and in larger aggregations of up to 30 individuals around permanent wells and close to towns (Wilhelmi et al. 2004). In Kenya, they were reported to be abundant near water-points in Garissa District, Kenya, in 1999 and 2004 (T. Butynski & A. Caron pers. comm.), common on the Laikipia Plateau and in the Samburu/Buffalo Springs/Shaba National Reserves area, but rarer towards Maralal and Marsabit in 2000 (S. Williams pers. comm.).

Measurements Phacochoerus aethiopicus Skull measurements:

Adaptations Morphologically, the Desert Warthog is the most specialized living suid, but nothing is known of its habits and physiology. As incisors are functionally absent, lips and gums are used to detach or pick up food items. Rootless third molars suggest adaptation to a more abrasive diet. The specialized condition of the zygoma and differences in wart shape suggest the style of combat between "" is different from that of the Common Warthog. Foraging and Food The potentially longer-lived and rootless third molar and absence of functional incisors imply significant differences in diet and feeding techniques from the Common Warthog. These await investigation. Social and Reproductive Behaviour Physiological adaptation to drier, and perhaps, colder environments than the Common Warthog probably influence, say, their use of shelters and adaptation to watering regime. These, in turn, are likely to be correlated with currently unknown differences in social structure.

Conservation IUCN Category: Least Concern. CITES: Not listed. The Desert Warthog is known to exist in Samburu, Buffalo Springs, Shaba and Dodori National Reserves, and in Tsavo West and East National Parks, Kenya. From the distribution of reliably identified specimens, it will probably be found in Malkamari N. P. and Losai, Marsabit, Arawale and Boni National Reserves of Kenya, and Lag Badana/BushBush N. P. in Somalia. In Ethiopia, Desert Warthogs are likely to occur in Babile andYabelo Wildlife Sanctuaries and associated Controlled Hunting Areas (the Lower Wabe Shebelle and Borana, respectively) in Ethiopia (d’Huart & Grubb 2001, Culverwell et al. 2008). Although the species is widespread in the Ogaden, there is a concern that hunting for bushmeat trade may become a threat (Wilhelmi et al. 2004).

P. a. aethiopicus GLS (""): 378.0 (366–393) mm, n = 4 GLS (!!): 330, 352 mm, n = 2 GWS (""): 224.2 (220–230) mm, n = 3 GWS (!): 179 mm, n = 1 South Africa P. a. delamerei GLS (""): 393.0 (369–407) mm, n = 6 GLS (!): 328.7 (325–331) mm, n = 3 GWS (""): 228.3 (209–245) mm, n = 6 GWS (!): 182.0 (169–197) mm, n = 3 Kenya and S Somalia (Jubaland) GLS (""): 357.0 (348–364) mm, n = 5 GLS (!): 301 mm, n = 1 GWS (""): 202.0 (192–227) mm, n = 5 GWS (!): 167 mm, n = 1 N Somalia Museum specimens (BMNH, NMK, PCM) and from literature; a single specimen in the AMNH from Galma Galla, Garissa District, Kenya, had the following measurements: TL: 1755 mm; T: 440 mm; HF c.u.: 275 mm; E: 142 mm (T. Butynski pers. comm.) The record tusk length (measured along the outer curve from the base to the tip) is unclear due to lack of distinction between the two warthog species in most record books (see Butynski & de Jong 2010) Key References Cooke &Wilkinson 1978; d’Huart & Grubb 2001, 2005; Ewer 1957; Grubb & d’Huart 2010;Tuijn & Van der Feen 1969; Van Hoepen & Van Hoepen 1932. Peter Grubb & Jean-Pierre d’Huart

Reproduction and Population Structure There is currently no information available for the species. 53

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Phacochoerus africanus COMMON WARTHOG Fr. Phacochère; Ger. Warzenschwein Phacochoerus africanus (Gmelin, 1788). Syst. Nat., 13th edn, 1: 220. ‘Habitat in Africa a capite viridi ad caput bonae spei’; restricted to Senegal, ‘Cape Verd [Verde]’ (Lydekker 1915: 373).

Common Warthog Phacochoerus africanus.

Taxonomy The Common Warthog is here treated as a species distinct from the Desert Warthog P. aethiopicus (Ewer 1957, Cooke & Wilkinson 1978, Grubb 1993b, 2005). Four geographical variants of P. africanus were provisionally recognized by Grubb (1993b) largely on the basis of skull size and proportions indicating a clinal species. Muwanika et al. (2003) investigated patterns of genetic variation in 181 specimens from 24 localities in Africa. They identified 70 mitochondrial haplotypes that clustered into three well-defined clades representing individuals from western, eastern and southern Africa. They concluded that differentiation at both mitochondrial and nuclear loci supported the existence of three lineages that resulted from Pleistocene climatic fluctuations. Synonyms: aeliani, barbatus, barkeri, bufo, centralis, fossor, haroia, incisivus, massaicus, sclateri, shortridgei, sundevallii. Chromosome number: 2n = 34 (Bosma 1978); the chromosome complement consists of 14 pairs of (sub)metacentric autosomes and two pairs of acrocentric autosomes (Musilova et al. 2010). Description At first glance an adult Common Warthog jauntily trotting across an open savanna with its tail and mane erect and large head held high resembles a miniature rhinoceros rather than a typical pig. The head is unusually large, the forehead, muzzle and snout broad and flattened, with the eyes placed high and wide. Ears are prominent and broad rather than pointed, and can protrude well above the level of the head. Warts, from which the animal derives its name, are formed of tough fibrous tissue and comprise the

infraorbital warts attached to the zygoma, mandibular warts along the angle of the lower jaw, and, in !!, supraoral warts situated on the snout behind the tusks. A dense, side-burn-like cover of long white bristles generally obscures the elongated mandibular wart. The neck is short and the nape, withers and back carry a mane of long black and brown bristles. The mane can be erected in displays associated with fight/flight situations. The Common Warthog’s dark grey, almost black colour is determined by its skin colour rather than by its sparse coat. The skin is sparsely covered with white bristles and can take on differing hues of grey or brown depending on the colour of the soil in its holes or wallows. The tail is long and slender with a flattened end carrying bristles. The legs are proportionately longer than in other suids and knees bear characteristic callosities that are present even in the foetus. The hooves are narrower than those of the Bushpig Potamochoerus larvatus. Both the preorbital area and the lip carry dermal secretory glands (Estes et al. 1982). Tusks and warts are markedly larger in !! and "" have one pair inguinal and one pair abdominal nipples. The skull slopes sharply forward from the braincase to the nostrils, and the rostrum is considerably elongated. The orbits are small, and there are distinct supraorbital ridges on the inner sides. The supraoccipital crest is well developed, lying at an angle backwards of the braincase, thereby leaving a wide area at the back for the attachment of the massive neck muscles. The paraoccipital processes are long, extending nearly to the level of the bottom of the angle of the mandible (Skinner & Chimimba 2005). The blunt upper

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wear while in the extinct Cape Warthog P. aethiopicus aethiopicus and presumably also in P. a. delamerei roots are formed only once all of the columns have erupted and are in wear (Shaw 1939, Cooke 1949, Grubb 1993b). Unlike any other African suids, the roof of the skull of P. africanus shows two deep sphenoidal pits behind the internal nares region (Grubb 1993b). Geographic Variation P. africanus africanus: Mauritania eastwards to S Ethiopia. P. a. aeliani: confined to Eritrea, N Djibouti and Somalia. P. a. massaicus: eastern and central Africa, in Zambia, Malawi, N Mozambique, Tanzania and S Kenya. P. a. sundevallii: SW Angola, N Namibia, Botswana, Zimbabwe, S Mozambique and parts of South Africa. Similar Species Phacochoerus aethiopicus. Only recently distinguished as a living

separate species based on cranial features and molecular evidence (Grubb 1993b, Randi et al. 2002; and see Grubb & d’Huart 2010). A supposedly consistent distinguishing feature in the field is the shape of the infraorbital warts in "": in the Desert Warthog they have a distinct anterior bend, whereas in the Common Warthog these generally point directly outwards (see also species profile for Phacochoerus aethiopicus). The differences between the two species are illustrated by d’Huart & Grubb (2005). Largely allopatric, but recently recorded in sympatry in Tsavo West N. P. (Culverwell et al. 2008). Potamochoerus larvatus. Also occurs in moister savanna habitats, but readily distinguished from warthogs in the field by its distinctly shaggy, reddish-brown coat, rounded back, pointed pixie ears, low slung head, more pointed snout without upturned prominent tusks or warts, and its loping gait. Unlike warthogs, the tail is never held erect while running. The more brightly coloured Red River Hog Potamochoerus porcus and the much larger Forest Hog Hylochoerus meinertzhageni, which has a well-developed black shaggy coat, occur in forests and their ranges have only marginal overlaps with the Common Warthog.

Lateral, palatal, dorsal and fontal views of skull of Common Warthog Phacochoerus africanus.

canines, or tusks, flare upwards from the snout and in older animals, particularly !!, curve inwards. The lower canine is triangular in profile, scimitar-like and is sharpened against the lower edge of the upper canine. Unlike the upper canine, its dentine core is covered by enamel. The key dental features distinguishing this species from the Desert Warthog are the presence of one pair of upper incisors and two to three pairs of lower incisors. Upper incisors are absent in Desert Warthogs and lower incisors, if present at all, are rudimentary (Grubb 1993b). The adult upper and lower incisors of the Common Warthog also have deep roots and prominent crowns, which converge and make contact. The structure of the third molar also differs between the species. In P. africanus roots are formed in the anterior molar columns before all the cusps have come into

Distribution Endemic to Africa. Their former distribution extended across the West African Guinea Savannah and Sahelian zone from Senegal, Gambia and extreme S Mauritania eastwards to Sudan, Ethiopia and Djibouti and from there southwards through East Africa to the savannas of southern Africa. The Tugela R. marks their former southerly limit of distribution in KwaZulu–Natal (Rowe-Rowe 1994). Historically, the Common Warthog did not occur in the arid Karoo of South Africa, where it was replaced by the extinct Cape Warthog P. aethiopicus aethiopicus. Today, the Common Warthog remains widespread in sub-Saharan Africa, being found in scattered populations in West Africa eastwards to Ethiopia and then southwards in protected areas and unsettled, or very lightly settled, areas in East and south-central and southern Africa. The continuous expansion of the Sahel-zone has resulted in a marked contraction in the species’ former range in the north since the early 1980s, and accounts for its probable extinction in Niger (J. Newby, in Vercammen and Mason 1993) although it is possible they may still persist in the south-central Aïr Mts (J. Newby pers. comm.). 55

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shifts in habitat densities occurred as these animals changed from wet season to dry season feeding grounds within their home-ranges in the Sengwa area. Common Warthogs will congregate on the flush of green grass following dry season burns.

Phacochoerus africanus

Common Warthogs are still recorded in the savanna woodlands of SW Mali (Chardonnet 2001a). They have been reintroduced to some parts of their former range in southern Africa (e.g. in Swaziland, where the indigenous population went extinct), and introduced elsewhere (e.g. in NE Algeria, as a result of an escaped captive population, and to the Eastern Cape Province of South Africa). Habitat Confined to moist and dry African savanna grasslands, open bushlands and woodlands and usually within range of perennial surface water (although water is not considered to be an essential habitat requirement; Skinner & Chimimba 2005). Usually absent from forests, extensive thickets, cool montane grasslands and deserts and succulent steppes. However, the population in the Goda Mts in Djibouti mainly occupies forested areas (Künzel et al. 2000). Although typically associated with lowland savannas, they reach elevations of 3500 m in the Ethiopian Highlands (and large numbers survive in Bale Mountains N. P.;Yalden et al. 1996). The need to sleep in holes in the ground at night links the distribution of this species to the formerly more widely distributed Aardvark Orycteropus afer, although in some parts of West Africa Bigourdan (1948) reported that Common Warthogs generally did not sleep in holes at night and only used Aardvark burrows as a last resort. They are capable of excavating their own holes and often modify existing holes extensively. Subterranean erosion gullies are also used and holes and gullies may be shared with porcupines that use them by day (Cumming 1975). In a quantitative study of habitat selection, Cumming (1975) found the highest densities of Common Warthogs in drainage line grasslands on nutrient-rich soils and in Acacia savannas on alluvial terraces. These habitats supported highly palatable grasses, such as Urochloa mossambicensis, which were grazed during the wet season and other species, such as Digitaria milanjiana, which provided an abundant store of rhizomes that were rooted during the dry months. Lowest densities occurred in miombo woodland habitats and in Combretum and Baikiaea thickets and woodlands on deep sandy soils. Seasonal

Abundance Seldom a dominant component, either numerically or in terms of biomass, within African large herbivore communities, although in some protected areas local densities may exceed 15–20 ind/km2. More typically, densities range between 1 and 10/km2 in protected areas, although local densities of 77/km2 were found on short grass in Nakuru N. P. (Radke 1985, 1991a). The overall number of Common Warthogs in southern Africa (Angola, Zambia,Tanzania and southwards) has been estimated at about 250,000 (Cumming 1999). Estimates for the numbers of Common Warthogs in West and north-central Africa are not available although they occur in most protected areas in the savannas. Populations in Benin, Central African Republic and N DR Congo were greatly reduced by rinderpest during the 1980s (Vercammen & Mason 1993). Populations are susceptible to droughts and resulting population crashes have been reported in several parts of Africa (e.g. Mason 1990a). However, the potentially high reproductive rate of Common Warthogs does allow for rapid recovery when conditions improve. Adaptations Differs from other suids in several important respects and these differences emphasize that Common Warthogs are suids beautifully adapted to dry savannas and steppes. The unusually large head, with eyes set high and wide, provides a wider field of vision than other suids.The snout and teeth are adapted to grazing and to rooting hard ground while the warts and tusks reflect displays and fighting style. Their gait, with head held high, is more akin to that of an antelope than the pigs. Common Warthogs are the smallest of the diurnal bare-skinned large mammals and, unlike any other ungulate, sleep in holes at night. The subcutaneous fat layer is very thin, if present at all, and, in contrast to other suids, they are markedly thermolabile and their daily range in body temperature can vary by as much as 7 °C (Cumming 1975). Piglets are unable to maintain their body temperature without huddling and access to shelter (Sowls & Phelps 1966) and even at one year old they will die if exposed to early morning temperatures of around 1 °C if denied access to holes (Cumming 1975). In winter, overnight and early morning temperatures approaching zero are common in arid savannas while the temperatures in occupied holes at that time of year are considerably higher (Cumming 1975). Daily temperature was close to 20 °C within Common Warthog holes in the Masai Mara National Reserve and Nakuru N. P. and temperature fluctuations within holes were markedly lower than fluctuations in temperatures outside the hole (Radke 1991b). Geigy (1955) found a temperature of 30 °C and relative humidity of 90% in a burrow close to Mofu in Tanzania. It thus seems probable that the cluster of features involving bare skin, absence of a subcutaneous fat layer, thermolability and sleeping in holes may reflect adaptations to conserving energy and water in arid habitats. Clearly, these adaptations may not be crucial in hotter, more humid areas. Physiological studies to test such a hypothesis have yet to be undertaken. Ewer (1957) characterized Common Warthogs as highly specialized ‘grass pluckers’ on the basis of the structure and wear on the incisors and her observations of them grazing short grass ‘lawns’ in

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Common Warthog Phacochoerus africanus juvenile.

Common Warthog Phacochoerus africanus juvenile myology.

KwaZulu–Natal. Their specialized bunodont molars are unlike those of other suids and with their flattened, grinding surfaces are adapted to a graminivorous diet (Ewer 1958). The variable number of incisors (Child et al. 1965) suggests that they may be dispensible and that Common Warthogs may use their lips, as do hippos, to graze. Watching the animals graze on short grass from a distance of a few centimetres has not clarified the question (D. Cumming pers. obs.). Common Warthogs change their diet seasonally and this is evident in the structure of the snout and the rhinarium. The dorsal edge of the rhinarium is relatively sharp, tough and firmly anchored to the nasals to allow them to scoop shallow depressions in hard, dry earth to unearth rhizomes – a feat that the Bushpig, with its more bulbous and flexible rhinarium, is incapable of performing (Cumming 1975). Common Warthogs are also capable of grazing efficiently on taller swards during the rains and of using their tusks to shred grass seeds when these ripen towards the end of the rains. The grinding surface of the molars is unlike that of other suids and is adapted to a highly lignified diet of grass leaves and rhizomes. The oft-repeated tale that Common Warthogs use their tusks to dig holes and root for rhizomes and tubers is untrue. The development of callosities on the fore-carpals on which they ‘kneel’ is clearly linked to grazing short grass and rooting for rhizomes. The callosities, absent in other suids, are evident in the embryo. A tame Bushpig that grew up with two Common Warthogs was occasionally seen trying to kneel in the same manner as the Common Warthogs, but once in this position it was clearly perplexed about what to do next and neither rooted nor grazed and soon stood up. Marked sexual dimorphism in the size of upper canines, or tusks, signifies their role in displays and inter-male rivalry. Linked to this is the sexual dimorphism of the supraoral and particularly the infraorbital warts.While the tusks may serve a similar function to horns in antelope, the three sets of warts (infraorbital, supraoral and mandibular) are situated so as to protect the eyes and underlying bones, tendons and muscles that are intimately associated with feeding. Warthogs are diurnal and generally emerge from burrows in the morning at about sunrise and retire at dusk. Feeding generally takes place during the cooler periods of the day in the morning and late afternoon with the mid-day period spent resting in the shade or basking out in the open if it is cool. In general, about six hours a day

are spent feeding, two hours walking and 2–3 hours resting with less than an hour spent on other activities such as drinking, wallowing and social activities (Clough & Hassam 1970, Cumming 1975). The remaining time of roughly 12 hours is spent in holes. However, times spent on different activities vary seasonally and with the reproductive cycle. Pregnant !! will spend a greater proportion of time resting up in late pregnancy and mothers suckling young in burrows during the postnatal period may spend as little as two hours a day feeding (Cumming 1975). The frequency of drinking and wallowing is strongly temperature-dependent. Foraging and Food Common Warthogs are almost entirely graminivorous, plucking green grass when this is available but also shredding grass seeds towards the end of the rains and rooting for grass rhizomes during the dry season (Cumming 1975). Reports from East Africa (e.g. Field 1970b, 1972) suggest that in some areas they may be predominantly grazers – a feature possibly linked to a dual wet season and availability of green grass through much of the year. In a recent study of the carbon and oxygen isotopes in the tissue of extant and fossil Suidae, Harris & Cerling (2002) characterize Phacochoerus as a hyper-grazer. In southern Africa a single wet season and long dry season spanning 7–8 months leads to clear seasonal shifts in diet and mode of feeding. In the first and so far only quantitative year-round study of Common Warthog diet and feeding behaviour, Cumming (1975) found that Common Warthogs grazed on at least 34 species of grass during the rains. As seed ripened towards the end of the rains they spent more time stripping seeds from the inflorescences of species such as Urochloa. During the dry season they rooted for the rhizomes of Digitaria milanjiana or Tristachya superba and occasionally Oryza barthii and Phragmites or the swollen leaf bases of Setaria species. Cumming (1975) reported 3–5 tons of rhizome (D. milanjiana and T. superba) per hectare in favourable dry season habitat in the Sengwa area – enough to support ten Common Warthogs per ha for the six-month dry season. Observed density was 0.28 ind/ha in such habitat. The only other larger mammal found using this major food resource was the Cape Porcupine Hystrix africaeaustralis. Common Warthogs are widely reported in the literature to eat fallen fruits, sedges, tubers and bulbs with the suggestion that the latter may 57

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Family SUIDAE

Common Warthog Phacochoerus africanus.

allow them to survive through droughts. At Sengwa fallen fruits formed only a very occasional dietary item and Common Warthogs were never recorded eating tubers and bulbs (i.e. underground storage organs of non-graminaceous plants) although Bushpigs, Chacma Baboons Papio ursinus and Cape Porcupines ate these. When offered ripe fallen Acacia pods, a favourite food item for many ungulates, tame Common Warthogs rejected them but a tame Bushpig readily ate them. Common Warthogs were regularly seen feeding on stomach and intestinal contents at predator kills in the Masai Mara National Reserve by Radke (1985) and have been recorded eating carrion elsewhere (Wilson 1975). They will, as many ungulates do, chew on old bones (osteophagia) and also exhibit geophagia and coprophagia (Cumming 1975, Mason 1982).The remains of a rodent were found in one of 600 faecal samples examined at Sengwa (Cumming 1975). Although not as large a problem as Bushpigs, Common Warthogs are sometimes a problem in agricultural lands, such as rice fields in Guinea-Bissau and peanut crops in E DR Congo (Vercammen & Mason 1993). Social and Reproductive Behaviour Common Warthogs are social animals, with the matriarchal sounder (family group), usually comprising an adult ! with her immediate litter and sometimes one or two female offspring of previous litters, forming the core social unit.Young "" leave, or are evicted from, matriarchal sounders at the end of their first year and form bachelor groups with other yearlings or older animals. Adult "" are usually solitary or in fluid bachelor groups of 2–4 animals. Breeding !! leave their sounder to farrow alone in holes but they may be joined after a week or two by yearling or subadult female offspring of previous litters. Durable social bonds between sows and their offspring were recorded in the Sengwa area. Where two or more related sows have bred they may join to form larger sounders of 12–18 animals, but sounder size is ultimately limited by the size of the burrows in which they sleep at night (Cumming 1975). More recently both solitary and cooperative breeding strategies within the same Warthog population have been described and analysed in terms of evolutionary pressures acting on sociality in the species (White 2008, 2010, White et al. 2010, White & Cameron 2011a, b). As in the Sengwa population (Cumming 1975), Common Warthogs in the Hluhluwe-iMfolozi Park in KwaZulu–Natal, South Africa, were dispersed in clans (White & Cameron 2009).

Home-range areas vary with locality. Average home-range areas of 24 ha in the Eastern Cape, South Africa, 171 ha in Zimbabwe and up to 600 ha in East Africa have been recorded (Somers et al. 1994, Cumming 1975, Radke 1991a, respectively). Home-ranges overlap and there is no evidence of boundaries, core areas or sleeping holes within the home-range being defended. The focus of activity within the home-range may shift seasonally, or boundaries may be extended temporally, in response to changing availability of food and water. Occasional exploratory forays beyond home-range boundaries may occur and young animals may disperse several kilometres from their initial home-range.There is no evidence of large-scale seasonal movements of Common Warthogs despite their occurrence in arid zones. Reports of daily distance travelled vary from 6 km in Uganda (Clough & Hassam 1970), to 3.4 km for matriarchal sounders in Sengwa in Zimbabwe (D. Cumming pers. obs.) and 1.6 km in a nature reserve in the Eastern Cape (Somers et al. 1994).They change sleeping holes frequently and there are usually many more holes available than there are Common Warthogs (Radke 1991b). In most parts of their range, and particularly in areas that experience a marked drop in temperature at night, Common Warthogs regularly enter holes at dusk and emerge at about sunrise with actual times of entry and emergence following the seasonal changes in day length (Cumming 1975). Holes may also be used as refuges from the rain. In the Sengwa area, few holes were used more than 70% of the time and animals frequently changed holes with the result that holes were used by more than one sounder. On occasion as many as 15 animals from related sounders, or members of larger filial groups, shared larger holes or burrows in erosion gulleys on the same night (Cumming 1975). On entering smaller and shorter Aardvark burrows, or holes that are being explored for the first time, Common Warthogs will generally approach the entrance, about turn and reverse into the hole (Geigy 1955, Frädrich 1965). Scent-marking begins in both sexes at 6–7 months of age (Cumming 1975) and is achieved by rubbing the damp patch in front of the eye (preorbital gland) or flange of the upper lip beside the tusks, on tree trunks, stumps and other solid objects. Adult boars will spray urine, and presumably secretions from the preputial gland, over the urine of !!. Marking with preorbital and lip glands is carried out more frequently by "" and particularly during fighting and during the mating season, while !! mark the edges of their home-ranges more

Silhouette of Common Warthog Phacochoerus africanus in ‘proud posture’.

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frequently (Radke & Niemitz 1989). Allo-marking is uncommon (Somers et al. 1995). There is no evidence to suggest that Common Warthogs use dung piles in marking. Intra-specific threat displays include standing tall, raising the mane and pressing the tail against the flank facing the opponent. Fighting between "" consists of highly ritualized head-on-head duels engaging the snout and tusks with sharp blows to sides of the head that are mostly taken on the warts. The mouth is closed and the sharp lower canines are seldom engaged in these encounters. However, they can be used, with typical suid sideways slashing movements, to good effect against predators. Greeting behaviour takes the form of naso-nasal or naso-oral contact and sniffing or touching areas of glandular secretion such as the upper lip and the preorbital area (Cumming 1975) and nasoanal contact between warthogs was reported by Frädrich (1975). Subsequent amicable behaviour may take the form of gentle frontal pushing, one animal placing its chin on the other’s back, and sometimes play fighting in young animals (Cumming 1975). Allogrooming, by drawing hairs through closed lips, occurs between family members and may serve to remove nits. Common Warthogs readily huddle together in cool weather and otherwise rest or lean on or against each other. Bouts of vigorous, playful sparring between littermates are frequent, particularly after wallowing. Warthogs are remarkably vocal and emit a wide range of grunts, squeaks and squeals that communicate, inter alia, alarm, threat-attack, greeting and submission. Courting "" emit a loud, rhythmic, almost explosive ‘chant de coeur’ that resembles the ‘phut-phut’ of a two-stroke engine. Estes (1991a) considers that clacking the canines together is a major component of the chant, but after watching and listening to a tame boar at close quarters (90%) of the population in this park (Hillman Smith et al. 2003, Lewison 2006). Adaptations  Common Hippos are highly adapted to their amphibious life-style. The epidermis is thin (up to 1 mm thick on the back), and as it is very well supplied with nerve endings, is very sensitive and delicate. Common Hippos reacted strongly when hit with drops of paint ejected from a medical syringe (for marking) (H. Klingel pers. obs). Slight scratches from bushes often result in bleeding wounds that heal quickly. The epidermis dries and cracks easily, so must be kept moist. The dermis is up to 60 mm thick on the back and flanks, thinner on head, neck and belly, and is composed of fibrous collagen. The skin accounts for about 18% of total body weight. Hippos have no true sweat glands. However, a viscous alkaline secretion (pH 8.5–10.5), ranging from colourless to reddish-brown, is produced by large, subdermal glands (distributed at a density of about 1/cm2). This secretion imparts a pinkish tinge to the body and has antiseptic properties, preventing infections and sunburn and helping in thermoregulation (like sweat) (Luck & Wright 1964, Wright 1964, 1987, Eltringham 1999, Saikawa et al. 2004). The skin serves to control body temperature primarily through regulation of evaporation, which is very high compared with other mammals, and is particularly high when the skin is wet with the secretion from the subdermal glands (Wright 1987).This led Eltringham (1999) to suggest that temperature control is not achieved through

1 6–10 2–5 No. of Hippopotamuses 0

11–20

21–60 0.5

1 km

Distribution of hippopotamus groups along a stretch of the Nile (near Chobe) (after Laws et al. 1975). Note solitary animals (nearly always males) scattered down both banks and large herds’ preference for shallows near the mouths of tributaries.

any mechanism akin to sweating unless the secretion functions like sweat, and that they may not be able to control the rate of water loss from the body.Whether or not the secretion serves the same function as sweat, Common Hippos maintain a constant core body temperature of around 36° C even while on land (though body temperature is certainly also controlled by using its aquatic environment as a means of cooling off) (Eltringham 1999). When dry it gives a shiny varnishlike appearance. When in the water, Common Hippos lift their slit-like nostrils up to the surface to breathe, at intervals of up to 6 min. A reflex response ensures that the nostrils and ears are closed as soon as they come into contact with water. By lifting the head above the water level, the ears, eyes and nostrils are allowing for visual, acoustic and olfactory perception. Because of the insulating properties of their thick skin, heat loss in the water is greatly reduced, the skin functioning as a ‘diving suit’. This, however, is an obvious disadvantage when moving on land as it reduces metabolic heat dissipation. Therefore, Common Hippos undertake the long excursions to their grazing grounds during the cooler night hours. Although Common Hippos spend much time in the water, they can neither float nor swim – at least there is no convincing evidence that they do in freshwater. They can even sink to the bottom fully inhaled, and they exhale under water or when they come up for the next breath. This can easily be observed in the wild. In Mzima Springs in Tsavo West N. P., Kenya, the Okavango in Botswana, and in some zoos, Common Hippos can be observed walking on the 71

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Common Hippopotamus Hippopotamus amphibius skeleton.

bottom. Depending on the depth of the water, the animals lie in sternal recumbency on the bottom, either fully under or with just the tops of their backs showing; in deeper water they stand on all fours, and in even deeper water they stand on their hindlegs.To reach the surface, they either just lift the head or they push themselves off the bottom. Shallow water for the adults is deep water for the infants who have the option of standing on their hindlegs, pushing themselves up for breathing, or of sitting on their mothers’ backs, apparently floating like a cork when the mother is submerged. It is quite possible that Common Hippos are able to swim in seawater, and that would explain their former appearance on Madagascar, Cyprus and Malta as well as on islands off the African coast. As the sea levels were lower during the Pleistocene than at present, the distances to be covered would have been considerably shorter. It is still a remarkable achievement for Common Hippos not only to have reached Madagascar, but also to have established viable populations there for tens of thousands of years. This achievement is even more impressive when one realizes that none of the many other ungulate species of Africa that are good to excellent swimmers ever managed to make it across to Madagascar. The colonizing effort and success of Common Hippos on dry land is also quite remarkable. Examples are the population of the Basodesh– Dongobesh–Hanang craters in the driest part of Tanganyika and of the Ngorongoro population. The founders must have negotiated their way over long and unknown stretches of waterless terrain, and for Ngorongoro they climbed the outer crater slope and down the inner one. A journey of ca. 1800 km over a period of three years over unknown country was well documented in the South African press when, in the late 1920s, Huberta the Hippo travelled from L. St Lucia in KwaZulu–Natal to near Port Elizabeth (Chilvers 1931). Common Hippos have a graviportal skeleton, being designed specifically to support their great weight. The bones of the skeleton are massive, particularly the spinal vertebrae, although the limb bones are not as heavy as may be expected in such a large animal (presumably because much of the weight of the Common Hippo is supported by water) (Eltringham 1999). When at ease, locomotion on dry land is a slow walk, each step being accompanied with a nod. When fleeing or attacking, Common Hippos perform a speedy trot and can exceed 30 km/h. Indeed, despite their heavy build, they are swift and agile and are able to climb steep slopes. Under water they move as on land except for the head nod, but, in addition, they can perform a series of forward jumps, pushing themselves up from the bottom, breaking the surface and diving in again, all in a seemingly dolphin-like manner and attaining impressive speeds that have not yet been timed.

Common Hippos spend most of the daylight hours resting and most of the night feeding. Activity pattern is influenced by weather. In dry, hot weather most animals come on land in the late morning to rest for several hours in the sun on the foreshore. In cool weather, and when it rains, they either stay in the water or come out to feed near the shore. At St Lucia in South Africa, Common Hippos, especially the subadults, bask mostly during the winter months when the water is coldest (R.Taylor pers. comm.).Where they are disturbed, generally outside protected areas, they stay in the water throughout the day. At dusk they leave the water and walk inland to their grazing areas, which can be up to 10 km from the shore (Kingdon 1979, H. Klingel pers. obs.). During the dry season when the grass is dry, coarse and scarce, Common Hippos often feed throughout the night and even stay on for some hours in the morning before returning to the water. When during the rains food is abundant, the Common Hippos often take a rest of up to several hours in the grazing area, lying down under bushes, or they even return to the water to rest there, and then walk back to the grazing area for another helping. During the wet seasons, in the Queen Elizabeth N. P., a portion of the population emigrated from the lake-shore and settled down in wallows within the grazing areas.When the wallows dried up the animals returned to the resting places from where they had departed. The shape of the muzzle is adapted to feeding unselectively, in a lawn-mower-like fashion. The grass is plucked with the horny lips, not with the front teeth, and torn off. The head is swung from side to side during the process. This is a rare feature, the only parallel being the White Rhino Ceratotherium simum. The stomach consists of four major compartments. The two anterior ones are diverticula and are fermentation chambers as is the next (third) chamber, the fourth serving for gastric digestion (Crisp 1867,Arman & Field 1973, Clemens & Maloiy 1982). The first three chambers are lined with papillae, the fourth is glandular. The anatomical and functional similarities with the stomach of ruminants are obvious, and Common Hippos are often referred to as pseudo-ruminants (Arman & Field 1973, Van Hoven 1978, Eltringham 1999). Ciliate protozoa have been identified as symbionts (Van Hoven 1974). A gall bladder is present (Ganzberger & Forstenpointner 1995). Foraging and Food  Common Hippos are predominantly grazers, although isotope studies show that many Common Hippos have a significant (more than ca. 15%) browse component in their diet (Cerling et al. 2008). Lists of species of food plants, seasonal differences in availability and preferences are given by Field (1970a, 1972); in this study, grasses consistently taken included Bothriochloa sp., Bracharia decumbens, Chloris gayana, Cynodon dactylon, Heteropogon contortus, Hyparrhenia flipendula, Sporobolus pyrimidialis and Themeda triandra. Dicotyledons are eaten accidentally where they are interspersed in the grass. In extreme situations Common Hippos feed substantially on woody plant browse (in Queen Elizabeth N. P. especially from Cassia sp.) and the fruit of some trees, such as the Sausage Tree Kigelia pinnata, are taken as well (Ansell 1965, H. Klingel pers. obs.). Aquatic vegetation is usually rare in Common Hippo waters and thus rarely taken, except locally for the Nile Cabbage Pistia stratioides, which, in the Queen Elizabeth N. P., is taken in small quantities during the dry seasons (Field 1970a, H. Klingel pers. obs.). In St Lucia, KwaZulu– Natal, Common Hippos eat fairly large quantities of Water Lilies

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Nymphaea cerulean and the submerged macrophytes Ruppia cirrhosa and Potamogeton pectinatus (R. Taylor pers. comm.). On the plains of L. Rutanzige in Virunga N. P., DR Congo, Mugangu & Hunter (1992) found that when dry season grasses failed to yield sufficient crude protein, Common Hippos were observed to feed on aquatic vegetation. According to a spatially explicit individual-based foraging model (Lewison & Carter 2004), Common Hippos appear to employ foraging strategies that respond to vegetation characteristics, such as vegetation quality, as well as spatial reference information, namely distance to a water source. Because of their wide, square muzzle, they feed unselectively with respect to individual plant species, but they do select the swards to a certain extent. Grazing grounds are used in an irregular pattern, and they also change with the quality and quantity of available food. The weights of stomach contents are remarkably low and have been determined at ca. 1% (dry weight) of body weight, which is only 50% of that of other large herbivores.The stomach contains two nights’ intake, and daily intake is in the order of 20 kg dry weight (Field 1970a, Arman & Field 1973, Eltringham 1999). Where they occur in high densities, Common Hippos shape the environment. Olivier & Laurie (1974) recognize a cyclical ecosystem in which Common Hippos play the key role.The heavy grazing pressure reduces the amount of combustible material, thereby suppressing grass fires and enhancing the spreading of bushes. This in turn will lead to a reduction of grazing area and eventually to a reduction in Common Hippo numbers and density. Reduced grazing will result in higher grass growth, fires will spread, bushes and trees will be burnt and destroyed, grassland expand and Common Hippo numbers increase again. In swamps, the paths of the Common Hippos can alter the pattern of water flow, like on the banks of L. St Lucia, and thereby have an important ecological influence (R. Taylor pers. comm.). In extensive swampy areas, like the Okavango, the movements of Common Hippos create and maintain deep, clear channels through the reedbeds and permit freer movement of the water. In backswamp areas, pathways lead to the development of new channel systems during channel avulsion (McCarthy et al. 1998) (see also Deocampo 2002). Exceptionally, Common Hippos have been observed to kill antelope such as Impala Aepyceros melampus and Common Wildebeest Connochaetus taurinus and livestock (e.g. La Hausse de Lalouvière & Wood 1989, Dudley 1996, 1998, Estes, in Eltringham 1999), and also to scavenge from them and even from the cadavers of conspecifics. The likely explanation is mineral deficiency in the ‘normal’ diet (Eltringham 1999). Licking the skin of conspecifics (R. Taylor, pers. comm.) may have the same cause. Young animals have the habit of eating dung of conspecifics; this also occurs in other ungulates and is considered to be the mechanism for transmitting intestinal symbionts. Adults are also known to feed on elephant dung, suggesting that partially digested plant material constitutes an important forage resource for Common Hippos during dry periods (Dudley 1996). By defecating in the water Common Hippos fertilize rivers and lakes. They are thus causal for the vast harvests of fish, for example, of Lakes George and Edward in Uganda and DR Congo. Social and Reproductive Behaviour  Except where noted, the following account is based on the author’s investigations from 1974 to 1979 in the Queen Elizabeth N. P., with some follow-up in the same area and in other parts of Africa, namely Mara R., Kenya;

Okavango Delta, Botswana; and Luangwa and Kafue Rivers, Zambia (Klingel 1979, 1989, 1991, H. Klingel pers. obs.). In two study areas in Queen Elizabeth N. P., more than 200 Common Hippos, the majority of the adult and subadult residents and including all key individuals, could be identified from various natural marks like scars, cuts in the ear pinna, missing ear pinnae, damaged or missing tails, colour and colour pattern. Seven Hippos were immobilized and marked with ear tags, streamers and paint, and 20 more were sprayed with paint for recognition at night. The social organization of the Common Hippo is based on mating territoriality. Although true dominance relationships do not seem to exist in Common Hippo groups, some of the adult bulls, of the order of 10%, occupy territories in which they are dominant over all conspecifics and where they have exclusive, but not unchallenged, mating rights.Among the other group members, vague and anonymous dominance relationships roughly correlated with size can be observed, but there was no evidence for individualized dominance. Territories extend along the shorelines of lakes and rivers in the water and include a narrow stretch of the bank.The sizes of territories vary considerably: in the Mweya Peninsula study area they measured from 250 to 500 m along the shore of L. Edward, while in the Ishasha R. study area only 50–100 m. In Ishasha, they included both banks of the river; in Mweya the outer, lake-ward boundary was not defined as there were no neighbours. In swamps, the territories were arranged in a mosaic pattern. Bulls keep their territories for long periods of time, and they never give them up voluntarily. But occupancy is always intermittent as the bulls, like the rest of the population, emerge at night to feed. Actual tenure is considered to depend on the number of competitors, chance and physical strength. In Mweya, four of six territories were occupied by the same bulls for the whole period of observation, i.e. 4.5 years. Two of the bulls were still territorial in their original territories in September 1982, almost eight years after they were first recorded, and one of them even after 12 years. In the Ishasha study area, recorded tenure was from a few months to over two years, i.e. throughout the investigation, changes in territory boundaries, takeovers and the establishment of new territories in previously unoccupied places could be documented. From the available data, the maximum tenure can be estimated to span the whole adult life of a bull, i.e. in the order of 20–30 years. Size of the territories, frequency of takeovers, boundary changes and length of occupancy seem to be inversely correlated with density. In Mweya, density was only 7 animals/100 m shoreline, whereas in Ishasha it was 33 animals/100 m. However, other factors are important as well. In Ishasha, some of the territorial changes were caused by changes of the course of the river, the water level and current. During the study one branch of the river fell dry and, consequently, the whole territorial set-up changed, as two out of seven bulls gave up and moved out of the area, and the others rearranged their territories. Later, during a flood, all the Common Hippos left the area. Some settled in a dead meander of the same river, others invaded a newly formed swamp, where they immediately started to establish new territories. Territories are advertised by the bulls through their presence, their dominant behaviour and their ritualized defecation combined with urination. By rapidly wagging their tails, faeces and urine are scattered in the vicinity. This type of defecation is repeated in the 73

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Common Hippopotamus Hippopotamus amphibius.

same places and results in impressive dung heaps, measuring several square metres in area. The dung heaps certainly do not function as boundary markers and they do not prevent other bulls from entering the territory, but they are thought to serve as orientation marks for the territorial individual as well as for other Common Hippos. Dung heaps are not produced by the territorial animal alone, but by virtually all passing ??, and they are also found outside the territories in the grazing areas and along the inland tracks. They are rarely found in the open grassland but are regularly produced and maintained at the edges of bushes, at narrow passages between bushes, sometimes at corners of houses. Territorial neighbours often display ritualized simultaneous defecation at their common boundary in the water. They then stand side by side a few metres apart facing in the same or opposite direction, both demonstrating strength and dominance by holding their heads high and ears forward. In this situation it is quite clear that the behaviour serves as a visual signal, as dung heaps are not produced. Smell is likely to be of importance as well. When a territorial bull moves across the boundary of his territory, he loses his status and behaves subordinately to the territorial neighbour. Fights between territorial neighbours are generally ritualized frontal combats combined with splashing of water, which serve to demarcate boundaries. Often, the combatants hardly touch each other, advancing and retreating in response to the opponent’s actions. However, serious fights occur for the possession of a territory or part of it. Then the bulls fight standing parallel facing in opposite directions and slashing with their lower canines at each other’s flanks. Though the hide is several centimetres thick, these attacks can result in serious injury and even death. In these fights the combatants are

usually a territorial defender and a non-territorial challenger, but occasionally two territorial neighbours are involved if one tries to extend his territory into the other’s area. Territorial owners are extremely tolerant of all conspecifics, including adult bulls as long as these behave subordinately. Tolerance goes to the extent of the territorial bull hosting groups of 100 and more fully mature, prime bachelor bulls in the territory. In this situation the function of mating territoriality becomes absolutely clear – only the territorial individual has access to any cow in the territory. The relations between a territorial bull and subadult bulls in his territory are of an extremely friendly nature.The subadults demonstrate their subordinance in an elaborate display: on land they approach from 100 m or more away, head down, the last few paces virtually crawling on the ground, then sniffing the bull’s genital region from the side or from behind without him seeming to take notice. A modified version of this behaviour is displayed in the water. Nasonasal contacts are common. The subadult bulls often stay for hours in the immediate vicinity of the territorial bull, and there is not the slightest indication of antagonism. Rarely, during the submissive display, the territorial bull defecates on the head and back of a subadult. It is not yet clear if this is incidental or if it has a functional significance. During the day, Common Hippos live in social groups of variable size and composition (Olivier & Laurie 1974, Delvingt 1978, Klingel 1979, 1989, 1991, Karstad & Hudson 1986, Viljoen 1995). Solitary animals are generally older ?? or // about to give birth. Group size is not a sociological parameter, but rather depends on density and the environmental situation. For example, Common Hippos avoid fast currents and prefer shallow waters with sandy, open foreshores. R. Taylor (pers. comm.) measured 1.3– 1.5 m as the preferred water depth for lying up during the day. In such situations the larger groups counted were over 200 animals strong, as in the Ishasha R. (Klingel 1991). Two types of social groups can be distinguished: nursery schools consisting mainly of // and their young, and bachelor groups consisting mainly of bulls. However, the distinction is not clear-cut, and there are often odd members of the other sex in a group. Neither of the groups is a stable social unit but both are loose associations in suitable resting places and the animals are socially but anonymously attracted to each other. Indeed, the existence of individual recognition among adults cannot be proved and it certainly does not play an important role in Common Hippo social life. However, mother and offspring do recognize each other individually, and mother–calf bonds are the only stable associations, usually lasting until the young are almost fully grown at 6–8 years of age. In the evenings the groups break up and the animals walk singly or in mother–offspring units to their grazing areas. The latter are not monopolized or defended by an individual or group of individuals (i.e. they are not subdivided into territories). Members of a school use an exit near their resting site, or they move along the shore in the water to a place closer to the grazing grounds.They stick to a network of tracks that form through repeated use over the years. At exits the tracks can cut a metre or more into the bank, and on steep inclines tracks erode to become steep gorges that, in the Queen Elizabeth N. P., are characterized by their special plant, bird and small mammal communities. Common Hippo trails can often be recognized from a grassy, belly-smoothed strip in the middle. Regular paths are also used in shallow lakes.

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After their nocturnal feeding trips the animals return preferentially to the sites from which they have departed the previous evening, resulting in a certain constancy of group composition. But each individual has, independently of the group, its own home-range and several resting places. In one study area, Mweya Peninsula, the majority of the known animals were located for years in the same day home-ranges of 100–200 m in length. Home-ranges do not coincide with the territories, nor are they restricted to a particular territory. Depending on territory size, a / can be at home in one or in several territories. Changes in group composition also occur during the day. Frequency depends on the spacing of the groups but also on other factors. Some individuals were particularly mobile and, possibly out of curiosity about the observer, their movements were correlated with his, and they moved parallel to the shore in the water, crossing territories and passing groups of conspecifics without any sign of antagonism and without any attempts by the bulls to prevent their departure. Generally, Common Hippos are quite philopatric and stay within their regular home-ranges. In Queen Elizabeth N. P., a portion of the residents, especially non-territorial ??, used to settle in wetseason pools, utilizing the grazing area nearby. At the beginning of the dry season they would return to where they had come from. When their rivers and wallows dry up, they are eventually forced to move, occasionally over large distances, to find suitable water bodies. Matings take place exclusively in the aquatic part of territories. Accordingly, for the ?, the possession of a territory is an absolute prerequisite for reproduction, although it is by no means a guarantee. Some bulls maintained territories in areas that were unattractive to // and they probably had no access to cows. However, in the Ishasha R. several ‘bad’ territories changed to ‘good’ ones when the course of the river and the water level changed and consequently some // moved in. Other territories changed for the worse. It is therefore a better strategy for a bull to occupy a bad territory than to be a non-territorial bachelor. Oestrus lasts for 2–3 days and up to six copulations occur per day. Although the // tend to return in the morning to the place and thus to the territory from which they had departed the previous night, they are promiscuous and likely to visit two or more territories in succession and mate with the respective owners. A few days before giving birth to a single young, the cows separate from the group. When the cow is undisturbed, birth is in shallow water at the waterline; when she is disturbed, birth can take place in deeper water. The young are precocious and within a few minutes after birth able to suck even under water and to follow their mothers into deeper water. Eltringham (1999) suggested that milk is injected into the calf’s mouth by muscular action on the part of the mother, as occurs in some cetaceans. However, young use their tongue and mouth roof around their mothers’ nipple while actively sucking, but not the lips, which are inflexible (Kingdon 1979). Mother and infant stay away from conspecifics and prefer to rest in close proximity at the edge of the water, the mother obviously without eating for several days. For about ten days she is intolerant of all conspecifics and attacks even the territorial bull and her own elder offspring should they approach. This behaviour, which is also observed in other ungulates, is considered to prevent false imprinting. It also serves to protect the infant from attacks by conspecifics. After this period, the mother will graze on the bank during the day with

the infant resting nearby. After several weeks mother and calf will walk, at night, to the grazing grounds where the infant will be parked under a bush whilst the mother feeds. She eventually returns, collects the calf and they return to the water together. Occasionally, calves aged 6–12 months stay behind in a crèche near the water whilst their mothers depart on their nocturnal foraging excursions. The birth of a sibling does not disrupt existing bonds, and // with several young, up to three, of different ages, can be observed. The marching order is by age, the youngest first, directly behind the mother. Numerous cases of infanticide have been reported, some directly observed, others based on circumstantial evidence, and are attributed to ??. However, in a recently filmed episode the infanticidal individual was clearly a subadult, of undetermined sex (M. Deeble & V. Stone pers. comm.). Verheyen (1954) considered adult ?? to be the major enemies of infants. Lewison (1998) analysed the evidence and concluded that infanticide in Common Hippos, as has been demonstrated for Lions, may be a reproductive strategy by which a new territorial ? kills his predecessor’s offspring to shorten the inter-birth intervals of the / and thus enhance the male’s reproductive success. However, the evidence is only circumstantial. In none of the cases was a change of territorial ownership and/or the status of the infanticidal individual observed. As all or most incidents occurred in the dry season, high densities and nutritional stress may have triggered the behaviour. Common Hippos command a rich repertoire of signals and means of communication, the actual significance of which is only partly understood. The impressive gape signals strength and presence; it is, however, very similar to the yawn, which is of shorter duration, and conspecifics do not seem to be impressed by either. Also, the yawn/ gape is displayed by all members of a group including infants, and it occurs with highest frequency just before they go on land in the evenings – suggesting that the yawn/gape may be an expression of excitement, and not a threat (Kingdon 1979, H. Klingel pers. obs.). When attacking, Common Hippos have their mouth only partly open, and this is clearly a threat that is understood by conspecifics. Head up signals dominance, head low, lip smacking and tail wagging with or without defecation are signals of submission, occurring in minor confrontations and by the / during pre-mating activities (Karstad & Hudson 1986, H. Klingel pers. obs.). Behaviour analogous to ‘flehmen’ has only recently been observed to occur in Common Hippos and probably has no communicative value (Zapico 1999). Several vocalizations can be distinguished, but interpretations of their significance are still largely speculative. The most common consists of a series of guttural honks preceded and followed by a high-pitched squeal. It is contagious, being started by one individual, repeated by others, and within seconds all members of a group, and soon the neighbours as well join in a cacophony that carries over several kilometres. Calls are individually different and may serve for individual recognition. Sometimes mother and young call and answer after having lost contact while on their walk to the grazing grounds. Other individuals occasionally utter this call at any time of day or night, and they sometimes get answers from far away, but often not (H. Klingel pers. obs.). When fighting and in distress, Common Hippos produce long high-pitched squeals. Barklow (1997) recorded and analysed three types of underwater sounds that are rarely audible on the surface: a whine signalling submission; a croak used by young; and clicks for communication (and see Barklow 2004). When facing 75

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each other they often make a low-pitched loud rumbling noise (R. Taylor pers. comm.). Many bird species, especially waders, use Common Hippos for perching; some run up and down the animals’ backs hunting insects and ticks (e.g. African Jacanas Actophilornis africanus), others catch insects that the Common Hippo has disturbed in the water vegetation (see Eltringham 1999 for summary). Hediger (1951) described a case of mutualism between Common Hippos and the Cleaner Fish Labeo velifer. M. Deeble & V. Stone (pers. comm.) have observed and recorded on film that Common Hippos actually invite the fish to clean any area of the body, including the mouth cavity, and that they visit cleaning stations where the fish are waiting. Erwee (1996) observed a Common Hippo saving an Impala Aepyceros melampus from drowning, by pushing it ‘carefully’ onto dry land, and there are other instances of Common Hippos displaying similar behaviour, including intervening in attacks on Impalas and other antelopes by crocodiles or large predators. Common Hippos are said to be responsible for more human deaths than any or even all other dangerous wildlife species combined. The truth is difficult to assess as there are no reliable statistics. Common Hippos are certainly dangerous, especially when cornered and wounded or provoked. Most of the casualties are of fishermen who fatalistically drive their boats right through the Common Hippo schools, oblivious to the danger. If the boats are attacked the fishermen will jump out, or may fall into the water, and drown because they generally cannot swim; they may also get bitten by the Hippos and may lose a limb or a life. On land, dangerous situations for humans arise when bumping into a Common Hippo, especially at night, and when inadvertently getting between a mother and her young. Normally, the relations between Common Hippos and people are quite relaxed, with animals idling in the water within as near as 20 m of people bathing or washing their clothes (H. Klingel pers. obs). In much of their range crop-raiding by Common Hippos is a big problem. Chain-link fencing is typically ineffective at excluding cropraiders, especially where it cuts across well-used tracks, because the animals simply trample the fences. Likewise, single strands of electrical wire are only partly effective, as some of the animals, upon contact, rush forward and break the wire. One solution involves constructing a simple barrier like a wooden rail or a shallow ditch about 50 cm in front of and parallel to the electric wire. Both obstacles help slow the animal’s pace, thereby serving as a warning before the animal touches the high voltage wire. In KwaZulu–Natal, fields are successfully protected by 1.2–1.5 m deep trenches (R. Taylor pers. comm.). Reproduction and Population Structure  Detailed infor­ma­ tion on breeding and population structure is available from culling schemes, in particular from Queen Elizabeth N. P. (Laws & Clough 1966), the Luangwa R. in E Zambia (Sayer & Rakha 1974, Marshall & Sayer 1976) and South Africa’s Kruger N. P. (Smuts & Whyte 1981). Average age of sexual maturity in ?? (based on testes weights) is 7–8 years (Laws & Clough 1966, Sayer & Rakha 1974, Smuts & Whyte 1981), although spermatogenesis may begin much earlier (as early as two years of age in animals from Kruger; Smuts & Whyte 1981). Based on follicular size, // reach sexual maturity at about seven years, although some // examined have been found to be sexually mature at much younger ages, including one at three years in Uganda (Laws & Clough 1966, Sayer & Rakha 1974). The age

female

male Common Hippopotamus Hippopotamus amphibius.

at which all // are pregnant varies between the three regions in which major studies on reproduction have been conducted: 11 years in Kruger, but 20 years in Uganda and Zambia, indicating different degrees of crowding and nutritional stress (Eltringham 1999). Captive animals may breed at an earlier age, and Dittrich (1976) recorded one / conceiving at only 2 years and 3.5 months. Males are sexually active throughout life, their body weight continuing to increase and testis continuing to grow. Births are recorded any time of the year; however, there is pronounced seasonality, and most births in Uganda (close to the equator) occur during the rains (Laws & Clough 1966). In Zambia, conceptions occurred throughout the year, but births only during the wet season (Marshall & Sayer 1976). Gestation lasts for about 240 days, and usually a single calf, weighing a mere 50 kg, is born (Laws & Clough 1966, Smuts & Whyte 1981, Eltringham 1999). Twins occur infrequently; in the sample of 276 culled specimens examined by Laws & Clough (1966), twins were recorded twice. Calving intervals are 2–3 years (see Smuts & Whyte 1981), and oestrous cycle is given as around 50 days (Laws & Clough 1966, Smuts & Whyte 1981, Eltringham 1999). There is some evidence of a postpartum oestrus, as 25% of // examined by Laws & Clough (1966) were both pregnant and lactating. Conception rates were close to 40% in Kruger N. P. (Smuts & Whyte 1981) and 27% in Queen Elizabeth N. P. (Laws & Clough 1966). Percentage of lactating // in Uganda was almost 60% (Laws & Clough 1966), and 78% in South Africa (Smuts & Whyte 1981). Weaning probably takes place between six and eight months, with most calves weaned by 12 (Eltringham 1999); however, calves as young as six to eight weeks often have considerable quantities of grass in the stomach (Laws & Clough 1966). Sex ratio is 1 : 1 in foetuses (Bere 1959, Laws & Clough 1966, Smuts & Whyte 1981), infants and young (H. Klingel pers. obs.). Reported adult sex ratios are likely to be inaccurate due to the segregation of sexes. Smuts & Whyte (1981) reported a ratio of 1 : 1.93 in Kruger N. P. (n  = 463), while Marshall & Sayer (1976) recorded 1 : 0.98 in 1970

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(n = 375) and 1 : 1.56 in 1971 in Zambia (n = 210), the difference in the proportion of ?? due to a greater effort during 1970 to take a more random sample. Laws & Clough (1966) found a nearly exact sex ratio in Uganda, and confirmed that there was a skewed distribution of sexes according to habitat: in small wallows, 313 ?? and 110 //, and on lake shores, 600 // and 821 ??. Population structures of Common Hippos from culling operations exist (e.g. Marshall & Sayer 1976), although samples are biased given the tendency for younger animals not to be culled (and see Eltringham 1999). Longevity is recorded at 61 years in captivity (Wiesner & von den Driesch 1996).

Common Hippopotamus Hippopotamus amphibius juvenile mouth showing adaptation of tongue and upper lip to suck maternal nipple.

Predators, Parasites and Diseases  Lions Panthera leo succeed occasionally in bringing down an adult Hippo, but they have no impact on the population. Also, young animals are preyed upon by Spotted Hyaenas Crocuta crocuta and Lions (Kingdon 1979). In one attack by five hyaenas on an infant, at night, the cow’s defence was quite inadequate: she repeatedly chased individual hyaenas that came close, but in the process she again and again left the calf unprotected. The hyaenas took turns in attacking the calf and fleeing from the mother and they eventually managed to bring the calf down. The mother stood for several hours over the body and protected it from being eaten, with the hyaenas sitting nearby. Eventually, after dawn, the mother walked the short distance to the water and the hyaenas had their meal (H. Klingel pers. obs.). Nile Crocodiles Crocodilus niloticus are potential predators of Common Hippos, especially young ones, but no records are known. Most observers report Common Hippos to be dominant to crocodiles (see Kofron 1993). In the wild, Common Hippos are susceptible to rinderpest, a viral disease that was introduced to Africa towards the end of the nineteenth century and which decimated large numbers of domestic and wild ungulates. It is not known if Common Hippos actually succumbed to the disease in any numbers, but they developed antibodies (Plowright et al. 1964). Laws (1968) correlated the presence of rinderpest antibodies in culled Common Hippos with

dated outbreaks of the disease and could thereby construct an ageing scale based on tooth eruption and wear, which is one of the most accurate for any wild animal species. A major and often deadly disease of Common Hippos is anthrax, caused by Bacillus anthracis. Outbreaks have been reported repeatedly from Uganda and Congo/DR Congo during the past 40 years, though they were not necessarily diagnosed correctly. One identified outbreak, in 1978, resulted in high mortality in some areas of Queen Elizabeth N. P., but had little impact on the population as a whole (H. Klingel pers. obs.). An outbreak in Zambia in 1987 resulted in the death of >4000 Common Hippos out of a population of 20,000 (Turnbull et al. 1991). In 2004, anthrax outbreaks were reported from Zambia and Uganda, the latter claiming about 300 Common Hippos (C. Tumwesigye pers. comm.). Other disease organisms afflicting Common Hippos are Brucella abortus (which causes brucellosis) and several species of Salmonella and Trypanosoma. Major parasites include flatworms (liver flukes Fasciola nyansae are found in the livers of most Common Hippos, while one species of blood fluke, Schistosoma hippopotami, is host-specific), nematodes, and ectoparasites, including ticks and a fluke Oculotrema hippopotami that lives on the outer surface of the eye (see Du Preez & Moeng 2004). See Eltringham (1999) for an overview. Conservation  IUCN Category:Vulnerable A4cd. CITES: Appen­dix II. Today Common Hippos are present in numerous protected areas across the continent, with important populations in Niokolo-Koba N. P. (Senegal), National Park of Upper Niger (Guinea), Comoé N. P. (Côte d’Ivoire), W N. P./Arly/Pendjari (Benin, Niger, Burkina Faso), Zakouma N. P. (Chad), Queen Elizabeth and Murchison Falls National Parks (Uganda), Salonga, Upemba and Virunga National Parks (DR Congo), Serengeti, Ruaha, Mikumi and Tarangire National Parks, Ngorongoro Crater Conservation Area and Selous G. R. (Tanzania), Amboseli, Lake Nakuru and Tsavo National Parks and Masai Mara and Samburu National Reserves (Kenya), Kasanka, Kafue, North and South Luangwa and Lower Zambezi National Parks (Zambia), Kasungu and Lake Malawi National Parks (Malawi), Hwange and Mana Pools National Parks (Zimbabwe), Chobe N. P. (Botswana), Gorongosa N. P. and Niassa G. R. (Mozambique) and Kruger N. P. (South Africa). However, across their range, and in particular in West Africa, many populations of Common Hippos are decreasing due to droughts, poaching and to habitat loss and habitat fragmentation from changing land use and damming of rivers (Eltringham 1993a, 1999, Lewison 2007, Zisadza et al. 2010).Two very large Common Hippo populations suffered particularly heavy losses through intense commercial poaching by military and civilian poachers. In Uganda, Common Hippos of Queen Elizabeth N. P. declined during the war and unrest of the late 1970s to 1980s from about 20,000 to 2000 (Eltringham 1999). Since then, an increase to only 2600 in 2004 has been recorded (C.Tumwesigye pers. comm). More recently, in DR Congo, an even more dramatic decline has been reported from the Virunga N. P. population, which, during the recent civil war and turmoil, declined by about 97%, from close to 30,000 in the 1970s (Delvingt 1978) to 1300 in 2003 (A. Plumptre pers. comm.) and to perhaps less than 400 in 2006 (Lewison 2006). Hunting pressure is presently mostly for meat; however, there is increasing demand for tusks, the ivory of which is said to be softer and easier to carve than elephant ivory. 77

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developed, and the responses of the animals are highly variable (H. Klingel pers. obs.). M.Woodford (pers. comm.) has suggested that the parasite load of the animal might be responsible for the inconsistent effects. In South Africa, Common Hippos are captured by pushing them into a funnel leading directly onto a lorry (R. Taylor pers. comm.). Kingdon (1997) sees much potential in Common Hippos being domesticated as a meat source. The dung also could be made use of in fish farms, thereby producing additional protein. Measurements Hippopotamus amphibius HB (??): 3120 (2600–3500) mm, n = 54 HB (//): 2990 (2590–3370) mm, n = 156 Sh. ht (??): 1500 (1290–1720) mm, n = 32 Sh. ht (//): 1440 (1100–1580) mm, n = 36 WT (??): 1546 (955–1999) kg, n = 86 WT(//): 1385 (995–1850) kg, n = 192 Kruger N. P. (Pienaar et al. 1966, I. Whyte, pers. comm.) Laws (1963, in Eltringham 1999), in Uganda, found the average weight to be 1536 kg for ??, with a maximum of 2065 kg, and 1386 kg for //, with a maximum of 1716 kg. Ledger (1968) reported an average of 1490 kg (range 1179–1714 kg, n = 4) for ??, and 1277 kg (range 1185–1401 kg, n = 4) for //. Based on the samples of 189 ?? and 186 // in the Luangwa Valley, // appear to approach an asymptotic weight at around 24 years of age, while ?? continue to grow throughout life (Marshall & Sayer 1976). The record tusk length is 1638 mm for animal from Congo (Rowland Ward).

Common Hippopotamus Hippopotamus amphibius seated.

An important conservation measure is the translocation of Common Hippos, although immobilization of animals, for research and/or for translocation, poses problems. It has to be done at night in the grazing grounds, to make certain that the drugged animal does not reach deep water and drown. Also, no safe drug combination has been

Key References  Eltringham 1993a, 1999; Kingdon 1979, 1997; Klingel 1979, 1989, 1991; Laws & Clough 1966; Marshall & Sayer 1976; Smuts & Whyte 1981. Hans Klingel

Genus Choeropsis Pygmy Hippopotamus Choeropsis Leidy, 1852. J. Acad. Nat. Sci., Philadelphia 2 (1): 207.

There is only one living species of Choeropsis, the ‘Pygmy’ or Liberian Hippopotamus, C. liberiensis. It is now restricted to lowland forests of Côte d’Ivoire, Sierra Leone, Liberia and Guinea, but in the recent past, it probably also populated coastal regions east of these countries as far as the Niger Delta. The Pygmy Hippopotamus (hereafter Pygmy Hippo) is similar in appearance to the well-known Common Hippopotamus Hippopotamus amphibius, but it is much smaller than the latter, its back slopes forward, and its eyes, ears and nostrils are situated lower down on the cranium. Its general proportions are also different, with relatively longer limbs and a proportionately smaller, narrower head. Osteologically, it differs by its sagittal crest sloping backward, its lachrymal separated from the nasal by an anterior extension of the frontal, its low-crowned cheekteeth and the presence of only two incisors on the mandible. Fossils of actual pygmy hippopotamids are known from several islands, including Cyprus, Malta, Crete and Madagascar. Continental

small hippos were discovered in the Neogene of eastern Africa: Hexaprotodon ? imagunculus in the western branch of the Great Rift Valley; aff. Hippopotamus aethiopicus and Archaepotamus lothagamensis in the eastern branch. For those continental species, small size probably permitted the use of ecological niches unsuitable for the abundant and much larger species found in the same deposits. In the case of Choeropsis, small size could, in fact, be a primitive condition (the oldest known hippos being about the same size or somewhat smaller) and/or could be an adaptation to its habitat, unusual for a hippopotamid. Indeed, very large size is not particularly beneficial for moving through dense forest vegetation and must be sustained by an abundant, nutritious and reliable food supply. Many peculiar traits of Choeropsis can in fact be interpreted similarly: its forward sloping back perhaps facilitates passage through dense vegetation, whereas in these conditions low orbits could offer better protection for the eyes. Furthermore, much of this vegetation is woody and

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Pygmy Hippopotamus Choeropsis liberiensis.

not especially nutritious. Choeropsis shows the same sun-sensitive skin histology as Hippopotamus, but can venture outside water more frequently in its shady and humid environment, and is less aquatic than its larger extant relative. The Pygmy Hippo was initially classified in the genus Hippopotamus by Morton (1844), then in its own genus (Leidy 1852). In 1977, on the basis of separation between the lachrymal and nasal bones, Coryndon (1977) merged Choeropsis into the genus Hexaprotodon including most fossil species. However, this morphology was shown to be primitive in the family (Harris 1991a), whereas Harrison (1997) favoured generic distinction of the Pygmy Hippo because of its peculiar overall anatomy. Accordingly, a recent review of the taxonomy and phylogeny of the hippopotamids has restricted the definition of Hexaprotodon and thus revalidated Choeropsis for the extant Pygmy Hippo (Boisserie 2005). Choeropsis has no known fossil record; its mosaic of primitive (e.g. low sagittal crest), derived (i.e. number of incisors) and unique (i.e. strong nasal spine of the palatine bone) characters among Hippopotamidae suggests that its ancestors separated from other lineages in the late Miocene (Harrison 1997, Boisserie 2005). At this time, hippopotamids underwent a limited radiation, becoming for the first time abundant and diversified in African ecosystems.

At least four main lineages emerged, including Choeropsis (Boisserie 2005). Among the three others, the first is recognized in eastern Africa and the Arabian Peninsula as genus Archaeopotamus. Its species exhibited an elongated, shallow and narrow mandibular symphysis (Coryndon 1977, Gentry 1999, Weston 2000). The second lineage, the extinct genus Hexaprotodon, was seemingly rare in Africa, but flourished in southern Asia, from India to Indonesia, and maybe in southern Europe. Its species were characterized by a robust, high and moderately wide mandibular symphysis. The third lineage, represented by the monospecific Saotherium from basal Pliocene in central Africa, has some affinities with Choeropsis. Both genera share an elongated braincase rounded in transverse section and relatively large orbits. These characters could be primitive, so may not express actual close phylogenetic relationships. In fact, the relationships between all these lineages – thus, the origin of Choeropsis – are still unclear. In any case, Choeropsis and Saotherium suggest a long and diverse hippopotamid evolutionary history in central and western Africa, despite a fossil record far more sparse than in the eastern part of the continent. Jean-Renaud Boisserie & S. Keith Eltringham 79

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Choeropsis liberiensis Pygmy Hippopotamus Fr. Hippopotame nain, Hippopotame pygmée; Ger. Zwergflusspferd Choeropsis liberiensis (Morton, 1849). J. Acad. Nat. Sci., Philadelphia 2 (1): 232. St Paul’s River, Liberia.

Pygmy Hippopotamus Choeropsis liberiensis.

Taxonomy  Two subspecies are recognized. The form C. l. heslopi, from the Niger Delta, is originally known from three skulls, one specimen shot by Heslop himself and two skulls obtained from hunters (Heslop 1945); a further two skulls were collected by J. B. I. Mackay in 1928 (Ritchie 1930). It was considered a distinct subspecies by Corbet (1969) and Coryndon (1977), based on variations in cranial anatomy. Synonyms: heslopi, minor. Chromosome number: 2n = 36 (K. Benirschke pers. comm.). Description  A secretive, nocturnal, highly solitary, pig-like quadruped resembling, but significantly differing in size, anatomy, behaviour and ecology from, the Common Hippopotamus Hippopotamus amphibius (see Similar Species). Overall colouration ranges from grey-brown to purplish-black with creamy-brown underparts. The body profile is raked; the legs less massive than the Common Hippo and the digits are more free and mobile. Nostrils and diminutive ears can be closed by muscular valves, as in Hippopotamus. The dental formula is I2/1, C1/1, P3/3, M3/3 = 34, which is reduced from the ancestral condition of 44. The incisors are peg-like and the canine teeth, which are used for defence and aggression rather than for feeding, are sharpened by occlusal wear and are open-rooted and continuously growing. The molariform teeth are low-crowned with a grinding function. Geographic Variation C. l. liberiensis: Upper Guinean lowland forests, in Guinea, Sierra Leone, Liberia and Côte d’Ivoire. C. l. heslopi (Heslop’s Pygmy Hippo): Niger Delta. Probably extinct. The Nigerian subspecies has been described as having a more horizontal dorsal profile, as contrasted with the more sloped profile characteristic of the nominate subspecies (I. R. P. Heslop pers. comm.); also differs in cranial features (discussed by Corbet 1969).

Choeropsis liberiensis

Similar Species Hippopotamus amphibius. Largely allopatric, although their ranges overlap in the upper stretches of some rivers in Côte d’Ivoire (see Roth et al. 1996, 2004) and perhaps in NW Sierra Leone (Grubb et al. 1998). Larger overall size, but with proportionally shorter limbs; nostrils and bony orbits more prominent; toes close-knit; two pairs of lower incisors. Typically solitary or in small groups, and confined to forested regions close to rivers. For a comparative review of Choeropsis and Hippopotamus see Eltringham (1999). Distribution  Endemic to Africa, where currently confined to the Upper Guinean lowland forests of Liberia, Côte d’Ivoire, Guinea and Sierra Leone.An isolated subspecies, C. l. heslopi, formerly occurred some 1800 km to the east, from the Niger Delta east to the Cross R. in Nigeria (Heslop 1945; and see Corbet 1969, Ansell 1972). Reports of this subspecies in the 1930s and 1940s were from the swamps of the Cross R. region of Calabar Province, and from swamps of the Niger R. and its tributaries in the British colonial administrative divisions of Kwale, Asaba, Onitsha and Aboada.The subspecies has not been reliably recorded in more than half a century, and its continued presence seems unlikely. Powell (1995) suggested that the only remaining possibility was in the Upper Orashi Forest Reserve. Their current distribution is more fragmented than in the past, largely due to the extensive forest loss that has taken place in the lowland rainforests of Upper Guinea. The core of the species’ range is centred on Liberia, which has the most extensive tracts of intact lowland forest in the region (Christie et al. 2007). Pygmy Hippos are also present in neighbouring E Sierra Leone (in the Gola Forest

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region), SE Guinea (including Ziama and Diécké) and SW Côte d’Ivoire (Eltringham 1993b, 1999, Grubb et al. 1998, Roth et al. 2004, Mallon et al. 2011). In Côte d’Ivoire, the most northerly occurrence of the Pygmy Hippo is between 7° 20' N and 7° 30' N in the Upper Cavally R. valley, just south of the Guinean border, and along the Upper Boan R., south of the Mt Nimba massif (Roth et al. 1996, 2004). They have been recorded from Azagny N. P. and along the Bandama R. in SE Côte d’Ivoire, just west of Abidjan (Roth et al. 1996, 2004), which is about the farthest east the nominate form has been recorded (4° 18' W). In Liberia, their distribution seems to be divided between the large remaining forest blocks in the south-east (centred on Sapo N. P.) and north-west of the country (Mallon et al. 2011). The farthest west the species appears to have been authentically recorded in the past is near Forécariah, in Kindia, Guinea (Dekeyser 1955; see Grubb et al. 1998), but there are no present-day records from further west than NW Sierra Leone (Grubb et al. 1998) where it is possible they may survive in Outamba-Kilimi N. P. in sympatry with the Common Hippo. Pygmy Hippos are also still recorded from the Loma Mts in the north of the country. Except for the reports from NW Sierra Leone (Grubb et al. 1998) and Côte d’Ivoire by Roth et al. (1996, 2004) and Hentschel (1990), such as on the Bandama R. as far north as the confluence of the Nzi R., no other locations have been verified where the ranges of Pygmy and Common Hippos currently overlap. Historically, however, it is likely that these two species did have small range overlaps; for example, in SW Liberia in the 1930s (Van Ness Allen 1939) and 100 years ago in the Junk River region in Liberia (Schomburgk 1912, 1913, 1922). Heslop (1945) reported seeing Pygmy Hippopotamus tracks within 20 feet of those of a Common Hippopotamus in the Niger Delta region. Records of the species from Gambia and Ghana were rejected by Grubb et al. (1998), while another isolated record from Guinea-Bissau (Cristino & Melo 1958) almost certainly refers to the Common Hippo. Habitat  Lowland primary and secondary evergreen forests, sometimes penetrating savanna regions along gallery forests. Prefers isolated areas with low human disturbance; riverine and swamp areas are frequented much more than upland forest sites. The habitat must offer suitable cover for resting, calving and nursing of calves. Pygmy Hippos often follow water-courses and dry-season forest gullies in their travels and do not hesitate to cross rivers and streams, usually fleeing water to seek refuge in the forest when encountered. Under-cut and eroded banks of forest streams and rivers are not uncommonly used as opportunistic retreats, a habit that is exploited by hunters (Schomburgk 1913, Robinson 1981); the latter author provides a detailed description of one such denning structure in Liberia. Seasonal climate within the species’ range is characterized by a major dry season occurring variably between Nov and Mar, with significant rainfall during the balance of the year. Abundance  The total size of the wild population is unknown, but was estimated in the thousands in the early 1990s with the bulk of the population in Liberia, and perhaps less than 100 animals in Sierra Leone (Eltringham 1993b, 1999). However, Roth et al. (1996, 2004) present evidence (see below) contradicting the traditional view that Liberia remains the only stronghold for this species. Using faecal

Lateral, dorsal and palatal views of skull of Pygmy Hippopotamus Choeropsis liberiensis.

droppings and a decay rate method, Hentschel (1990) estimated mean population densities in Taï N. P., Côte d’Ivoire at 3.6 ind/km2 in primary forest and 2.9 ind/km2 in secondary forest. Based on these density estimates, Hentschel (1990) estimated a total population of about 14,200 Pygmy Hippos in Taï N. P. and its surrounding areas, although Roth et al. (2004) caution that this was probably an overestimate. Subsequently, recent studies in the eastern part of Taï N. P. have revealed that densities decreased to 0.3 ind/km2 in 1998 and to 0.2 ind/km2 in 2001. Densities in the interior and in the western part of the park ranged from 0.8 ind/km2 in 1995 to 1.4 ind/km2 in 1998. In the northern adjacent N’zo G. R., densities of up to 2.5 ind/km2 existed in 1998. From 1999 to 2001 the Hippo population there increased to 4.1 ind/km2 (Roth et al. 2004). The latter authors concluded that the total number of Pygmy Hippos in Taï N. P. and its adjacent forest areas was certainly less than 10,000 animals, and estimated the total number in Côte d’Ivoire as probably no more than 15,000 in total. Notwithstanding errors associated with extrapolating population density estimates from dung pellet counts, these figures suggest that the total population of Pygmy Hippos is probably larger than that indicated by Eltringham (1993b, 1999). Adaptations  Compared with the Common Hippo, the Pygmy Hippo is more adapted to terrestrial locomotion. The toes of the feet are less webbed than the Common Hippo, and better suited to frequent movement in and out of rivers and streams and forest travel (Robinson 1970). Ocular orbits are less prominent, consistent with a less aquatic existence. The body profile slopes from the rear to fore facilitating movement through riparian vegetation. As in the Common Hippo, the skin is well lubricated with mucus-like droplets 81

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left:

Sketch of neonate. Pygmy Hippopotamus Chroeropsis liberiensis myology of head.

above:

exuding from pores that cover the body. The secretions may take on the appearance of a foamy film over the body surface when rubbed against vegetation. The digestive system is characterized by a four-chambered stomach (microbial fermentation probably taking place in the first three chambers), lacks a caecum and, like the Common Hippo, has simple small and large intestines (Macdonald & Hartman 1983). They are classified as pseudo-ruminants, as they do not regurgitate and chew their food. Newborn individuals have a lactational groove that diverts milk directly into the glandular portion of the stomach, resembling the same functional morphology found in ruminants (A.A. Macdonald & W. Hartman pers. comm.). Pygmy Hippos are primarily, albeit not exclusively, nocturnal and crepuscular, spending the daylight hours resting (Robinson 1970, Bülow 1988); evidence from camera traps shows that they may be active throughout the night and also during the day (Mallon et al. 2011). Foraging and Food  Pygmy Hippos consume a wide variety of herbaceous materials (forbs, sedges, ferns and fallen fruit), as reported by Hentschel (1990) and Robinson (1970, 1971, 2003). Hentschel (1990) concluded that the feeding spectrum is wide and consists mainly of herbs and fruits. His feeding experiments, direct observations and examinations of damage at feeding sites showed the use, as food, of 17 ferns, 26 dicotyledonous plants, 16 monocotyledons and the fruits of 24 tree species. Feeding preferences varied per individual and some ferns and monocots were readily eaten by all captured animals. The most preferred food plants had high sodium content. Captive diets frequently consist of green leafy vegetables, lucerne, grass hays, carrots, apples, oats, bread and supplements of trace minerals and alpha tocopherol. Robinson (2003) consistently found that a small, recumbent, vine-like forb, widely known as ‘Deewinkon’ (Geophila sp.), is a preferred food throughout Liberia. They do not appear to eat aquatic vegetation or animal matter in the wild. Heaped dung and tail-splattered excrement are commonly found along trailside vegetation. They very occasionally consume crops during wet seasons in remote agricultural areas, but are not regarded as a crop pest. They have been observed standing on their hindlegs (with their front legs against the stem) to reach ferns growing within Raphia palms (Bülow 1988).

Social and Reproductive Behaviour  Pygmy Hippos are extremely difficult to observe, and even in primary forest areas of low human population density, they are rarely seen. It has been rarely photographed, even with the recent advent of camera-trap technology (e.g. see Collen et al. 2011). Consequently, the behaviour of this species in the wild is poorly known, and detailed information is lacking on home-range, territoriality, sexual behaviour and maternal–young interactions.They are known to be highly solitary, and maximum group size reported is three, when a /, ? and a juvenile offspring may keep transient company while breeding (Robinson 1970).When frightened, they seek forest or rivers for escape from intruders, and are formidable at close quarters if injured or trapped. Considerable time is spent travelling/foraging along meandering, tunnel-like paths that they create in streamside vegetation and through forests and swamps (Robinson 1970). Hentschel (1990) recorded a home-range size of 50 ha for a radiocollared /, and a daily range of 1.8 ha; Bülow (1988) recorded a home range of 180 ha for a ?. Home-ranges of several female Pygmy Hippos may overlap, and they appear to be very residential. Homeranges seem to depend on the presence of small streams with submerged trees, root hollows, swampy depressions, and size and density of ground vegetation, rather than nutritional factors or proximity of rivers (Roth et al. 2004). In captivity, copulation has been observed on land and in water. However, parturition, contrary to the Common Hippo, is confined to land. Unlike the Common Hippo, young calves in captivity are not in the constant company of their mothers, but remain in seclusion and suckle intermittently (Galat-Luong 1981).This explains why tracks of young animals are hardly ever found (Hentschel 1990), because young start following their mothers at around 3–5 months of age when they are already quite large. Reproduction and Population Structure  No distinct parturition season has been observed in captivity or in the wild, although some hunters report that newborn animals are observed more commonly in the early dry season between Nov and Jan (Robinson 1970). Oestrous cycle is about 35 days, with oestrus lasting one to two days (Tobler 1988). Gestation period is reported to be 6–7 months in captivity (Zschokke & Steck 2001). They have an

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epitheliochorial placenta (K. Benirschke, pers. comm.). A single young is born out of the water with a birth-weight of 4.5–6.2 kg; twinning does occur, but is rare (less than 1% of pregnancies in captivity result in the birth of twins; Hlavacek et al. 2005). Weaning occurs at 6–8 months. Reproductive maturity is reported to be 3–5 years (Lang 1975); in captivity, the youngest / to give birth was 2 years 10 months, and the youngest ? to mate, 3 years 4 months (Hlavacek et al. 2005). Captive adult body size is achieved by three years. Maximum age in captivity is in the order of 35–40 years; one wild-caught captive / lived to nearly 44 years of age (A. Conway pers. comm.). While no data exist for wild animals, it is expected to be substantially less. Bülow (1988) recorded an instance in which a young Pygmy Hippo died as a result of drowning in the Azagny N. P., indicative that swampy areas such as the Raphia swamps in Azagny are not optimal for reproduction because young are not accomplished swimmers. Zschokke (2002) noted that the total captive population of the Pygmy Hippo at the time had a highly female-biased sex ratio at birth (41% ??), exceeding most other known distorted sex ratios in captive mammals. This deviation was not compensated for by a higher juvenile mortality in //, but seemed to be related to high feeding intensity and ‘hands-on’ husbandry that favoured production of daughters. Predators, Parasites and Diseases  The main predators, particularly for young, are Leopards Panthera pardus, Nile Crocodiles Crocodylus niloticus and African Rock Pythons Python sebae. Hentschel (1990) obtained a photograph of a juvenile Pygmy Hippo killed by a Leopard. Disease in wild animals is undocumented. In captivity, they are relatively hardy with competent husbandry, and most health problems relate to husbandry shortfalls. The author is aware of several cases of dental malocclusion (and see Johnston 2002) and solar dermatitis resulting from exposure to excessive solar radiation (e.g. Olivier 1975). Conservation  IUCN Category: Endangered C1. CITES:Appendix II (as Hexaprotodon liberiensis). The range of this species has diminished dramatically in the past 100 years (Robinson 1970, 1971, 2003), but most acutely in the past 50 years. Forests within the historical range have been steadily logged, farmed, converted to plantations (rubber, coffee and oil palm) and settled, resulting in a fragmented distribution. Unchecked forest destruction and other damaging human disturbances, such as mining activities, pose serious threats to all remaining Pygmy Hippos and their forest habitats. Sierra Leone has important remnant populations in the Gola Forest region (Gola Rainforest N. P.) bordering Liberia (and also in the vicinity of Tiwai I. and in the Loma Mts), while Guinea contains isolated populations along the Liberian frontier (including in the Ziama Biosphere Reserve, and Diécké and Mont Béro Reserves) that are declining rapidly due to the impact of displaced Liberian war refugees. Côte d’Ivoire has lost most of its original forest cover, and, aside from remnant populations bordering Liberia, most hippos reside within Taï N. P. and N’Zo Faunal Reserve and surrounding areas and in Azagny N. P. Liberia contains the most significant remaining forests suitable for Pygmy Hippos, with the greatest numbers in the central and southeastern regions, where Sapo N. P. is the only protected area for this

species. However, this refuge has experienced significant encroachments from illegal mining and other human activities between 2002 and 2010, and the effects on Pygmy Hippo numbers remain undetermined. Extensive road-building, logging and settlement activities, accelerating in the 1990s, are destroying and isolating the remaining forest areas of SE Liberia, particularly in the Cestos and Senkwehn riversheds (Robinson & Suter 1999). Due to the exodus of rural Liberian refugees to urban settlements and neighbouring countries starting in 1989, wildlife populations in general rebounded during the Liberian civil war, but that trend is now reversing (P.T. Robinson & J. Suter pers. obs.). The presence of the Pygmy Hippo may be a highly sensitive indicator of the degree of environmental disturbance; logging, hunting and settlement activities readily cause its disappearance, and seldom will it maintain a sustained presence within 5–10 km of such activities. However, Hentschel (1990) reports they will reoccupy secondary forests derived from logged areas. Primary methods of hunting this species are at night with headlamps and 12 gauge shotguns using solid metal projectiles, and with wire snares and pit traps. Captive animals are ordinarily pit-trapped or taken as young after killing the mother. Some wild-caught individuals have been said to become remarkably tame after very short periods in captivity (Schomburgk 1913). The Basle Zoo in Switzerland records the most captive births and maintains the captive breeding studbook.Animals are readily maintained in zoos and breeding has been generally successful, with Basle, Washington and Pretoria zoos the most successful breeders (Hlavacek et al. 2005). The number held in captivity in 1988 was 340 animals, although the number of zoos exhibiting this species had decreased (Tobler 1988). As of Dec 2009, there were 332 (133 ??, 196 // and 3 individuals of unknown sex) known Pygmy Hippos in 134 public zoos and private collections. The captive population is descended from 61 wild-caught founder animals, unequally represented in collections. The last wild-caught individual was imported on 22 September 1982. The current captive population shows a skewed, female-biased, sex ratio and a relative lack of young animals (Mallon et al. 2011; and see Hlavacek et al. 2005). In Nov 2010, a workshop was convened in Monrovia, Liberia, attended by representatives from all range states, government agencies, and local and international NGOs working on Pygmy Hippo conservation to develop a conservation strategy for the Pygmy Hippo with the goal to ensure the effective protection of, and connectivity between, known populations (Mallon et al. 2011). Measurements Choeropsis liberiensis TL: 1420–1570 mm, n = 4 WT: 179–273 kg, n = 7 Measurements from across the range (Lang 1975) Height at the hindquarters for a ? and /, measured by Lang (1975), was 810 mm and 830 mm, respectively The longest tusk on record measured 304 mm (Rowland Ward) Key References  Bülow 1988; Eltringham 1993b, 1999; Hlavacek et al. 2005; Hentschel 1990; Mallon et al. 2011; Robinson 1970, 1971, 1981. Philip T. Robinson 83

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Suborder RUMINANTIA

Suborder RUMINANTIA – Ruminants Ruminantia Scopoli, 1777. Introductio ad historiam naturalem, Prague, pp. 493-496.

In the taxonomic upheaval that has accompanied the application of genetic analysis to traditional nomenclature, Ruminantia has emerged as the primary category for most, but not all of the two-toed, predominantly herbivorous mammals. The ruminants are subdivided into what might, in functional terms, be called ‘proto-ruminants’, the chevrotains orTragulina, and the ‘true ruminants’ or Pecora, embracing the superfamilies Giraffoidea (giraffes), Cervoidea (deer) and Bovoidea (bovids). This division is well supported by molecular studies (e. g. Hassanin & Douzery 2003, Hassanin et al. 2012). The presence of the third forestomach compartment, the omasum, is a primary distinction between chevrotains, which lack this organ, and the Pecora or true ruminants. Extant tragulid species have become restricted in body mass and, although they were abundant and covered a wider body size range before the bovids evolved, they only survive today as relatively small forest dwellers in warm and humid climates. For more than 100 years, scientists have known that only specific microbes can produce the enzymes necessary for breaking down the complex carbohydrates in plant material, and convert them into the coals of the fire of life, the sugars.The capacity to digest plant material is shared by many mammal groups, such as rodents, horses, rhinos and elephants, in which bacterial fermentation takes place in the hindgut (‘hindgut fermenters’), and hippos, kangaroos or colobus monkeys, in which the microbes are housed in a forestomach (‘foregut fermenters’). In foregut fermenters, the bacteria flowing out of the fermentation chamber are not lost in the faeces, but enter the glandular stomach (called the abomasum in ruminants), where they are killed by the hydrochloric acid produced in this compartment.The dead bacteria, consisting of highly digestible protein, thus contribute significantly to nutrition in foregut fermenters. The products of bacterial fermentation of structural carbohydrates such as starch, cellulose, hemicellulose and pectin are always sugars, which are immediately used by the bacteria themselves; the metabolic products of their activity, namely three volatile fatty acids (VFAs), acetic, propionic and butyric acids, are absorbed by the mucosa of the fermentation chamber, be it the hindgut or the foregut. In ruminants, the rumen papillae increase the total mucosal surface, thus vastly enhancing the rumen’s absorptive capacity. The rumen papillae are efficiently served by an intricate vascular system, which immediately transports these VFAs to the liver via the portal vein. The liver is vital not only for detoxification, but primarily, as in all herbivores, for resynthesizing sugars, especially glucose, from these VFAs, a process known as gluconeogenesis. The problem of toxins, or substances such as polyphenols that suppress digestibility (and are present in many plants, except grasses), may represent a particular advantage for a foregut fermentation system. In the foregut, the gut bacteria can metabolize toxic substances before they reach the sites of absorption in the small intestine. In ruminants, additional support comes from the salivary glands, which produce particular proteins that bind, for example, to the polyphenolic compounds in browse forage.Therefore, many non-grazing ruminants have a more viscous, protein-rich saliva produced in large salivary glands, whereas grazing species, which do not need this kind of defence,

Ruminant stomach. Rumen (R) derives from cardiac region of stomach. Reticulum (ret) is a muscular sac derived from a loop in the body of the stomach. Omasum (o) is a lesser curvature of gastric primordium. Abomasum (ab) represents pyloric atrium (after Young 1962).

have smaller salivary glands that, nevertheless, produce a copious, but more watery saliva. The ruminant digestive system represents a unique evolutionary step forward by combining both the advantages of conventional microbial forestomach fermentation with a selective retention (and subsequent reprocessing) of large ingesta particles. This intensive use of ingested plant parts means that ruminants gain a comparable amount of daily energy from a distinctly lower amount of forage than non-ruminant herbivores. Ruminants ‘chew the cud’, but while we are so used to this well-known fact, we should not forget that this revolutionary adaptation necessitates a series of complicated morphophysiological adaptations. The sphincter cardiae (the barrier that prevents vomiting in horses and other animals) had to be removed. The smooth (i.e. involuntary) oesophageal musculature had to change into striated (voluntary) tissue. And in order for the rumination process to make sense, a sorting mechanism in the forestomach had to evolve to separate those particles that needed to be re-chewed from those that had already been digested and could pass on into the lower digestive tract. Just how this sorting mechanism works is still a matter of debate. It is assumed that it is associated with the ‘honeycomb’ cells of the second forestomach, the reticulum, and the third forestomach, the omasum; both structures are much more pronounced in grazing ruminants. While the omasum provides a mucosal surface of about 2000 cm² in large browsers such as the Moose Alces alces or Greater Kudu Tragelaphus strepsiceros, this surface is enlarged up to more than 35,000 cm² in some grazing bovines! And it is in these species that evidence for a particularly effective and selective mode of particle retention has been generated. The fact that grazers excrete very fine particles in their faeces (in contrast to browsing species such as Moose or Greater Kudu) supports the notion that the mechanisms of particle retention and rumination are more effective in the more recently evolved grazing ruminants. Thus, the forestomach anatomy of the tragulids (chevrotains or ‘proto-ruminants’) represents an evolutionarily older state, with no omasum at all. In true ruminants, the omasum was small to begin with, with few laminae, a sieve-like structure between the particlesorting reticulum and the glandular abomasum. The evolutionary

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success of the omasum structure can only be guessed at or inferred, partly by noting the displacement of tragulids from nearly all their previous ecological niches by true ruminants. A time scale for ruminant evolution has been inferred by Hassanin et al. (2012), who used a Bayesian molecular clock approach as well as evidence from the fossil record. According to their calculations, the primary split between chevrotains and Pecora took place about 51.6 mya, which was well before grasses began to spread widely during the Miocene. So, rumination did not evolve directly in order to digest grass, as has sometimes been assumed. Thus, a high proportion of extant species (approximately 40% of all ruminants) have retained what could be called the early ‘ruminant blueprint’, both in the anatomy and the physiology of their digestive tract. These, often conservative, species are highly selective and avoid forage rich in cellulose, especially mature grass, which they are incapable of breaking down into a digestible form. They select fruit, litter and foliage with a low cellulose content, but rich in soluble nutrients, and with indigestible, lignified fibre. They chew with more vertical bites (puncture crushing), rather than grinding their teeth sideways.There is little stratification of ingesta in their rumen, in contrast to what occurs in grazers. This category of ruminants has been characterized as ‘concentrate selectors’ (Hofmann 1973); of the antilopine tribes recognized in this work, the Neotragini, Madoquini, Oreotragini, Raphicerini and Cephalophini are all ‘concentrate selectors’. Only some 25% of about 180 extant ruminant species are true ‘grass and roughage feeders’ (Hofmann 1973, 1989). These species have refined their ruminal physiology and anatomy to the point where they derive most of their energy from slow ruminal fermentation. This type of rumination mainly processes the cellulose that is provided by the grasses and composites that are now ubiquitous in the extensive grasslands of Africa and other continents. The more these larger ruminants adapted to cellulose digestion, the less important detoxification became, and this has resulted in diminished salivary glands and a relatively smaller liver. There is also a large and significant category of opportunistic species that are intermediate between the two extremes mentioned above (Hofmann 1989). These ‘intermediate strategists’ comprise about 35% of all ruminants, and they are able to make use of both strategic avenues. They are especially well adapted to strong seasonality in African and other continental habitats because they can switch from grasses to browse material and vice versa (e.g. most cervids, Antilopini and Caprini).A recent literature review has synthesized dietary information for 78 species of African Bovidae (Gagnon & Chew 2000). The evolution of ruminant diversity is far from being a linear process; rather, it is more like a bush (or even a baobab tree!) since completely different permutations and variations occur, some of them retaining ‘older/conservative’ features, while others incorporate innovative morphophysiological adaptations, especially when grass is their main resource. There are evolutionary changes in the behaviour and morphology of those ruminants that ingest an increasing proportion of grass in their diet. Consuming more slow-fermenting cellulose fibre is accompanied by an increase in the size and capacity of both the rumen and the omasum (as well as changes in the molar teeth). In grazing species, rumination occupies a major portion of the daily activity budget. The exigencies of rumination and grinding cellulose fibre have caused changes in the shapes of mandibles, the fibre arrangement

Leptomeryx skeleton (primitive Oligocene chevrotain-like ruminant) (after Scott 1940).

of the masticatory muscles and the basic foraging apparatus. For example, pointed, delicate muzzles with manipulative lips transform into wide and rigid rostra, supported by the powerful gathering tongues that typify grazing bovines. Some ruminants select forage with a high content of rapidly digestible (and maybe even soluble) nutrients and a rather indigestible (lignified) fibre component. These species, even when large in body size, have retained a relatively small rumen, permitting only a short retention of their ingesta.This results in short (intermittent) rumination periods and frequent foraging bouts. There are striking convergences between bovids and deer in the more conservative species; thus duikers resemble muntjacs in being small ‘concentrate selectors’, while the Greater Kudu and the Moose are both large browsers with poor cellulose digestion. In contrast to grazers, many non-grazing ruminants show solitary and cryptic behaviour. There have been attempts to impose conceptual generalizations and categories on the overall trend from small, solitary, territorial browsers to large, social and hierarchical grazers. For example, Jarman (1974) concluded that both body mass and group size is negatively correlated to feeding selectivity, and group size is correlated with ‘antipredator behaviour’. However, the diversity of specific adaptations ensures that there are many exceptions. For example, the largest extant ruminant does not fit into the scale of body mass interpretation: the Giraffe Giraffa camelopardalis is a browser with a short retention time and has a poor capacity for cellulose digestion. It is true that grazing ruminants are frequently on the large side – but rarely above 1000 kg (compared with a maximum of about 1500 kg for the Giraffe!). Likewise, the Oribi Ourebia ourebia is a grazing ruminant that only weighs 15–20 kg, another ‘exception’ that illustrates that ruminants have evolved unique forms for almost every exigency. As the most widely distributed forage resource across all the continents, grass has become progressively more widely used by ruminants. Following long periods of adaptation across a wide span of the globe, the morphophysiological diversity of ruminants and the detailed structure of the rumination process in ruminants has changed and accommodated to many local permutations of climate, vegetation and ecology. Thus, the antecedents of grass-eating bovids had, almost 85

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The Okapi Okapia johnstoni is one of several ruminants that have conspicuous chins. Such patterns effectively advertise the activity of chewing cud.

certainly, begun to differentiate on the basis of other biotic parameters well before the major expansions of grasslands (Kingdon 1982). The spectrum of surviving ruminant types offers many insights into evolution, especially among bovids, and has implications for classification and nomenclature that are still very largely unexplored. The briefest of surveys of living ruminants reveals that the digestion of grass was a specialization adopted independently by later members of several lineages (notably some Reduncini and Caprini and virtually all Hippotragini, Alcelaphini and Bovini) with concomitant development

of broad, deep molars. These ‘hypsodont’ species do not represent a single closely related entity, as was supposed until recently (Schlosser 1904, Eisenberg 1981). In spite of substantial research on domestic stock, the livestock industry has failed to invest in research on the full adaptive range of ruminants and rumination, leaving the field to a handful of dedicated researchers in wildlife biology (e.g. Hofmann 1989, Jiang & Takatsuki 1999, Cerling et al. 2003, Sponheimer et al. 2003b, Hummel et al. 2005, Clauss et al. 2006). In the meantime, the politics of livestock promotion are destroying both ruminant diversity as well as the ecological diversity of African ecosystems. Because ruminant adaptive plasticity is an intensely interesting and useful process in its own right, and ruminant evolution appears to be still going on, a change of heart is long overdue. The livestock industry must modify its blinkered attitudes if conservation efforts are to succeed in retaining Africa’s ruminant diversity within their many native habitats; and change is overdue, not only in Africa. Reino R. Hofmann & Jonathan Kingdon

Infraorder TRAGULINA – Chevrotains Tragulina Flower, 1883. Proc. Zool. Soc. Lond. 1883: 184.

Lateral view of skull of late Eocene Hypertragulus (from Scott 1940).

Recent, molecular-led reformations in mammalian taxonomy support the majority of classifications that divide the order Ruminantia into two infraorders: the true ruminants, or Pecora (generally including those members possessing horns, antlers or ossicones), and the proto-ruminant chevrotain lineage, Tragulina (Hassanin & Douzery 2003, Hassanin et al. 2012; and see Hernández

Fernández & Vrba 2005). Since their divergence, some 51 mya (Hassanin et al. 2012), the pecoran lineage has diverged into more than 20 major lineages, which have continued to diversify right up to the present. Meanwhile, the chevrotains, after first registering in the fossil record in the late Eocene (in the form of Archaeotragulus from Thailand; Métais et al. 2001, and see Métais et al. 2007), remained a diverse and widely distributed group through the Miocene. They subsequently declined until only one representative remained in Africa, the Water Chevrotain Hyemoschus aquaticus. This decline was clearly strongly influenced by a less efficient digestive system than that of pecorans. Their likely ~50-million-year survival as a single lineage calls for the chevrotains’ continued existence to be registered at both infraordinal and superfamily level. In this work, as in most traditional treatments, tragulid peculiarities are described under the Family Tragulidae. Jonathan Kingdon

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Superfamily TRAGULOIDEA chevrotains Traguloidea Gill, 1872. Smithsonian Misc. Coll. 11 (230): 9.

The superfamily Traguloidea accommodates the ancient and once diverse chevrotains and mouse-deer. Chevrotains probably occupied a spectrum of early ruminant niches from at least the late Eocene

to the middle Miocene, but have been in decline ever since. The characteristics of chevrotains are summarized under their family profile, Tragulidae.

Family TRAGULIDAE Chevrotains Tragulidae Milne-Edwards, 1864. Ann. Sci. Nat. Zool. Paris, ser. 5, 2: 157. Hyemoschus (1 species)

Water Chevrotain

p. 88

The family Tragulidae occupies a threshold taxonomic position as ruminants. Most members of the suborder Ruminantia are also members of the infraorder Pecora comprising the extant families Moschidae (musk deer), Cervidae (deer), Antilocapridae (North American Pronghorn), Giraffidae (Giraffe and Okapi) and Bovidae (cattle, sheep, goats, antelopes). However, the more primitive tragulids are in their own infraorder, Tragulina. Nine tragulids occur in South-East Asia: six species in the genus Tragulus (Meijaard & Groves 2004, Grubb 2005) and three in the Indian genus Moschiola (Groves & Meijaard 2005). Only a single larger-sized species occurs in Africa, the Water Chevrotain Hyemoschus aquaticus, living in parts of the west and centre of the continent. Tragulids are secretive little forest-dwelling animals with an unusual association with water.They have a cannon bone (lengthened and fused metapodials III and IV) in their back legs, but not always in the front legs. Their legs, especially in front, are not very long and the animals have a low crouched appearance when standing. Each foot has four digits, although in Hyemoschus the side digits do not touch the ground when standing. Tragulids lack the horns or antlers found in nearly all pecorans. The dental formula is I 0/3, C 1/1, P 3/3, M 3/3 = 34. Males have curved upper canine tusks as in one or two pecorans such as musk deer, but of different cross-sectional shape; the lower canines look like incisors. Their molar teeth are less fully selenodont (with crescentic cusps) than in pecorans and the few cusps on the lower premolar teeth lie in an anteroposterior line and without additional or ‘new’ cusps on the lingual side (inside) of the original line. They have a compound stomach like that of pecorans, but without the omasum chamber, and rumination takes place in their digestion of food. Flower (1875) provided an exemplary account of the differences between the musk deer (a pecoran without horns or antlers) and tragulids. The first fossil tragulid to be named was Dorcatherium naui from early in the late Miocene (about 10 mya) of Germany (Kaup & Scholl 1834). Fortunately, the material included a complete skull. This skull differs from the living Hyemoschus aquaticus, which in 1834 had yet to be scientifically described and named, by being bigger (about onefifth longer), having longer nasals, a preorbital fossa and a first lower premolar which had largely disappeared in ruminants even before 10 mya. Miocene Tragulidae are known from European countries other

than Germany, and also in East Africa, Arabia, India, South-East Asia and China. The largest African fossil species and some of the European species from about 18–10 mya differ from later tragulids by their less selenodont molars with more signs of their former bunodont (with low rounded cusps) condition. It seems likely that the family evolved in southern Eurasia or, just possibly, in Africa.They are known from early in the African Miocene and Dorcatherium is generally thought to have migrated into Europe at about 18 mya. For about 170 years Dorcatherium was accepted as the earliest tragulid but, interestingly and anomalously, it is not known as far back in geological time as are pecorans. It occurs no earlier than about 22– 20 mya, almost back to the start of the Miocene and a long time after the earliest pecorans, at around 34 mya. The pre-Miocene ruminants, going back to around 45 mya, constitute a varied array of animals with many primitive attributes, the earliest of them even possessing upper incisors and a long tail. They were evolving selenodont teeth with every sign of convergences and parallels between the lineages. Moreover some non-ruminants were also evolving various patterns of selenodonty. The earlier Oligocene Lophiomeryx and its allies at around 30 mya lagged somewhat in the advance to selenodonty and were sometimes considered to be near tragulids. This could be true if it turns out that Tragulidae did not develop their apparently primitive characteristics as secondary reversals. Métais et al. (2001; and see Métais et al. 2007) made a new and substantial contribution to this ongoing discussion by describing late Eocene (ca. 40 mya) fossil dentitions from Thailand, named by them Archaeotragulus krabiensis, as a member of the Tragulidae. However, ideas about the concepts and contents of early ruminant and nearruminant families remain far from stable. Tragulidae have been contracting their range since the middle of the Miocene epoch. By the late Miocene they were less common in Europe and by the end of that epoch they were extinct there. In East Africa, the latest cited occurrence is Miocene. In China, at the present day tragulids range only into a small southern area. This contraction is like that seen in the anthracotheres, of which the last survivor was the Indian Merycopotamus at around 3 mya. These declines must be linked with long-term climatic change leading to diminution of suitable wetforest habitat and perhaps to the post-Miocene success of pecorans. Alan Gentry 87

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Genus Hyemoschus Water Chevrotain Hyemoschus Gray, 1845. Ann. Mag. Nat. Hist. ser. 1, 16: 350.

Hyemoschus includes only theWater Chevrotain H. aquaticus of theWest and central African rainforest. Water Chevrotains are small animals with short legs, large lateral hooves, a moderately long bushy tail and naked rhinarium. The limb bones are less reduced and co-ossified than in typical ruminants, with the ulna complete and separate from the radius, and metapodials II and V slender, but complete. Fusion

of the median metapodials is less advanced than in the related IndoMalayan Moschiola and Tragulus, with metatarsals III and IV fused to form a cannon bone, but the suture is not complete and still visible, and metacarpals III and IV are unfused; all metapodials are very short. Peter Grubb

Hyemoschus aquaticus  Water Chevrotain Fr. Chevrotain aquatique; Ger. Hirschferkel Hyemoschus aquaticus (Ogilby, 1841). Proc. Zool. Soc. Lond. 1840: 35 [1841], Sierra Leone, Bulham Creek.

Taxonomy  No subspecies have been recognized. The species is apparently monotypic across a large range, which is separated by the Dahomey Gap between the Upper Guinea and Congo Basin forest blocks. Synonyms: batesi, cottoni, typicus. Chromosome number: not known.

Description  A squat, short-limbed, small-headed ungulate, with a compact body form, largest at the hindquarters, which are often held higher than the shoulders, and strongly tapered toward the head. Body covered with long-haired pelage overall umber brown to reddish-brown, grizzled with paler hairs, and marked with variably

Water Chevrotain Hyemoschus aquaticus.

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Water Chevrotain Hyemoschus aquaticus.

distinct pale to white stripes and spots. The most distinctive white markings include longitudinally oriented white malar and neck patches offset by a darker line of pelage along the side of the neck, extending from below the chin and behind the gape area onto the neck. One to two variable longitudinal white stripes extend roughly in parallel along the sides of the body, gradually disappearing on the rump, and a series of pale dorsal spots, 1–3 cm in diameter are arranged as 6–7 irregular broken bands across the shoulder through the middle back. The small but somewhat elongated head narrows on the crown above large eye orbits and between relatively small ears that are dark on the outside with lighter hairs lining their inner surfaces. The hair on the crown is dark intermixed with variably lighter pelage around the eyes and on the muzzle. The eyes are dark brown and produce a whitish reflection when seen at night under torchlight. The chin and underparts of the neck are white, variably set off by dark brown line of pelage extending posteriorly from the upper chest. The lower chest and abdomen are grizzled brown. Tail dark brown above, white below, extending into a white fringe. The udder with four teats is covered with white hairs and located between the hindlegs. Hooves similar to other small to medium-sized ungulates, though slightly more rounded than those of the similarly sized forest duikers. Prominent hoofed metacarpals are present on all four limbs, though seldom visible in tracks. Skull not unlike that of Indo-Malayan tragulid genera but more than one-and-a-half times longer, highest at the back and sloping down to the snout. The first incisors are broad. Other incisiform teeth have very narrow crowns; the cheekteeth are bunoselenodont, with relatively large sectorial premolars. Distinctive thin and strongly curved upper canine teeth (tusks) occur in both sexes, but are larger and externally visible in ??.

Lateral, palatal and dorsal views of skull of Water Chevrotain Hyemoschus aquaticus.

Cuniculus paca, which is only slightly smaller than theWater Chevrotain, has a similar pelage and body form, a remarkable case of convergence in two unrelated species with comparable ecology and cursorial mode of existence (Dubost 1968a, Eisenberg & McKay 1974). Distribution  Endemic toWest and central Africa; the distribution ranges across the forest belt from Sierra Leone and SE Guinea, through Liberia, S Côte d’Ivoire into SW Ghana. Grubb et al. (1998) discuss the validity of supposed records from west of Sierra Leone, concluding that, unless a record from the Kounounkan Forest in Guinea is substantiated (see Barnett et al. 1996), the furthest west the species has been reliably recorded is Sierra Leone. However,

Geographic Variation  None recorded. Similar Species  The only tragulid representative in Africa, the Water Chevrotain is unmistakable. It shares its forest habitat with up to six species of duikers, which are similar in overall body size and share its primarily frugivorous diet. Water Chevrotain tracks might be confused with mid-sized duikers, especially as the print of the ‘dew hooves’ in their tracks is usually not evident. The Water Chevrotain is the only artiodactyl that can recline on its hind metatarsals and frequently does so as it settles to rest (Dubost 1968a, and see illustration in Kingdon 1979). The Neotropical caviomorph rodent, the Paca

Hyemoschus aquaticus

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there are recent photographic records of the species from SE GuineaBissau in the Boé (P. Wit pers. comm.). The species is apparently absent from the Dahomey Gap, from E Ghana through Togo and Benin. Although formerly widely distributed in S Nigeria, East (1999) mapped it only east of the Niger R., from whence it ranges east through the central forest block, across S Cameroon, Gabon, Cabinda (Angola), Congo and DR Congo to extreme W Uganda, where it is now believed extirpated (East 1999). Crawford-Cabral & Veríssimo (2005) report a record from Angola from the Lunda Norte Province, near the Cassai R., which is the southernmost record of the species from the continent. The Water Chevrotain’s occupancy of this range is highly discontinuous, being limited to the vicinity of streams and rivers. Forest loss and continuing forest degradation have further reduced suitable range in many areas, in particular in the remnant Upper Guinea forest block. Pickford et al. (2004) recently recovered remains of the Water Chevrotain from the early Pliocene in the Kenya Rift Valley. Habitat  Water Chevrotains are confined to closed-canopy, moist, tropical lowland forest, and within this habitat, they only occupy areas in the vicinity of streams and rivers. When disturbed, they frequently move to water, often using well-established trails and paths. If pursued, they may conceal themselves in the water by remaining still and almost entirely submerged beneath debris or under overhanging stream banks. The Water Chevrotain is not a swamp specialist, nor strictly limited to riverine forest, and often ranges, forages and even rests, in mature upland forest areas. But these areas are almost always within several hundred metres of steams and rivers large enough to provide cover (Dubost 1978). Water Chevrotains do not range out into gallery forests, or into forest savanna mosaic, even along river courses. This underlines the primary importance of large areas of mature evergreen forest for the survival of this species. Abundance  Overall distribution and abundance are limited by the proximity of suitably sized water-courses. In the Ituri Forest, densities have ranged from 1.5 to 5.0/km2 (Hart 1985, 2000). Hoppe-Dominik et al. (2011) reported a density of 0.5/km2 in Taï N. P. based on transect counts. Much higher densities have been reported in Gabon, where up to 28/km2 were recorded on an island in the Ivindo R. (Dubost 1978). East (1999) estimated the total population size at around 278,000 animals. However, given the differences in reported densities, and with current uncontrolled bushmeat hunting over much of its range, any attempt to compute overall current numbers is unrealistic. Adaptations  The Water Chevrotain, along with the Asian tragulids, is considered among the most primitive and conservative of ruminants. Tragulids are thought to have separated from the pecoran ruminants in the early Eocene (Hassanin & Douzery 2003). The Water Chevrotain retains many traits thought to characterize the earliest of the ruminants, including small body size, forest habitat and a selective ‘concentrate’ diet (Kingdon 1979). Many of the skeletal traits of the species are identical to those of the upper Miocene tragulid, Dorcatherium, which was discovered as a fossil before the living Hyemoschus was discovered in Africa (Turner & Anton 2004). Water Chevrotains exhibit some behavioural traits of the Suidae, including similar roles for olfactory signals and use of canines in intra-specific combat (Dubost 1975, Kingdon 1979). The

Water Chevrotain Hyemoschus aquaticus open-mouthed head myology.

overall adaptations of the Water Chevrotain suggest a species that is specialized for a stable but low energy trophic niche, coupled with a unique strategy for predator avoidance. Water Chevrotains are nocturnal and spend the daylight hours resting, most often in concealed locations such as between tree buttresses, or in thick cover in tree falls, but surprisingly often in more open settings (Dubost 1978). This species relies on a complex pattern of light and dark colouration combined with immobility to provide camouflage and avoid detection. Resting animals remain immobile and may be approached to within just a couple of metres before they flush. Water Chevrotains differ from sympatric duikers in their use of water as an avenue of escape. Black-fronted Duikers Cephalophus nigrifrons have specialized on swamp forests (an adaptation that is probably influenced by lower predator pressure), but Water Chevrotains, unlike Black-fronted Duikers, do not have elongated hooves and are the only small forest ungulates to actively conceal themselves by submersion in water. Water Chevrotains swim freely, although they are not sustained swimmers. Cinematography of the species under water has revealed that submerged animals move along the surface of the stream-bed using a walking gait reminiscent of submerged hippos (A. Root pers. comm.). The slit nostrils, which bear some resemblance to the valved nostrils of hippos, appear to be adapted to this aquatic escape mode. Water Chevrotain dentition differs from that of similar sized frugivorous forest duikers in having thinner, finer-edged selenodont teeth that are weakly bedded in the skull, adapted to soft foods and ill-adapted to handle hard or coarse forage. The method of chewing resembles the shearing and cutting movements of some carnivores, and does not resemble the grinding and crushing mastication of other ungulates and suids (Dubost 1964). Water Chevrotains have a well-developed rumen and reticulum as in other foregut fermenting, pecoran ruminants but the omasum, at the posterior end of the fermentation chamber, is minimally developed. Thus the capacity to

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Water Chevrotain Hyemoschus aquaticus skeleton.

Water Chevrotain Hyemoschus aquaticus myology.

retain food in fermentation chambers is limited (Hofmann 1973). The ratio of rumen volume to body weight is lower in the Water Chevrotain than it is in similar-sized sympatric frugivorous duikers, providing further evidence that the rumen is designed for rapid food turnover. This suggests that foregut fermentation in this species has evolved less for efficient breakdown of complex plant carbohydrates than to provide important detoxification functions for a diet that contains a wide range of digestion inhibitors and toxins. Chevrotains produce abundant saliva: this may capture and immobilize tannins, and other digestion deterrents and plant defensive toxins. While the ability to handle plant toxins allows theWater Chevrotain to exploit a wide range of food sources, detoxification nevertheless incurs energetic and other costs. Water Chevrotains differ from many other ruminant ungulates in the strikingly pale colour of its flesh, in particular the musculature of the limbs. This is indicative of lower levels of myoglobin and lower metabolic capacity in the muscles. Unless they can reach water and find refuge quickly, Water Chevrotains that are flushed from hiding can be readily run down and captured, a characteristic exploited by the Mbuti hunters of the Ituri Forest, who sometimes capture animals by hand after flushing them from hiding and blocking their access to water.

diet. In West Africa, the fruits of the abundant Marantaceae have been listed in the diet (Malbrant & Maclatchy 1949). Dubost (1964) has suggested that meat may be a significant dietary item. In the Ituri Forest, Water Chevrotains as well as larger duikers, and Red River Hogs Potamochoerus porcus are known to feed on carcasses. Consumption of rotting carrion may even be advantageous for the ruminants, as the ammonia produced by the decaying protein is the most immediately available source of nitrogen for anaerobic microbial flora of the rumen. There are also reports that Water Chevrotain eat fish, insects and even aquatic plants (Dekeyser 1955). Rahm (1966) reports that Water Chevrotain are sometimes baited into traps with dead crabs or rats. His captive chevrotains readily ate live freshwater crabs offered to them. No evidence of these food sources was found in the rumens that were studied in the Ituri Forest (Hart 1985).

Foraging and Food  Fruits, seeds and fallen flowers, eaten from the forest floor, dominate the Water Chevrotain’s diet. A wide variety of plant species have been reported in the diet from across its range (Gautier-Hion et al. 1980, Dubost 1984). A study of rumen contents taken from ungulate frugivores in the Ituri Forest (Water Chevrotain and six species of duikers) revealed that the Water Chevrotain had the most diverse diet on an annual basis (n = 22 stomachs) (Hart 1985). Important plant families in the diet include: Irvingeaceae, Sapotaceae, Euphorbiaceae, Apocynaceae, Sterculiaceae, Annonaceae, Ulmaceae, Sapindaceae and Meliaceae. Soft unripe fruits and unripe seeds were especially prominent in stomachs and included a number of species such as the Irvingiaceae, that were not recorded in diets when ripe. These food sources are all likely to contain protective chemical defences. Mast seed-fall of caesalpinaceous dominant canopy species, Julbernardia seretii and Cynometra alexandri, were also important seasonally. Animals captured by hunters during periods of mast seedfall had layers of fat around their kidneys that were absent at other times of year. In Gabon, dicotyledonous families also dominate the

Social and Reproductive Behaviour  Water Chevrotains are primarily solitary. Dubost (1978), describing a population on an island in the Ivindo R. in Gabon, reported stable and mostly exclusive female home-ranges of 13–14 ha, while ?? had larger home-ranges of 20–30 ha that overlapped those of //, although they spent little time with them. In the Ituri Forest, in an area of small rivers and creeks, two radio-collared // utilized home-ranges of 22 ha and 28 ha over the course of a year (J.A. Hart pers. obs.). They are irregularly active during the night, with periods of frequent movement alternating with periods when the animals are quiescent (Dubost 1978). In the Ituri Forest, radio-collared // spent on average over 40% of each night resting. Water Chevrotains have remarkably dense and thickened skins. This may protect the animals from injury in their encumbered environment; however, it also appears to protect against injuries in confrontations between conspecifics (Kingdon 1979). The canines are sharp and exposed. One ? caught in the Ituri Forest had a deep gash on its lower neck that might have been caused by a canine slash from a conspecific. Water Chevrotains have numerous epidermal glands, notably in the area under the chin. These are likely used in intra-specific marking, possibly in association with mating, but may also leave chemical signals at bedding sites. As yet there is no evidence for ritualized marking of home-ranges (faecal piles, rubbing posts, etc.). Dubost (1965) described a courting ? making brief cries as it approached and followed a receptive /. Local Mbuti 91

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hunters attributed ‘chuffing’ barks heard at night in the Ituri Forest to this species. Reproduction and Population Structure  A single young is born after a gestation variably estimated at between four and nine months, and most likely in the region of 220 days (see Dubost & Feer 1992). In NE Gabon, births were recorded throughout the year, but with higher numbers in Dec, Jan, Apr, Aug and Sep (Dubost 1968b; and see Dubost & Feer 1992). In the Ituri Forest, births were also widely dispersed over the year. Of seven adult //, three (May, Jun and Sep) either carried full-term foetuses, or had recently given birth.Two young animals, both caught in Jan had been born in Dec. Four // were not pregnant (Apr, Aug, Oct and Sep); another had been recently impregnated in Sep. The largest full-term foetus weighed 630 g (about 5% maternal weight). In the forests of Kivu (Utu area), Rahm (1966) reported that a / pregnant at the time of her capture and kept in captivity with a ? gave birth in early Mar. Sixteen months later, in late Jul, the same / died and was found to be bearing a full-term foetus. Water Chevrotains appear to grow quickly. In the Ituri Forest, // with three molars and deciduous premolars had begun to breed, but no

animals with only two erupted molars were in breeding condition. Based on tooth eruption estimates by Dubost (1978), the estimated age of female maturation would fall between 20 and 24 months. Predators, Parasites and Diseases  Documented predators include Leopards Panthera pardus and possibly African Golden Cats Profelis aurata. In one study of Leopard diets in the Ituri Forest, Water Chevrotain remains were not as frequent in scats as those of nocturnal duikers (Hart, J. A. et al. 1996). It is possible that small ranges and limited movement byWater Chevrotains, even at night, in conjunction with their use of water as a refuge, may reduce vulnerability to predation. There is no information on parasites or diseases. Conservation  IUCN Category: Least Concern. CITES:Appendix III (Ghana). Water Chevrotains appear to be rare, and may have disappeared from much of their historic range in West Africa. However, the species is still frequently encountered in many areas of central Africa, and is one of the preferred bushmeat species in the Kisangani region where its ‘white’ flesh is compared, somewhat exaggeratedly, to fish. In the central Ituri Forest, Water Chevrotains are regularly caught by the Mbuti net hunters (Hart 1979, 2000), consistently representing about 5% of total catch, even in areas that have been hunted for years This low capture rate is probably due to the hunters’ avoidance of riverine areas where chevrotains are most likely to occur. Water Chevrotains are particularly vulnerable to snares. One /, caught by Pygmy net hunters in the southern Ituri Forest, had a gangrenous snare wound on its front leg. Despite this, the animal had normal body weight and a full-term foetus. Water Chevrotains occur in a number of protected areas, including: Sapo N. P. and Grebo National Forest (Liberia), Taï N. P. (Côte d’Ivoire), Lobeke N. P. (Cameroon), Lope N. P. and Minkebe N. P. (Gabon), Maiko N. P., Kahuzi-Biega N. P. and Okapi Faunal Reserve (DR Congo) and Odzala N. P. and NouabaléNdoki N. P. (Congo) (East 1999). Ultimately, Water Chevrotains will only be conserved by the protection of large areas of mature, undisturbed forest. Measurements Hyemoschus aquaticus HB (sexes combined): 620–1020 mm* T (sexes combined): 75–150 mm E: 60 mm Central Africa (Rahm 1966, Dubost 1975) *Mean 725 mm (??), 768 mm (//) Dubost (1975) cited a range of 7–15 kg with a mean 9.7 kg for ?? and 12.0 kg for //. A sample of 13 adult body weights from the Ituri Forest (J. A. Hart pers. obs.) ranged from 9 to 13.5 kg with a mean of 10. 3 kg for ?? (n = 6), and 12.1 kg for // (n = 7) The longest tusk on record, from Cavallo R., Liberia, measured 57 mm (Rowland Ward)

Water Chevrotain Hyemoschus aquaticus.

Key References  Dubost 1964, 1965, 1968a, b, 1975, 1984; Hart 1985, 2000; Rahm 1966. John A. Hart

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Infraorder Pecora

Infraorder PECORA – Horned Ruminants Pecora Linnaeus, 1758. Syst. Nat., 10th edn, 1: 65.

The Pecora embraces all the extant families of horned ruminants: Cervidae (deer), Giraffidae (Giraffe and Okapi), Bovidae (bovines and antelopes), Antilocapridae (pronghorns) and the hornless Moschidae (musk deer). The last are Asian, while Antilocapridae are exclusively North American. The Giraffidae are now entirely African, although they were also known in Eurasia until the late Pliocene. Inter-relationships between and within these families have been explored in some detail by several investigators using molecular and other techniques (e.g. Hassanin & Douzery 2003, Hernández Fernández & Vrba 2005, Hassanin et al. 2012; and see Prothero & Foss 2007). The Pecora probably arose in the Eurasian land mass during the Eocene: the earliest pecorans were small and hornless (with narrow bladed upper canines in the ??) and resembled modern tragulids (chevrotains) in size and general form, although both tragulids and pecorans are derived in their own different ways from the ancestral traguloid ruminant stock. According to Hassanin et al. (2012), the chevrotains branched away from the lineage leading to the Pecora in the Early Eocene, about 51 mya. There are substantial discrepancies between this molecular clock date and the first appearance of modern pecoran families in the fossil record at the start of the Miocene, around 23 mya (Cote 2010). However, there is a record during the Late Eocene of Thailand of a modern tragulid (Métais et al. 2001, 2007) and fossils are known from the Late Eocene and Oligocene of traguloids more derived than tragulids (e.g. leptomerycids), and primitive small, hornless pecorans (e.g. gelocids) (Prothero & Foss 2007). The latter would have resembled chevrotains in appearance, but shared a parallel-sided astragalus with other pecorans (Webb & Taylor 1980), indicative of greater cursorial specializations. Modern pecorans are usually of larger body size, and are typified by the possession of horns, ossicones or antlers in ?? (and even in the // of some species and populations). The evolution of horns can sometimes be correlated with living at high densities, sometimes with territorialism, or with larger body masses. Remarkable as it may seem, it is likely that horns (or horn-like organs such as antlers and ossicones) evolved convergently several times, perhaps independently in each horned pecoran family. The living moschids are primitively without cranial appendages (Davis et al. 2011). The mid-Eocene saw several climatic changes in the higher latitudes, but major changes in climate and vegetation only took place in the late Eocene (Hooker 2000, Janis 2007). The issue of timing is important because pecoran ruminants have some physiological advantages and anatomical features that can be correlated with at least two significant changes that are known to have taken place in the late Eocene. One was the development of more seasonal climates; another was an increase in leguminous plants (see Volume 1). Janis (2007) has suggested that the rise of the ruminants corresponded with an innovation in the way plants responded to changes in climate. Before the onset of marked seasons most plants grew thick and durable perennial leaves or leaflets. Up to this time herbivores were therefore adapted to bulk feeding on foliage that had cell walls rich in cellulose. These animals were fast throughput, hindgut digesters,

typified by the perissodactyls that were dominant during the Eocene, so what seems to be at issue in the late Eocene is the beginnings of a decline in bulk processing versus the initiation of selective feeding, possibly with a reduction in the quantity of available vegetation (Janis 2007). While late Eocene climatic changes seem to correlate with the diversification of ruminants in general, this cannot be tied specifically to the type of pecorans that we know today (the rise of the contemporary pecoran fauna is better correlated with changes in the early Miocene; Solounias & Dawson-Saunders 1988). It is only during the Pliocene that the species richness and composition of fossil bovid species starts to resemble the present (Bobe & Eck 2001). Following the Eocene a great increase in deciduous leafing offered advantages to herbivores that could select out smaller quantities of higher quality foliage and extract more nutrients out of less material, even if the processing of food took longer to pass through the system. The ancestral Pecora may not have been as efficient at extracting and processing nutrients from deciduous foliage as some of the living, highly derived Bovidae or Cervidae, but the primary innovation in Pecora was development of a more advanced digestive capacity which is expressed today in the possession of a fourth forestomach compartment, the omasum. The omasum, situated between the fermentation chamber of the rumen-reticulum and the true stomach (abomasum), increases the digestive capacity of the foregut by enlarging its absorptive surface with extra papillae and laminae (the details of ruminant physiology are outlined in a later section). A primary innovation of the pecoran omasum is probably its conformation into a ‘sieve’ that helps sort and separate incompletely crushed particles from those digesta that are already sufficiently processed to proceed on their journey through the gut. This innovation helps make rumination much more efficient, but the exact course of its evolution must remain speculative. While we cannot date the omasum back further than the common ancestor of extant pecorans (which certainly would not be as early as the late Eocene), it is, perhaps, by around the mid-Oligocene when animals such as Amphitragulus (an identifiable cervoid) appear as fossils and such a supposition becomes more secure. Rumination has the additional advantage of nitrogen cycling: dietary protein is fermented to form ammonia, which is converted to urea in the liver, and then fed back to the rumen bacteria.The protein actually digested by the animal is from the rumen microbes, which frees ruminants from needing to find all the essential amino acids in their diet. This cycle also ties in with waste urea, which allows for the conservation of urinary water, allowing ruminants to survive in drier habitats. The craniodental morphology of early pecorans supports the notion of a relatively soft browsing diet and they would likely have had a rudimentary omasum and probably preferred highly nutritious vegetation, such as new shoots and fruit (a preference that is still evident in some bovids, cervids and giraffes). The selection of grasses as food only took place much later and this major shift into graminivory (mainly seen in the bovids, less so in the cervids) led to massive increases in the size of the omasum and the storage capacity of the foregut, together with appropriate modifications of salivary 93

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Superfamily Giraffoidea

glands, livers, teeth and chewing apparatus. The development of selenodont cheekteeth seems likely to be coincident with the development of foregut fermentation, the former first appearing in the late Eocene. It has been pointed out that browsing ruminants can be as much as 400 kg heavier than the largest grazing ruminant (giraffe // may average about 1200 kg): this is because, with enlargement, ‘packing constraints’ and the production of methane within the abdomen eventually constrain roughage grazers (Clauss et al. 2003). Thus, the African Buffalo Syncerus caffer, with an enormous rumen, is probably not very far short of that threshold. Deer are only peripheral to Africa, being particularly successful in temperate latitudes of Eurasia, but with extensions into lower latitudes

in the Americas and Eurasia. Bovids have the broadest geographic range, although unlike cervids they never extended their range into South America. The early Miocene African fossil record is sparse, and at the moment we cannot be certain when bovids entered the continent. Some primitive pecorans of unknown taxonomic affinity are known from around 20 mya (Cote 2010), but definitive bovids are not present until around 17 mya, and it was not until around 15 mya that they (along with giraffids) became a significant part of the fauna (Gentry 2010). The radiation of antelopes, the most successful of pecorans in Africa, is discussed elsewhere in this volume. Christine Janis & Jonathan Kingdon

Superfamily GIRAFFOIDEA Giraffe, Okapi Giraffoidea Gray, 1821. London Med. Repos. 15: 307.

The giraffes are one of four extant groups of ruminating herbivores in Africa: chevrotains (Tragulidae), deer (Cervidae) and bovids (Bovidae) being the others; the Pronghorn Antilocapra americana (Antilocapridae) and musk deer (Moschidae) are exclusively American and Asiatic. Today differences are obvious, but from comparing their anatomy (especially teeth), and from their fossil history, it is clear that giraffoids share a common origin with deer (more distantly with the bovids and more distant still, the chevrotains). Molecular clocks estimate the divergence between protogiraffes and protocervid/ bovids as occurring about 23.4 mya, while the tragulid/pecoran divergence was closer to 51 mya (Hassanin et al. 2012). It is possible that a combined examination of differences in geography, climatic adaptation, body-size, limb proportions, fossils, phylogeny and genes may eventually help explain the origins of differentiation between cervoids and the earliest giraffoids. The region of origin for cervoids is undoubtedly Eurasia, probably temperate Eurasia. Because the first known fossil giraffe had been found in the 1840s from the Siwalik Hills (G. sivalensis), there has long been a presumption that giraffes might have originated in the subtropical Indian sub-continent. Given that fossil giraffoids are diverse and well represented in India, this remains a possibility, but the genus Giraffa is far too late (evolving only 7–8 mya) to indicate the origins of the superfamily. None the less, the genus Giraffa may well have originated in India, and Harris (1991b) considered G. punjabiensis the oldest member of the genus (7.3–7.1 mya) and the one most like Bohlinia, a widely distributed proto-Giraffa. Giraffoidea had branched into two different families long before the 8-million-year-old G. sivalensis and G. punjabiensis. The Climacoceratidae are long extinct while modern Giraffes and Okapis exemplify the Giraffidae. Some resemblance between the earliest cervoids and giraffoids are exemplified by the deersized, deer-shaped Climacoceras africanus (a social herbivore that is common in East African deposits of about 17–15 mya), which had very long, spike-like male ‘antlers’ with short ‘tines’ sprouting along their length; however, it should be noted that Harris et al. (2010) recognize Climacoceratidae and Giraffidae as separate families.

In spite of superficial similarities, the physiological origins of giraffoid bony ossicones and non-bony deer antlers differ, although both structures probably derive from damage-minimizing or healing processes on the crania of ancestral stocks. Selection for what began as wound-generated lumps and bumps (that incidentally served defensive purposes) eventually became selection for species-specific cones, of dermal origin, with growth patterns that, in the giraffes, transformed cranial structures into genetically programmed, explicitly offensive weapons. If correct, this evolutionary progression illustrates the primacy of defensive behaviour and structures, while weapons tend to be secondary structures that become integrated into earlier defensive structures. Most giraffoid genera have ossicones in both sexes. The sivathere Helladotherium had // without ossicones but that seemed to be the exception rather than the rule. The earliest known fossil form of giraffine is Canthumeryx sirtensis. This fossil was discovered in deposits near Gebel Zelten some 200 miles south of Tunis, which is now desert but at the time of Canthumeryx was a flourishing alluvial/flood-plain/savanna/river basin habitat. Canthumeryx was a medium-sized antelope, about the same size as a Fallow Deer Dama dama, and with the characteristic bilobed giraffoid lower canines (Hamilton 1973, 1978). Body-size differences clearly separate modern giraffes and deer; even greater contrasts existed between the well-named Sivatherium giganteum and any fossil cervid. So, at some quite early stage of the divergence between the giraffe and deer lineages, size may well have become a decisive factor. Large size would have included the advantage of higher feeding zones: thereafter, the elongation of both limbs and necks would have been selectively favoured, because it offered still greater access to high-quality food resources that were out of reach for other herbivores. There are interesting comparisons here to be made with the Gerenuk Litocranius walleri, which stands on its hindlegs to augment an already exceptional reach. A summary of the characteristics of living Giraffoidea can be found in the family and generic profiles and warrants no repetition here. Jonathan Kingdon & John M. Harris

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Family GiraffidAE

Family GIRAFFIDAE Giraffe, Okapi

Giraffidae Gray, 1821. London Med. Repos. 15: 307. Giraffa (1 species) Okapia (1 species)

Giraffe Okapi

p. 96 p. 110

European peoples of the past were familiar with deer (family Cervidae) in the wild places of their own continent. They would also have known of Bovidae as sheep, goats and cattle in their farmed countryside and as ibexes and chamois in the mountains. Some had heard of gazelles in the great desert belt south of the temperate lands, and when Africa was explored it was found to be full of antelopes (Bovidae again). So Cervidae were found in Eurasia, while Bovidae concentrated in Africa. In addition, a single, spectacular large African pecoran ruminant, the Giraffe Giraffa camelopardalis, belonged to a different family, the Giraffidae, and in 1901 a second giraffid, the Okapi Okapia johnstoni, was discovered in Africa. The Giraffe and Okapi are large ruminants with long legs and long necks. Both species have somewhat toughened skin over their bony horns, present in both sexes in the Giraffe, but only in ?? in the Okapi. Both have a dental formula of I 0/3, C 0/1, P 3/3, M 3/3 = 32 (Singer & Boné 1960), with small and low-crowned cheekteeth, bilobed lower canines and prehensile tongues. They browse off shrubs and trees well above ground level, yet the Giraffe is an open-country or bushland inhabitant, sometimes in quite dry areas, whereas the Okapi is a true forest inhabitant. It is interesting and instructive that such similar cheekteeth can be found in species of such contrasted habitats. The earlier relatives of Giraffes go back to early in the Miocene epoch, before 20 mya, in Africa (Morales et al. 1999) and perhaps elsewhere. Conclusions about their ultimate continent of origin are much influenced by molecular phylogenies that still do not agree on which pecoran family is most closely related to Giraffidae. Price et al. (2005) and others (see Meredith et al. 2011, Hassanin et al. 2012) have even suggested the North American Antilocapridae, which, it has to remembered, descend from immigrants from the Old World. According to Harris et al. (2010), the family Giraffidae contains the subfamilies Canthumerycinae, Bohlininae, Giraffokerycinae, Sivatheriinae, Palaeotraginae and the living Okapiinae and Giraffinae. The first unequivocal giraffids, animals like Canthumeryx and Injanatherium, have been found as fossils from Africa to Pakistan, also in south-eastern Europe and their descendants to other parts of Asia. These early giraffids were large ruminants for their period (17–10 mya) and had widely divergent horns with spreading bases broadened in an anteroposterior direction and flattened mediolaterally. Sometimes they had a small anterior pair of horns in front of the orbits in addition to the main pair, but the nature of the skin covering of the horns is unknown. Their cannon bones or metapodials were longer than in the various deer and antelopes living alongside them. Coincidental with the arrival of three-toed hipparionine horses from North America between 11 and 10 mya, Old World ruminant faunas changed. Two subfamilies of more advanced giraffids appeared: Giraffinae and Sivatheriinae. Sivatheres started grazing with the spread of C4 grasses (J. Harris pers. comm.) and became very large and

Sivatherium giganteum skull in anterior view (left); skull of Miocene giraffid Giraffokeryx punjabiensis in dorsal view (top right) and lateral view (bottom right).

bulky, heavy-headed animals with branched or bulbously-tined horns, retaining an anterior pair of horns, and re-evolving somewhat shorter cannon bones. They persisted into the Pleistocene in India and Africa, in the latter until 500,000 years ago or possibly later. They became extinct presumably for the same reason that caused the demise of the large grazing bovids (Pelorovis, Megalotragus) and giant grazing pigs. The common Eurasian late Miocene giraffines are a separate development from whatever stock gave rise to modern Giraffa. Some of them approach hypsodonty in their teeth, and microwear studies have shown that they must have grazed. In this they are very unlike Giraffa, as also in that their legs were not invariably lengthened to the same great extent. After flourishing for several million years they declined and later became extinct in the Pliocene. The precise phylogenetic position of Okapia is unclear, the key question being whether it split recently from Giraffa or whether it has a long-separate ancestry. Its most obvious difference from Eurasian Miocene giraffes is the closeness of the two horn insertions. This might be considered adaptive for forest dwelling, except that the same character is found in Giraffa. There are wide disparities between fossil, molecular and other estimates for the divergence between Giraffa and Okapia. Molecular clocks suggest a divergence at 15 mya (Hassanin et al. 2012), while other reviewers have posited 8 mya (Mitchell & Skinner 2003). It can be surmised that giraffid ruminants survived by becoming larger than other ruminants, but their Eurasian radiation was curtailed after the end of the Miocene. In Africa both Okapia and Giraffa specialized in a capability for high-level browsing, one in forests and one in savannas, but sivatheres failed to survive to the present day. Giraffa was also found for a long time in India but the date of its extinction there is not known. It can almost be thought of as an unfortunate historical or evolutionary accident that Giraffidae have become confined to the continent of their probable origin. Alan Gentry 95

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Family GIRAFFIDAE

Subfamily GIRAFFINAE – Giraffe Giraffinae Gray, 1821. London Med. Repos. 15: 307.

Subfamily Giraffinae is a monotypic subfamily, represented by a single surviving species, the Giraffe Giraffa camelopardalis.

Genus Giraffa Giraffe Giraffa Brisson, 1762. Regn Anim., 2nd edn, pp. 12, 37.

The Giraffe, Giraffa camelopardalis, is the only extant representative of the genus Giraffa. Together with the only other living giraffid, the Okapi Okapia johnstoni, they represent the extant members of a previously more diverse group. Not long after the description of the species, Levaillant in 1790 described the southern giraffe as a separate species (Dagg & Foster 1982). This separation between northern and southern species

Giraffe Giraffa camelopardalis skeleton.

continued through the 1800s (Owen 1841, Lesson 1842; although see Ogilby 1836). Thomas (1901) separated the reticulated giraffe Giraffa reticulata as a separate species from all other forms of Giraffe, and this was maintained by Lydekker (1904). Subsequent authors followed either classification scheme: Dollman (1929) and Allen (1939) separated the reticulated giraffe as a separate species, while Stott (1959) maintained the northern versus southern species split. However, for much of the latter half of the twentieth century, only a single living species of Giraffe has been recognized. More recently, Brown et al. (2007) identified six genetically distinct lineages, with little evidence of interbreeding between them, and proposed that some may represent distinct species. The most obvious features of Giraffa are the long limbs and neck. Where paired cranial appendages occur in ruminants they are on the frontal bones in a lateral supraorbital position. In Giraffa the paired ossicones are dermal in origin and attach to the skull more posteriorly over the frontoparietal suture. These ossicones are straight with rounded ends and vary individually in form (Singer & Boné 1960). In extant Giraffes the parietal ossicones show strong sexual dimorphism (Dagg 1965, Seymour 2001). The lower incisors and canines are robust and arranged in a semi-circular arcade. The lower canine is typically bifid and occasionally trifid. The cheekteeth are variable in size between species, with premolars relatively complex. Basicranial and basifacial planes typically are not parallel and skull flexion varies individually in the species (Seymour 2001). The genus has an Asian origin, with the earliest known African specimens of Giraffa originating from fossil beds dating to the late Miocene and early Pliocene of Kenya and the early Pliocene of South Africa (Churcher 1978). Churcher (1978) described four extinct members of the genus Giraffa in Africa besides the extant species: G. jumae, G. stillei, G. gracilis and G. pygmaea. More recently, Gentry (1997) – following Harris (1987, 1991b) – accepted three extinct species, with G. gracilis included in G. stillei. Giraffa jumae is known from localities in East Africa and South Africa (though Harris [1976] gives Giraffa cf. G. jumae at Langebaanweg) dated from the early Pliocene to middle Pleistocene. The South African specimens are isolated teeth and ossicones, while the East African material is more substantial. The species is founded on a nearly complete skull and mandible with a significant proportion of the postcranial skeleton. Giraffa jumae specimens are typically larger than contemporary G. camelopardalis. While the dimensions of the skull are greater, the teeth are similar in absolute size in comparison between the two species. The ossicones of G. jumae originate directly behind the orbital rim, further forward than in G. camelopardalis, and extend back parallel to the plane of the skull. The parietal horns end

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Giraffe Giraffa camelopardalis skull outlines (bottom) compared with PlioPleistocene Giraffa jumae (top).

in knobs. No median horn has been identified in specimens assigned to G. jumae. No secondary bone deposition is apparent and the occipital ridge is not enlarged. In accord with the larger cranium, the mandible is more substantial than in G. camelopardalis, curving upwards through the diastema and downwards in the incisor region. Giraffa stillei, from the early Pliocene to early middle Pleistocene of East Africa, was first described as an Okapi based on dental characters and forelimb anatomy. Harris (1976) rediagnosed these specimens to Giraffa, according to the premolar morphology. Churcher (1978) suggested that the dental characters used to diagnose this taxon were highly individually variable and showed greater affinity to another giraffid genus, Palaeotragus. Churcher (1978) suggested that, without additional material, the affinities of the specimens currently assigned to G. stillei remained controversial. However, Harris (1987, 1991b) and Gentry (1997) accepted G. stillei and subsumed G. gracilis into it. Specimens of Giraffa gracilis have been found in East Africa and possibly in South Africa from beds dating from the late Pliocene to late Pleistocene. The limb bones and neck are of similar absolute length to G. camelopardalis but are more lightly constructed and show finer proportions in all parts of the skeleton. Dental characters also differ. The bases of the ossicones are oval in cross-section, smaller than in G. camelopardalis or G. jumae and oriented at the same angle as in the modern giraffe. Secondary bone deposition occurs over the ossicones. The area between the orbits is convex (Harris 1976) and may or may not have a median horn present. Churcher (1978) suggested that, despite generalizations about the relative size of G. jumae, G. camelopardalis and G. gracilis, the skeletal elements overlapped in size between these purported species making size alone an unreliable character for species identification. A Giraffa species with ossicones smaller and more delicate than those of G. gracilis and flattened on the posterolateral surfaces occurs in the early Pleistocene of East Africa. Giraffa pygmaea shows (presumed) sexual dimorphism with secondary bone deposition occurring in the (presumed) ??, increasing the proportions of the ossicones relative to the (presumed) //. Dental characters are typically giraffine, but differ from other species by their small size. Churcher (1978) considered this species to be inadequately defined for certain recognition as a separate species.

Giraffe Giraffa camelopardalis myology.

Many of the African Giraffa species are based on relatively little material and the status of some species may be equivocal. Many fossil specimens were originally attributed to G. camelopardalis and have subsequently been reassigned to the extinct species, bringing forward the earliest recorded fossils of the modern giraffe. Contemporary giraffes vary individually, sexually and geographically in size while skull morphology is also highly variable within the extant taxon. As many of the described differences (particularly in size) may represent extremes of continuous variation further fossil material is necessary to substantiate the status of the species within Giraffa. Mitchell & Skinner (2003) reviewed the origin and evolution of Giraffes, while Mitchell (2009) provides a review of the scientific study and classification of Giraffes. Russell Seymour

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Giraffa camelopardalis Giraffe Fr. Girafe; Ger. Giraffe Giraffa camelopardalis (Linnaeus, 1758). Syst. Nat., 10th edn, 1: 66. ‘Habitat in Æthiopia et Sennar’; identified as Egypt, in captivity at Cairo (Thomas 1911: 150); restricted to Sudan, Sennar, by Harper (1940: 322).

Dagg & Foster (1982).They identified six genetically distinct lineages (peralta, rothschildi, reticulata, tippelskirchi, giraffa and angolensis), with little evidence of interbreeding between them, and 11 genetically distinct populations. According to these authors, neighbouring subspecies as well as those that are geographically separated are essentially reproductively isolated, suggesting that some may represent distinct species rather than a single polytypic form. Synonyms: aethiopica, angolensis, antiquorum, australis, biturigum, capensis, congoensis, cottoni, giraffa, hagenbecki, infumata, maculata, nigrescens, peralta, renatae, reticulata, rothschildi, schillingsi, senaariensis, thornicrofti, tippelskirchi, wardi. Chromosome number: 2n = 30 (FN = 54). The 28 paired autosomes are mainly metacentric and submetacentric with one pair of acrocentric chromosomes, which are small; the large X chromosomes are submetacentric and the small Y chromosomes metacentric (Wallace & Fairall 1965, Gallagher et al. 1994).

Masai Giraffe Giraffa camelopardalis tippelskirchi.

Taxonomy  The Giraffe is here considered a polytypic species comprising eight subspecies (and see genus profile). Numerous subspecies have been recognized (e. g. Lydekker 1904), although most recent treatments have acknowledged nine (Ansell 1972, Dagg 1971, Dagg & Foster 1982), eight (Kingdon 1997), or six (East 1999, Grubb 2005). Based on pelage patterns, skull morphology and mitochondrial DNA analysis, Seymour (2001) described six or seven subspecies, and the existence of a distinct northern and southern clade. Considerable uncertainty surrounds the geographic and taxonomic limits of all described subspecies, as well as intergrades or hybrids between supposed subspecies (especially between reticulata and tippelskirchi in E Kenya; Kingdon 1979, Stott & Selsor 1981). Pelage patterning within subspecies highlights this uncertainty. As a definable characteristic, it is unreliable due to the high degree of individuality and variability within a population, ranging from albino and pure white through pale brown and unspotted to black (Mitchell & Skinner 2003, Fennessy 2004). Many of these variations exist within regionally definable patterns, although these boundaries are often far from clear. Brown et al. (2007) analysed mitochondrial DNA sequences and nuclear microsatellite loci from six of the nine subspecies defined by

Description  The tallest of all animals, the Giraffe is characterized by its greatly elongated neck and long limbs. Males, which are taller than //, range between about 4 and 5 m in height, with records of individuals up to 5.9 m (Shortridge 1934). Shoulders appear higher than the croup, a characteristic enhanced by the long spines of the thoracic vertebrae (see Adaptations). Neck fringed with a short, thick mane. Ears narrow and pointed, eyes large. Pelage short, with a highly characteristic pattern of large, irregularly shaped, dark patches separated by a network of light-coloured bands. This patterning, which extends from about the chin to the limbs, varies predominantly from chestnut-brown to nearly black. Kingdon (1979) considered that all Giraffe network patterns were products of interaction between two different, and genetically controlled, spot formation systems whereby light spots never occur in isolation, whereas dark spots are always isolated unless clustered within a single ‘island’ of ground colour. Leucistic Giraffes have been recorded (e.g. Tarangire N. P. and Rukwa in Tanzania, and Masai Mara in Kenya; C. Foley pers. comm.). The individuals (both ??) in Tarangire N. P. were predominantly white, though they had regular brown colouration on the lower legs and some black markings on the upper body (C. Foley pers. comm.). Colour and pattern also varies individually, being unique to each Giraffe (although local lineage characteristics can sometimes be recognized). Overall patterns, especially the dark patches, generally tend to darken with age. Towards the hooves the limbs are generally lighter in tone, but there are regional distinctions in the extent of patterned or unpatterned ‘socks’.The skin pigmentation is uniformly dark grey. Tail with black terminal tuft. There are no compound odoriferous glands. Two pairs inguinal nipples. The skull is long, sometimes more than 700 mm, and is characterized by extensive pneumatic sinuses. A characteristic feature of adults is the two or three short, blunt ‘horns’, more correctly termed ossicones, which rise from the top of the skull. These comprise a parietal pair, and a single median ossicone. Ossicones are covered with skin and hair, and are present in both sexes, but are thinner and with more prominent

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Giraffa camelopardalis

Lateral, palatal and dorsal views of skull of adult male Reticulated Giraffe Giraffa camelopardalis reticulata.

tufts of hair in // and young.The median ossicone, which originates primarily on the frontal bone and the posterior portion of the nasal bone, is less prominent in // (sometimes just represented by a bulge) and represents a secondary sexual male character (Spinage 1993). The Giraffe is the only ruminant with paired parietal horns present in the foetus. These ossicones begin as fibro-cartilaginous discs and are not attached to the skull (Owen 1849): they lie against the skull and present no obstruction at birth (Naaktgeboren 1969). They progressively ossify and enlarge, fusing to the skull later in life (about 4–4.5 years in ?? and 7 years in //) by way of secondary bone deposits (Spinage 1968). The median horn develops in the same manner as the parietal horns (Spinage 1993). Secondary bone deposits may accumulate on male skulls, producing further bony growths. Male skulls often develop rugosities that can resemble supernumerary horns, especially on the nuchal crest (see Adaptations). The permanent teeth erupt at about three years of age, except for the canines that erupt at around six years (Hall-Martin 1976). No dental abnormalities have been recorded (Colyer 1936, Miles & Grigson 1990). Geographic Variation  The treatment of subspecies presented here is provisional, and largely follows East (1999) except for separation of giraffa and angolensis (Seymour 2001, Fennessy 2004) and peralta and antiquorum (following Hassanin et al. 2007). G. c. peralta (Niger or West African Giraffe): Niger. Wider pale bands; the edges of the spots, though somewhat crenulate, are strongly delineated from the paler ground colour. Males have the strongly formed median horn (as in all northern giraffes), but it tends to be more cylindrical (as opposed to conical). Recognized as a distinct subspecies by Hassanin et al. (2007). G.c.antiquorum (including congoensis) (Kordofan Giraffe): N Cameroon, S Chad, Central African Republic, W Sudan and presumably NE DR Congo. The blotches, which often appear coarsely divided or constricted, extend below the hocks.

G. c. camelopardalis (including rothschildi) and cottoni (Nubian, Rothschild’s, or Baringo Giraffe): Ethiopia, Kenya, Sudan, Uganda. The blotches are more widely separated than in reticulata. The inner sides of the legs are unspotted, with the legs pure white below the hocks. Generally very large body size, especially ??. Although provisionally included in G. c. camelopardalis here, the form rothschildi may represent, and is often treated as, a distinct subspecies (e.g. see Brown et al. 2007). G. c. reticulata (Reticulated Giraffe): Ethiopia, Somalia, Kenya. The large, almost polyhedral patches are placed closely together with only a fine network of light colour dividing them. Patterning may be very dark in old animals. Generally smaller body size. Integrades with G. c. reticulata and G. c. tippelskirchi have been recorded (Stott 1959, Stott & Selsor 1981). G. c. tippelskirchi (Masai Giraffe): Kenya and Tanzania.The blotches are typically deeply dissected, forming all shapes of sharply differentiated leaf or stellate designs. The patterning always continues down to the hooves. G. c. thornicrofti (Thornicroft’s Giraffe): LuangwaValley, Zambia. Slightly stellate spots, which become oblong on the neck.The neck is usually lighter in colour than the body and the legs are fully patterned. G. c. angolensis (including infumata) (Angolan Giraffe): Namibia, SW Zambia, N Botswana, extreme W Zimbabwe; formerly in Angola, but now probably extinct. Considered as a distinct subspecies based on recent molecular evidence (Seymour 2001, Fennessy 2004, Hassanin et al. 2007). G. c. giraffa (including capensis and wardi) (Southern or Cape Giraffe): South Africa, S and SE Zimbabwe and SW Mozambique. The blotches, which extend down to the hooves, are more or less round. The colouring may be very dark. Generally large body size. Similar Species  Given its distinctive features, the Giraffe is unlikely to be confused with any other animal. Distribution  Endemic to Africa. Historical Distribution Widespread in N and W Africa, including the Sahara, until the Neolithic. Rock drawings, petroglyphs and skeletons confirm their presence on the banks of the Nile and the plateau of Messak until 4000 BC and later in the Western Sahara and Mauritania (Le Quellec 1999). Giraffes were historically distributed widely throughout the Sahelian regions of West Africa from C and E Senegal, N Guinea and SE Mauritania through C Mali to C/S Chad, N Central African Republic, Sudan (west of the Nile) and NE DR Congo. Nubian/Rothschild’s Giraffes occurred in SW Eritrea and W and SW Ethiopia, south through Sudan, east of the Nile R., Uganda and W Kenya. Reticulated Giraffes ranged from S Ethiopia south through Kenya, east of the Rift, adjacent S Somalia to north of the Tana R. Masai Giraffes occupied S and E Kenya and much ofTanzania north of the Rufiji R.Thornicroft’s Giraffes occurred only in the Luangwa Valley in Zambia (Ansell 1978). There is one exceptional occurrence of a Giraffe being recorded in Malawi (a single individual killed in Sep 1976), presumably a vagrant animal from the Luangwa Valley (see Ansell & Dowsett 1988). The historical ranges of the two southern subspecies (giraffa and angolensis) covered Namibia (including the semi-desert areas of Kaokoland in the north-west) and adjacent parts of S Angola (in two 99

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2

1

3

5 4 6 7 8

9 Giraffa camelopardalis

well-separated populations in the SW and SE), N and C Botswana, SW Zambia, W and S Zimbabwe, S Mozambique, and N and NE South Africa (East 1999). Although some have proposed that Giraffe never occurred south of the Komati R. in Swaziland or KwaZulu– Natal (Goodman & Tomkinson 1987), Giraffe were probably present 1000 BP in KwaZulu–Natal, but had died out or been extirpated by ca. 220 BP (Cramer & Mazel 2007). Indigenous San drawings suggest that Giraffe once occurred throughout parts of S South Africa: near Queenstown, in the Tarka area, near Graaff Reinet and on the Tsomo R. around 1650. However, over a period of 300 years, the limit of their range was reduced by nearly 900 km (Dagg & Foster 1982). Current Distribution Today no Giraffes survive in North Africa, though some may have remained in Morocco as late as 600 AD (Schomber & Kock 1961). The beginning of the twentieth century marks the sharp decline of Giraffes in the West African region. The only surviving population in West Africa is in SW Niger, an area of approximately 15,000 km2 delineated as the ‘giraffe zone’ in a broader Biosphere Reserve. This population co-habits with subsistence agricultural communities and seasonally migrates throughout its range in search of available forage. During the wet season (Jun–Sep), the animals are located on the Koure and Fandou Plateau, but during the dry season they move to the Dallol Bosso North (a sandy agricultural region with permanent water in numerous pools) and the regions of Loga and Dogondoutchi (Ciofolo 1995, 2002, Boulet et al. 2004). In 1996, five Giraffes were observed in the Ansongo-Menaka Partial Faunal Reserve in Mali, on the border of Niger (Le Pendu & Ciofolo 1999, East 1999); however, this population is assumed to be extinct. The Kordofan Giraffe is still present in N Cameroon, S Chad and Central African Republic (mainly Manovo–Gounda–St Floris N. P.). East Giraffe Giraffa camelopardalis distribution interpreted as four major types with mixed or intermediate populations in between (reproduced from Kingdon 1997).

right:

10

11 12

Key to Giraffe patterns: 1. Nubian Giraffe G. c. camelopardalis Ca 2. Reticulated Giraffe G. c. reticulata (dark morph) Re 3. Reticulated Giraffe G. c. reticulata (light morph) Re 4. Kordofan Giraffe G. c. antiquorum P 5. Hybrid Reticulated Giraffe G. c. reticu­ lata ∞ Masai Giraffe G. c. tippelskirchi (between Tana & Galana Rivers) X

6. Masai Giraffe G. c. tippelskirchi T 7. Nubian Giraffe G. c. camelopardalis (dark morph) Co 8. Nubian Giraffe G. c. camelopardalis (Rothschild’s Giraffe) Ra 9. Masai Giraffe G. c. tippelskirchi (dark morph) T 10. Angolan Giraffe G. c. angolensis A 11. Southern Giraffe G. c. giraffa G 12. Masai Giraffe G. c. tippelskirchi T

above: Giraffe patterns display both regional characteristics as well as individual variation. Patterns shown on the page opposite come from localities indicated in the map (below) and correspond to the key (above).

Somali arid (camelopardalis and reticulata) Ca, Re Saharan (peralta) P N savanna (cottoni) Co S savanna (tippelskirchi, giraffa, etc.) T, G, A Cameroon (hybrids?) Rothschild (hybrids?) Ra Galana hybrids X

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Giraffa camelopardalis

Giraffe Giraffa camelopardalis (see key opposite).

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of this, there is a paucity of information, with no information available from Sudan, west of the Nile; they were eliminated from Radom N. P. and were not recorded at all from Southern N. P. or Shambe G. R. during recent surveys (Fay et al. 2007).Their main remaining nucleus is said to be Garamba N. P. in NE DR Congo (East 1999). The Nubian/Rothschild’s Giraffe survives in SE Sudan (in Boma N. P. and Bandingilo Reserve, but extinct in Dinder N. P. since 1985), SW Ethiopia, Uganda (Murchison Falls N. P. and Kidepo Valley N. P.) and Kenya, predominantly in protected areas; it is now extinct to the north and in Eritrea. The Reticulated Giraffe still occurs in S Somalia and Ethiopia, and occupies some of its historical range in N Kenya, mainly outside protected areas. Likewise, the Masai Giraffe also occupies parts of its historical range, although populations in Tsavo East N. P. in Kenya and Serengeti N. P. in Tanzania, as well as other protected and unprotected areas, have suffered declines; Masai Giraffe have been introduced to Rwanda (East 1999). Thornicroft’s Giraffe is still found in large numbers in the Luangwa Valley of Zambia (East 1999). Having been reintroduced to many parts of the range from which they were eliminated, Angolan and Southern Giraffes are currently common both inside and outside a number of protected areas in Namibia, Botswana, Zimbabwe and South Africa. In Angola, the Giraffes that historically occurred in the south-west of the country had disappeared completely by the 1940s, while the population in the south-east is now assumed to be extinct (Crawford-Cabral &Veríssimo 2005). East (1999) reported that the Angolan Giraffe still survived in small numbers in Sioma Ngwezi N. P. in SW Zambia. In Mozambique, a few individuals of the Southern Giraffe still occur in Coutada 16, adjacent to Kruger N. P. (J. Anderson pers. comm.). Giraffe have been introduced to Swaziland with stock from South Africa and Namibia (although Namibian animals have done poorly; see Monadjem et al. 2003), and have also been introduced to a number of reserves in South Africa, including in KwaZulu–Natal and the Eastern Cape. Habitat  Giraffes are typically associated with Acacia, Commiphora and Combretum savannas. They occur marginally in the miombo Brachystegia woodland, while an isolated population (G. c. thornicrofti) occurs in the Luangwa Valley (where mopane, Acacia and Combretum are widespread), and in the Isoberlina woodland in Cameroon. Giraffes are today absent from true deserts and from rainforests; they also are absent from coastal savannas and large savanna mosaics (such as Lopé, in Gabon). Nevertheless, in NW Namibia they reside throughout the northern Namib Desert, where annual rainfall is less than 100 mm; here, they are amongst the best adapted ungulates in this desert environment (Scheepers 1992, Fennessy 2004). Giraffes in the northern Namib Desert rely on the riparian forests along the banks of ephemeral rivers, which include Acacia, Faidherbia, Combretum and Commiphora species, and surrounded by open gravel plains and sand dunes (Fennessy 2004). East of the Nile, in S Sudan, Giraffes are never found in the permanently flooded areas of the Sudd, but aerial surveys (Mefit-Babtie 1983) showed that they were widely distributed over the entire floodplain throughout the year. During the height of the flood season, Giraffes were relatively few in the better drained woodlands yet, paradoxically, showed a perceptible seasonal preference for quite deeply flooded areas close to the permanent swamps. Observation revealed that the attraction for Giraffes at this time was vegetation growing on large termite

mounds, which are exceptionally abundant in the seasonally flooded grasslands (or ‘Toic’) and provide islands of soil raised above the surrounding floodwaters (J. Kingdon pers. comm.). In addition to climbers, such as Ipomea, the foliage of Balanites, Acacia, Cassia, Grewia and Ziziphus trees are all in an actively growing state at this time. Freezing temperatures coupled with wind and rain can kill Giraffes (Dagg & Foster 1982); however, they can tolerate temperatures in excess of 40 °C (e.g. in Namibia and Niger). Spatial and temporal environmental factors, e.g. rainfall, affect their distribution as well as influence their seasonal movements. In Niger, Giraffes aggregate during the wet season where forage resources are limited (Le Pendu & Ciofolo 1999), while in East Africa, they commonly disperse during the wet season, aggregating near rivers in the dry season (e.g. Leuthold & Leuthold 1978a, Estes 1991a). Abundance  East (1999) estimated the total population at about 140,000 animals, predominantly in areas dominated by Acacia woodlands and shrublands, including 3500 Niger and Kordofan Giraffes (peralta and antiquorum combined), approximately 500 Nubian/Rothschild’s Giraffes, 36,000 Reticulated Giraffes, 60,000 Masai Giraffes, 1200 Thornicroft’s Giraffes (more numerous than at any time during the last 50 years) and more than 40,000 Angolan and Southern Giraffes (angolensis and giraffa combined). The most recent estimates put the total population at less than 100,000 animals (International Giraffe Working Group pers. comm.); efforts are underway to census the continent’s populations more accurately (Fennessy 2007). The population in Niger (estimated to number 79 animals in 1999; Ciofolo et al. 2000) has since increased (Boulet et al. 2004, Suraud & Dovi 2006), with 193 animals photographed and identified in 2008 (Suraud 2009). Giraffe density appears correlated with the distribution and density of high-quality forage: where appropriate food plants are abundant, Giraffes can live at densities of up to 2.6/km2 (Pellew 1983a, b, 1984b), although Pratt & Anderson (1982) reported extraordinarily high densities in the small Arusha N. P. (3.96 ind/km2). Across its range Giraffe densities can vary between populations with 0.7/km2 observed in Kruger N. P. (Hirst 1975) and 0.85/km2 in Lake Manyara N. P. (Van der Jeugd & Prins 2000). On the other hand, where the abundance of high-quality forage is scarce, densities are markedly lower: for example, in Southern N. P. in Sudan, where Giraffes probably no longer occur, the population density was estimated at 0.05/km2 (0.03–0.1/km2) (Boitani 1981) and in Tsavo East N. P. at 0.2/km2 (Leuthold & Leuthold 1978a).The lowest densities have been recorded in the desert-dwelling populations of Niger and Namibia (0.01/km2; Fennessy et al. 2003, Fennessy 2004). Adaptations  The Giraffe displays a range of remarkable physiological and morphological adaptations, many linked directly to its standing as the world’s tallest animal, perhaps foremost of which is its long neck.The osteology of the neck has been studied by anatomists since Owen (1841). Brownlee (1963) summarized theories concerning its evolution, and more recent contributors include Solounias (1999), Mitchell & Skinner (2003) and Simmons & Altwegg (2010). As is the case in all mammals (except manatees Trichechus spp. and sloths Bradypus spp. and Choloepus spp.), there are seven cervical vertebrae. Solounias (1999) suggested that neck length could be attributed to the presence of an 8th cervical vertebra, but as this vertebra supports a rib it must

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Giraffe Giraffa camelopardalis.

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by definition be a T1 (Mitchell & Skinner 2003, Badlangana et al. 2009). In addition, it would seem that the role of articulation between the thoracic and cervical sections of the vertebral column normally performed by C7 is at least partially taken over by T1 (Mitchell & Skinner 2003). Elongation of the neck is the result of uniform lengthening of the entire cervical series, including C7 (Badlangana et al. 2009), which is not a foetal process but occurs after birth (van Sittert et al. 2010). The neck vertebrae have characteristically pronounced opistho­ coelous (or ball-and-socket) joints. Backward retraction of the base of the cervical column (so that it emerges well behind the sternum) and upward bowing of the thoracic vertebrae permits the forelegs to carry a high proportion of the head and neck weight, creating the ‘chesty’ profile so typical of Giraffes. The Giraffe has 14 thoracic vertebrae (as noted above, the first thoracic somewhat resembling a cervical vertebra), five lumbar vertebrae and four sacral, although a vertebra may be lost to the tail, which is variable, ranging from 12 to 20. The pelvis is shorter than in most other Pecora and the sacral region is shorter compared with some bovids. Other large species like African Buffalo Syncerus caffer and ox have shorter thoracic regions, with 13 vertebrae, a longer lumbar region and similar length sacral. Camels have 12 : 7 : 4, thus their lumbar region is longer than the Giraffe’s and the thoracic shorter. The Okapi Okapia johnstoni has 24 in the ratio 14 : 5: 5, thus the sacral region is longest. In terms of length of the cervical vertebrae relative to overall vertebral column length, the neck vertebrae of the Okapi comprise 35% of the total vertebral column, while those of the Giraffe comprise 54% (Badlangana et al. 2009). The Okapi is approximately 55% shorter in height at the shoulder than the Giraffe, about 1.65 m compared with 3 m (C. Spinage pers. comm.). The spines of the thoracic vertebrae are markedly elongated, forming points of origin for some of the neck muscles and for the ligamentum nuchae (Mitchell & Skinner 2003). These spines also enlarge the purchase (and therefore the swing of the entire forelimb) for muscles attached to the shoulder blade, notably the trapezius and rhomboid. Hall-Martin et al. (1977b) estimated weight for the head and neck combined at about 250 kg, and leverage of such an extended mass is facilitated by several adaptations, including the thoracic spinous processes, the presence of a large and extensive frontal cranial sinus (adding volume, but not mass to the skull; see Badlangana et al. 2011) and a reduction in the size of ossicones, compared with some extinct shorter-necked giraffe species.Van Schalkwyk et al. (2004) noted that cervical vertebrae mass decreases towards the head (largely a result of the size of each vertebra as the density actually increases), indicating that the bulk of the head and neck weight is supported at the base and that the distal end of the neck and head is comparatively light and more manoeuvrable than would otherwise be the case. Adaptations that reduce the mass and weight of the head and neck apply much less to adult ??. Their principal intra-specific weapon is the head and its offensive/protective protuberances, which is swung by the neck like a pendulum (Owen 1841, Kingdon 1979). The sometimes violent ‘necking’ (see Social and Reproductive Behaviour) has a correlation with accumulations of secondary bone growth and heavier skull weight for ?? (up to ~15 kg; Dagg 1965, Seymour 2001), which suggests that male rivalry has resulted in selection for greater head weights (Simmons & Scheepers 1996). Osteoblasts build up and reinforce ossicones, presumably under the influence of male hormones, but osteoblasts also become active in

direct response to trauma. In the latter case they plaster over cracked bones and, in response to bruising, form encrustations on the outermost margins of the frontal bones (which overhang and thus help protect the orbits) along the exposed bridge of the nose (forming lumps on the nasal bones) and, most prominently, on the topmost ridge of the occipital bone. Understandably, occipital or ‘nuchal’ knobs are often mistaken for ‘ossicones’ and such giraffes are often described as ‘five horned’. An interesting detail in male Giraffe skulls is the occasional occurrence of bony encrustations on only one side of the head, probably resulting from repeatedly striking with that side of the head (J. Kingdon pers. comm.); indeed, Lydekker (1904) described a subspecies (cottoni) from a single specimen citing an ‘orbital horn’ over the right eye. Males normally appear to be ‘ambidextrous’, but they also have a choice as to the direction in which they will deliver or absorb blows. When they first engage, the contestants can either stand side by side or face in opposite directions. In the side-by-side position, one fighter swings to the right, one to the left. In the second choice, the same side is involved in both animals. In a small sample of 13 contests, J. Kingdon (pers. comm.) observed six pairs facing in opposite directions, four with their right shoulders engaged, two with the left. It would be interesting to learn whether advantages or disadvantages could accrue for ?? with a particular preference and if there are regional patterns of preference. Mitchell & Skinner (2003) discussed the possible advantages conferred by height, including protection from predation, increased vigilance, an enlarged surface area for thermoregulatory heat loss, and, in ??, sexual dominance and access to food. These authors considered that the hypothesis that Giraffe height evolved as a response to the advantages conferred by feeding height stratification holds true only for adult ??. For example, Du Toit (1990a) compared preferred heights of Giraffes with Greater Kudu Tragelaphus strepsiceros, and found that Giraffes tended to feed at heights of 1.7– 3.7 m (?? feeding higher than //) and Greater Kudu at heights of about 1 m, and occasionally up to 2 m. Greater Kudu were, therefore, competitive with female (and young) Giraffes. Other dietary studies have noted that Giraffes frequently feed well below their maximum foraging height (Leuthold & Leuthold 1972, Pellew 1984a, Young & Isbell 1991). None the less, analysis of the feeding competition hypothesis suggests that a long neck confers at least some advantage when lower leaves of trees have been eaten by these shorter browsers (Cameron & du Toit 2007). An alternative theory suggested by some authors (Churcher 1978, Simmons & Scheepers 1996) is that the long neck may have evolved in response to the sexual advantage it confers, with ?? using their necks and heads to achieve sexual dominance. However, this ‘necks for sex’ theory was not supported by empirical data that showed that morphological differences between the sexes are minimal and that any differences that do exist can be accounted for by the larger realized mass of ?? (Mitchell et al. 2009a, van Sittert et al. 2010; but see Simmons & Altwegg 2010). In reviewing these two primary competing hypotheses, Simmons & Altwegg (2010) noted that the main challenge for the competing browser hypothesis is to explain why Giraffes have remained about 2 m taller than competing browsers for over one million years given the costs involved, while the sexual selection hypothesis fails to account for the long neck of female Giraffe (even though they are shorter than ??).

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Giraffe Giraffa camelopardalis rothschildi head of subadult female.

Some (Wright 1871, Brownlee 1963) have suggested that the Giraffe’s long neck evolved because it assists the detection of predators: Giraffes have excellent sight, and their auditory and olfactory senses are acute. They can detect danger and conspecifics from a greater distance than other savanna species and seem to stay in contact by sight at a distance of over one kilometre (especially in open environments) thanks to their height advantage (Dagg & Foster 1982). Predator avoidance may also explain the coat markings in Giraffe, which could serve as camouflage for calves during the hiding stage, as calves are vulnerable to high rates of predation during the first few months of life (Langman 1977) (see Reproduction and Population Structure). Social factors are a significant modifier of vigilance in Giraffes: in Kruger N. P., bulls scan the most when they are in groups with larger bulls and least when they are with smaller bulls. Conversely, predation risk does not appear to be a significant modifier of vigilance: cows do not change their vigilance behaviour when alone or accompanied by calves (Cameron & du Toit 2005). In addition to the extension of the neck, there is also elongation of the limbs. However, the elongation of the limb bones is independent of a corresponding increase in their diameter. The more distal limb bones, particularly the metapodials, but not the humerus or femur, are extraordinarily elongated and slender for an animal of such size, suggesting that the increase in length must be compensated for by an increase in strength (Mitchell & Skinner 2003). The latter authors posited that this increase in strength was facilitated by the high density of the lower limb bones, such that about 80% of all skeletal calcium is in the leg bones, with the radius/ulna and metacarpal bone having the highest relative density of any bone at 150% and 178%, respectively. That the weightbearing bones are denser than others is supported by the study of Van Schalkwyk et al. (2004), who also propose that, in addition to increased density, Giraffe limb bone strength is achieved by the bones being much straighter and having much thicker walls than in other artiodactyls. However, Van Schalkwyk et al. (2004) also presented evidence that the skeleton of the Giraffe is not unique with respect to bone density. Mitchell et al. (2005) have since shown that the Giraffe skeleton contains three times more absolute amounts of calcium and phosphorous than is found in the African Buffalo skeleton. This translates into a 1.5- to 2-fold higher calcium requirement for Giraffes, with which they seem to cope effectively by selecting for calcium-rich, dicotyledonous, browse. However, sources to meet phosphorous requirements are less obvious and a

seasonal deficiency of phosphorous is a probable cause of instances of observed osteophagia (see Foraging and Food). Both neck and limb elongation requires an unusual cardiovascular and respiratory system. Physiological adaptations of the circulatory system have been investigated in detail by numerous authors (e.g. Amoroso et al. 1947, Lawrence & Rewell 1948, Goetz 1955, Goetz & Keen 1957, van Citters et al. 1966, Kimani 1983, Badeer 1986, 1997, Hargens et al. 1987, Kimani et al. 1991a, b, Pedley et al. 1996, Mitchell et al. 2006, 2008, 2009b). The hydrostatic pressure exerted by gravity on the column of blood in the neck (the heart is ~2 m away from the head; Mitchell & Skinner 1993) necessitates an average systemic blood pressure of ~200 mm Hg compared with the norm of 100 mm Hg in other terrestrial mammals (see Mitchell & Skinner 2009 and references therein). Although widely reported as having a very large heart relative to overall mass, heart mass in Giraffe is only 0.5% of body mass – the same as that in other mammals (Mitchell & Skinner 2009). Instead of cardiac enlargement or an increase in cardiac output, cardiac hypertrophy of the left ventricular and interventricular heart wall muscles (their thickness is linearly related to neck length) is the key to maintaining cerebral blood flow. Further, hypertrophy of the arteries and arterioles at or below the level of the heart helps control blood flow to the organs and, during drinking, the thick-walled arteries help prevent blood from rushing to the head (Mitchell & Skinner 2009). Valves in the jugular vein direct the large amounts of blood returning to the heart via the inferior vena cava into the right atrium, and prevent it regurgitating into the jugular (Mitchell et al. 2009b; contra Hargens et al. 1987). When a Giraffe raises its head, a momentary pause during head lifting and intense extracranial vasoconstriction helps to prevent fainting (Mitchell et al. 2008). Microcirculation haemodynamics, the thick skin and thickened arteries, an autonomic nervous system and innervation of the limbs help prevent oedema (Hargens et al. 1987, Pedley 1987). Other physiological adaptations include an efficient nasal cooling system that regulates brain temperature (up to 3 °C lower than body temperature) and reduces respiratory water loss, while respiratory dead space volume is minimized by a variable diameter of the trachea (Mitchell & Skinner 1993). Whole body thermoregulation might be passively maintained by skin patches acting as heat dissipating windows facilitated by vasodilation of the blood vessels beneath the dark skin surface (Mitchell & Skinner 1993, 2004). In Etosha N. P., Namibia, Giraffes have been observed to face the sun when the temperature is high, probably in order to diminish the surface exposed (Kuntzsch & Nel 1990). Many of the extant Giraffe’s adaptations are essentially those of a very large animal not just a very tall one. While their height has facilitated several detailed physiological adaptations, their size also increases nutritional demands. For example, ?? require some 20 g of calcium per day; by comparison, a human weighing one-tenth the weight of a giraffe has a daily calcium requirement of one-fortieth (Mitchell & Skinner 2003). In order to fulfil their daily nutrient requirements, Giraffes must consume a large amount of browse in the form of legumes. Their molars are heavy, brachydont and selenodont. The lower incisors combine with the bi- or tri-lobed canines to form an arc to comb the leaves and bend/break the thorns off shoots. Horny papillae help protect the lips and tongue from thorns, and thick saliva protects the tongue and buccal cavity. 105

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Giraffe Giraffa camelopardalis myology of head.

The tongue is long and mobile, up to 45 cm, allowing the Giraffe to precisely select food items. Although Giraffes drink when water is available, they can survive for a long time without water. In the north of the Namib Desert, Scheepers (1992) observed Giraffes drinking twice during the course of six years of observation, and J. Fennessy (pers. comm.) only ten times during a five-year study period. Giraffes seem to demonstrate a total lack of dependence on available bodies of water and get their water from preformed water in food and dew on the plants (Scheepers 1992, Fennessy et al. 2003, Fennessy 2004). A gall bladder may or may not be present. Giraffes present a biphasic activity budget: energy consumptive activities like feeding and walking occur more post-dawn and pre-dusk, while energy conserving activities such as resting and ruminating occur more during the hottest period of the day (Leuthold & Leuthold 1978b, Pellew 1984a, Fennessy 2004, 2009). Contrary to most other quadrupeds, Giraffes swing both legs on the same side at almost the same time during their ‘pacing’ walk (this pattern of limb movement is abandoned when the Giraffe breaks into a gallop). A galloping Giraffe can reach 56 km/h, young running faster than adults. Their long legs enable them to clear fences 1.5 m in height (Dagg & Foster 1982). Giraffes emit a scent that can be detected by keen-nosed humans over considerable distances.Wood & Weldon (2002) analysed extracts of hair samples from adult Reticulated Giraffe ?? and // and found two highly odoriferous compounds (as well as a number of other major compounds) that appeared to be primarily responsible for the Giraffe’s strong scent. They suggested that these compounds may deter microorganisms or ectoparasitic arthropods, as most of these compounds are known to possess antibacterial or fungistatic properties against mammalian skin pathogens or other microorganisms. The levels of p-cresol, one of the compounds detected, in Giraffe hair are sufficient to repel some ticks. Foraging and Food  Giraffes are selective browsers, feeding predominantly on leaves and buds on trees and shrubs. Acacia species form the bulk of their diet throughout their range, as well as species of the genera Balanites, Commiphora, Detarium, Boscia, Combretum, Ziziphus and Grewia. However, many other species are eaten: at least 25 in Willem Pretorius G. R., South Africa (Kok & Opperman 1980), 29 in the Hoanib R. region, N Namibia (Fennessy 2004), 45 in Serengeti N. P. (Pellew 1984a) and in Niger (Ciofolo & Le Pendu 1998), 69 in Tsavo East N. P. (Leuthold & Leuthold 1972) and 77 in the Middleveld of Zimbabwe (Lightfoot 1978). Seasonally, they feed

on a range of species and plant parts, including herbs (e.g. Hibiscus asper) and climbers and vines (species of the genera Ipomea, Cissus). The proportion of grass species in the diet is very low. Giraffes are highly selective with regards to the plant’s growth stage and phenology. The diet varies according to the season, influenced by the availability of plant species and their growth stage (Hall-Martin 1974, Ciofolo & Le Pendu 2002, Fennessy 2004). Across their range this involves small-scale seasonal movements: in southern and East Africa, Giraffes often feed on deciduous trees, shrubs and vines during the wet season, and on evergreen species, near streams and rivers, during the dry season (Leuthold & Leuthold 1972, Kok & Opperman 1980). In N Tanzania, the ?? move to key resource areas during the dry season (Van der Jeugd & Prins 2000). In West Africa they feed on the shrubby acacias in the tiger bush during the wet season, and on the woody species of a sandy agricultural region in the dry season (Ciofolo & Le Pendu 2002). Pellew (1984a) found that the diet of adult // was nutritionally richer than that of bulls that consumed significantly higher proportions of fibres and lignin; furthermore, species-selection and the nutritional quality of diet were not obviously related and energy budgets suggest that Giraffes can maintain a positive energy balance during most stages of the female reproductive cycle (Pellew 1984c). In Niger, nursing // seem to avoid high levels of tannins even though it means giving up the highquality forage preferred by the ?? (Caister et al. 2003), while in N Namibia // avoided areas of human populations and thus forego richer nutrient food sources in preference for safety (Fennessy 2004). Pellew (1983a) demonstrated that, when Giraffes are not too numerous, their impact can actually stimulate shoot production in Acacia species, which soon declined when the browsing stimulus was withdrawn.Young & Okello (1998) experimentally excluded Giraffes and Savanna Elephants Loxodonta africana from access to Acacia, thereby reducing consumption of new shoots of the higher branches by 63%, which also induced the growth of longer spines. Pellew (1983a) suggested that Acacia species have evolved a high resilience to browsing.

Twigs of Acacia drepanolobium before and after Giraffe Giraffa camelopardalis browsing (plus gall ants). right: Giraffe Giraffa camelopardalis tongue pulling twig into mouth (from film). left:

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Coe (1998) found that the pods of A. eriobola were significantly larger above the browse-line for those trees that had been exposed to Giraffe feeding. In Ithala G. R. (South Africa), some Acacia species such as A. caffra and A. karoo showed high mortality attributable to Giraffe browsing while others including A. tortilis did not seem to be affected (Bond & Loffell 2001). In Kruger N. P., Giraffes were highly destructive and detrimental to the overall fecundity of A. nigrescens during the season examined (Sep), resulting in a significantly reduced fruit set at heights accessible to Giraffes (Fleming et al. 2006). Carnivorous ants that are symbiotic with some Acacia species reduce the amount of time that young Giraffes can spend browsing on any one plant (Madden &Young 1992). On the other hand, Acacia seed consumption by Giraffes favours seed dispersal into non-shaded habitats and enhances the potential for seed germination because digestion processes decrease seed infestation by bruchids (Miller 1994, 1996). It has also been suggested that Giraffes play a role in pollination; for example, Du Toit, J. T. (1990b, 1992) posited that Giraffes in Kruger N. P. may be providing a pollination service for Acacia nigrescens, although empirical evidence suggests that Giraffes are only flower predators (Fleming et al. 2006). Giraffes sometimes lick soil near termite mounds and chew bones (osteophagia) (Western 1971,Wyatt 1971, Leuthold & Leuthold 1972, Hall-Martin 1975, Ciofolo & Le Pendu 1998). Although Western (1971) maintained that this behaviour was unlikely to appreciably offset any mineral deficiency, Langman (1978) suggested that osteophagia might mitigate seasonal deficiencies or imbalances in calcium, phosphorus and/or magnesium, and Mitchell et al. (2005) predict that bone chewing may be a response to phosphorous deficiencies. Nesbit Evans (1970) also observed a Giraffe eating the stomach contents of a Common Eland Tragelaphus oryx carcass, and Western (1971) reported them scavenging on the carcass of a Grant’s Gazelle Nanger granti. Feeding is the dominant diurnal activity of Giraffes and differs significantly between sexes, season and populations. Females spend more time feeding than ?? in Serengeti N. P. (72% cf. 55%; Pellew 1984a) and in Tsavo East N. P. (Leuthold & Leuthold 1978b), while bulls spend more time feeding in N Namibia: 59% cf. 51% (Fennessy 2004). In Lake Manyara N. P., solitary ?? spent more time feeding than ?? in groups; Giraffes on average spent only 35% of their time on browsing (Van der Jeugd & Prins 2000). In Niger, time spent browsing doubles during the dry season compared with the wet season: 46% cf. 23%, respectively, probably due to variation in forage quality and its spatial distribution (Ciofolo & Le Pendu 2002). The other principal diurnal activities are walking, resting, ruminating and, to a lesser extent, vigilance. According to Ginnett & Demment (1997), there is no sex difference in rumination time but ?? spend more time in activities other than foraging and rumination, such as walking. Giraffes are also active at night, but feed significantly more during moonlit nights (34% cf. 22%, ?? versus //, respectively) and ruminate more during dark nights (49% cf. 40%, ?? versus //, respectively) (Pellew 1984a). Social and Reproductive Behaviour  Giraffes are sociable, but herd composition is fluid, unstable and highly variable, comprising any combination of sex and age classes at any given time and rarely consisting of the same individuals for any length of time (e.g. Pellew 1984b, Le Pendu et al. 2000, Van der Jeugd & Prins 2000, Fennessy 2004). Mean herd sizes are varied, often ranging between 2.6 and 8

Giraffe Giraffa camelopardalis mating behaviour.

(Dagg & Foster 1982), depending on population density and food availability: Katavi N. P. (Tanzania), 3.6 (Caro 1999a), Luangwa Valley, 3.6 (Bercovitch & Berry 2010; and see Berry 1973), NW Namibia, 3.8 (Fennessy 2004), Tsavo East N. P., 4.1 (Leuthold, B. M. 1979), Lake Manyara N. P. (Tanzania), 4.8 (Van der Jeugd & Prins 2000), and Niger, 6.9 (Y. Le Pendu & I. Ciofolo unpubl.). Maximum herd size also varies between populations (for example, from 13 to 47), although as many as 240 individuals have been observed in a herd (Pellew 1983b). Males tend to be solitary or associate in bachelor herds, becoming increasingly solitary as they mature, and often wandering between herds monitoring the reproductive status of //. Giraffes are not territorial, but occupy home-ranges that vary in size according to sex/age classes, competition, forage and resource availability (Leuthold & Leuthold 1978a, Le Pendu & Ciofolo 1999, Van der Jeugd & Prins 2000). In Tsavo East N. P., male and female mean home-range sizes were very similar (164 km2 and 162 km2, respectively), but smaller during the dry season than during the wet season (Leuthold & Leuthold 1978a). Seasonal home-ranges were exclusive in Niger where their mean size during the dry season (90.7 km2) was twice as large as that during the wet season (46.6 km2). Other mean annual home-range sizes recorded for adult // (using minimum convex polygon method) were 41 km2 in Timbavati Private N. R., South Africa (Langman 1973), 68 km2 in the Luangwa Valley (Berry 1978), 200 km2 in N Namibia (Fennessy 2009), 285 km2 in Kruger N. P., South Africa (du Toit 1988) and 367 km2 in Niger (Ciofolo & Le Pendu 1998). Home-ranges recorded for ?? are: 37 km2 in Timbavati Private N. R. (Langman 1973), 82 km2 in Luangwa Valley (Berry 1978) and 842 km2 in Niger (Ciofolo & Le Pendu 1998). The largest home-ranges recorded for ?? are for animals living in desert-dwelling populations: 1950 km2 and 1627 km2 for two bulls in N Namibia (Fennessy 2009) and 1564 km2 in Niger (Le Pendu & Ciofolo 1999); the largest cow home-range reported was also in Niger (1379 km2; Le Pendu & Ciofolo 1999). Giraffes in Niger have been observed to exhibit the longest range of movement, 107

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up to 300 km (Le Pendu & Ciofolo 1999). The longest linear range of movement measured in other populations was 86 km in the Luangwa Valley (Berry 1978), 70 km in N Namibia (Fennessy 2009) and 50 km in Tsavo East N. P. (Leuthold & Leuthold 1978a).The mean daily linear movements of two adult bulls (equipped with a GPS satellite collar) in N Namibia varied between 4.3 and 7.5 km (Fennessy 2009). Males exhibit flehmen when testing the reproductive status of //. If the // are receptive, dominant ?? actively pursue and fend off competitors. Often such behaviour results in ‘necking’ bouts, which are assumed to be dominance-driven (Coe 1967). Males with thicker, longer necks appear dominant during violent necking encounters with other ??, and subsequently are more successful in courting // (Pratt & Anderson 1982, 1985, Simmons & Scheepers 1996). Serious fights are rare, but when observed they have been brief and violent; Downey (in Kingdon 1979) saw one bull concussed during one such fight, whereupon the fallen animal was hammered with both head and hooves while lying prostrate. Dagg & Foster (1982) also reported an instance where a bull was knocked unconscious and took 20 minutes to recover. Association between adult ?? was random in Lake Manyara N. P., but // showed clear non-random association (Van der Jeugd & Prins 2000). The ? follows the oestrous / closely, sometimes resting his head on her rump and/or butting her gently with his horns. Males exhibit laufschlag (the ? tapping between or alongside the hindlegs of the / with his forelegs), causing the / to move slightly forward. During copulation the bull stands with head held high, mounting by sliding his forelegs loosely onto her flanks, and then standing bolt upright while at the same time delivering an ejaculatory thrust that also moves the cow forward (Estes 1991a). Females usually leave the herd to calve. Young are born while the cow is still standing, her back legs slightly bent so as to lessen the fall, which, at two metres in height results in the breakage of the umbilical cord. Calves may stand within half an hour. Lightfoot (in Van Aarde 1976) observed that a Giraffe made no attempt to eat the placenta after giving birth, although Dagg & Foster (1982) and Kristal & Noonan (1979) noted otherwise. Kok (1982) described a free-living Southern Giraffe / scavenging on her dead recently born calf. The mother showed intense interest in the carcass over a period of weeks, no doubt indicative of the mother–calf bond. Neonates are hidden for a few weeks and thereafter form nursery groups only in populations where feeding and drinking resources are distant from place of birth (Langman 1977, Pratt & Anderson 1979). Mothers and young remain strongly associated for 14–22 months, until around the time of the next parturition (Pratt & Anderson 1982). Young calves are playful and peaceful, but frequent ‘necking’ occurs between subadult ??. Vocal folds and laryngeal ventricles are entirely absent, supporting the view that Giraffes are mute (Owen 1841, Hahn & Mayhew 1999). Nevertheless, they infrequently emit various loud sounds, notably under stressful conditions. They can grunt, snort, cough and whistle, and the calves bleat, though what such vocalizations communicate remains undetermined (Dagg & Foster 1982). Captive Giraffes emit infrasonic vocalizations around or below 20 Hz (von Muggenthaler et al. 1999). Bashaw (2003) undertook efforts to demonstrate the communicative function of these vocalizations in captive Rothschild’s Giraffes by submitting playbacks of infrasonic vocalization; however, this initial work showed no evidence of Giraffes modifying their behaviour.

Reproduction and Population Structure  Giraffes breed throughout the year, although birthing peaks have been observed in numerous populations, including: Nairobi N. P. in Kenya (Aug and Sep; Foster & Dagg 1972), Waza N. P. in Cameroon (Nov–Jan, when Acacia seyal is flowering; Ngog Nje 1983), Serengeti N. P. (Sep; Sinclair et al. 2000) and in N Namibia (Dec; Fennessy 2004), and at different times of the year in the same area (such as Kruger N. P.; see Dagg 1971). Birthing peaks often correlate with rainfall (Hall-Martin et al. 1975) and seem to coincide with the production of new Acacia shoots that have a high protein content (Sinclair et al. 2000, Fennessy 2004).Van der Jeugd & Prins (2000) suggest that differences in ecological circumstances create variability in mating strategies. Giraffes in captivity also breed aseasonally (Backhaus 1961, Bercovitch et al. 2004). A single calf is born, although twins have been recorded (Dagg & Foster 1982, R. Brenneman pers. comm.). Mean gestation period is approximately 457 days or 15 months (Backhaus 1961, Skinner & HallMartin 1975, Dagg & Foster 1982); one individual in captivity gave birth to consecutive infants only 420 days apart (Bercovitch et al. 2004). Neonates are heavier in the wild (101 kg) compared with those in captivity (55 kg, Skinner & Hall-Martin 1975). The mean height at birth is 1.8 m and 1.9 m for // and ??, respectively (Pellew 1983b). Calves nearly double their height within the first year (Pratt & Anderson 1982), which probably serves as an anti-predator strategy because calves are extremely vulnerable to predation (Estes 1991a; see later). Giraffe milk is rich in fat (13–17%), protein (6%) and ash content, although lactose concentration is lower than in bovine milk (Aschaffenburg et al. 1962, Hall-Martin et al. 1977a, Dagg & Foster 1982). In the wild, Pratt & Anderson (1979) found that male and female calves suckle at the same rate. Timing of weaning varies: exceptionally as long as two years, but typically around 9–12 months, although calves are able to eat solid food after the third or fourth week and begin ruminating at between three and four months (Langman 1977, Dagg & Foster 1982). Calving intervals are 19–20 months (Hall-Martin & Skinner 1978, Dagg & Foster 1982, Pellew 1983b, Ciofolo et al. 2000, Bercovitch & Berry 2009). Intervals are similar in captivity (n = 61), with a range of 14–38 months (Bercovitch et al. 2004). The interval between parturition and conception is usually 4–9 months. First conception has been recorded at 50 months in Serengeti N. P. (Pellew 1983b) and 56 months in southern Africa (Hall-Martin & Skinner 1978). In captivity, average age at first parturition was 57 months (n = 12; Bercovitch et al. 2004). Males are sexually mature at 3.5 years of age (and even earlier in captivity), but in the wild are probably excluded from reproduction by older bulls (Hall-Martin et al. 1978). Adult sex ratios have been recorded as being female-biased in Serengeti N. P. (Pellew 1983b), Nairobi N. P. (Dagg & Foster 1982), Katavi N. P. (Caro 1999a), Luangwa N. P. (Berry 1973) and in Niger (Ciofolo et al. 2000), while the opposite was true in Tsavo East N. P. (Leuthold & Leuthold 1978a). In N Namibia, sex ratios differed between subpopulations, although was unbiased across the overall population (Fennessy et al. 2003, Fennessy 2004). Sex ratio at birth is 1 : 1 (Dagg & Foster 1982, Bercovitch et al. 2004, Bercovitch & Berry 2009). Limited research restricts any real assessment of such variation. Between 54% and 62% of a given population is more than four years old. Mortality rates are strongly shaped by Lion Panthera leo predation, especially on neonates. In Serengeti N. P., 58% of young Giraffes died during their first year of life while only 2% died at the

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(Dagg 1971, Boomker et al. 1986, Krecek et al. 1990; and references therein). Bengis et al. (1998) recorded three species of Sarcocystis from muscle fibres of a Giraffe in South Africa (Sarcocystis giraffae, S. klaseriensis and S. camelopardalis). Larvae of the oestrid fly Rhinoestrus giraffae have been collected from Giraffes (Laurence 1961). Giraffes are vulnerable to anthrax, gastroenteritis and rinderpest, the latter killing 40% of the Giraffes in N Kenya in 1960 (Dagg 1971 and references therein). Other recorded parasite-induced diseases include listeriosis (Cranfield et al. 1985) and brucellosis (Jensen 1999). Polyarthritis and polyosteomyelitis were recorded in a juvenile (Jacobson et al. 1986), while a case of Mycoplasma-associated polyarthritis was recorded in a captive Reticulated Giraffe (Hammond et al. 2003). Various skin diseases and lesions have been observed.

Neck-sparring and fighting in male Giraffes.

age of four (Pellew 1983b); this is slightly lower than the figure of 75% given by Foster & Dagg (1972) for the same general area. In Kruger N. P., first-year mortality was also high at 48% (Hall-Martin 1975). A breeding / in Luangwa gave birth at 24 years of age (Bercovitch & Berry 2009), and these authors suggest that lifetime reproductive success (which ranged as high as 11 in Luangwa) is more dependent on longevity and calf survivorship than on reproductive rate. The maximum lifespan of male Giraffes, based on individuals in Luangwa Valley, is shorter (22 years) compared with // (28 years) (Berry & Bercovitch 2012); maximum longevity in captivity is recorded at 39 years (Weigl 2005). Given the uniform sex ratio at birth, comparable mortality rates, suckling rates and inter-birth intervals among juveniles, and the similar growth rate of calves while nursing, Bercovitch et al. (2004) suggest that adult // invest equally in their offspring, regardless of sex, and that ?? surpass // in size only after the period of dependency. Predators, Parasites and Diseases  Lions are the main predator (Pienaar 1969a, Pellew 1983b, Hayward & Kerley 2005), although Giraffes are formidable opponents with their long strong legs and hooves; reports of Lions killed or injured by Giraffes are not uncommon. Spotted Hyaenas Crocuta crocuta, African Wild Dogs Lycaon pictus and Leopards Panthera pardus (Scheepers & Gilchrist 1991) have been reported preying on calves occasionally, while observations of Nile Crocodile Crocodylus niloticus and Cheetah Acinonyx jubatus kills have also been recorded. Ectoparasites are particularly numerous around the anus and the genitalia. Common tick species recorded from animals in southern Africa include Amblyomma hebraeum, Boophilus decoloratus, Hyalomma aegyptium, H. marginatum, H. truncatum, Rhipicephalus appendiculatus, R. camelopardalis, R. evertsi and R. longiceps (Horak et al. 1983c, 1992a, 2007). Helminth parasites include: Cooperia pectinata, C. punctata, Fasciola gigantica, Haemonchus contortus, H. mitchelli, Moniezia expansa, M. nullicollis, Monodontella giraffae, Parabronema skrjabini, Pseudofilaria giraffae, Setaria labiatopapillosa, Skrjabinema sp., Trichocephalus gracilis, Trichuris globulosa, T. giraffae, Trichostrongylus sp. and Echinococcus sp.

Conservation  IUCN Category: Least Concern (G. c. peralta – Endangered D; G. c. rothschildi – Endangered C2a(i)). CITES: Not Listed. East (1999) estimated the total number of Giraffes in Africa to exceed 140,000 (of which 40% were in or around protected areas and private lands); such numbers were thought capable of being maintained were they adequately protected (East 1999). Current estimates put the population at less than 100,000, evidencing declines in some populations. For example, poaching and armed conflict across the range of the Reticulated Giraffe in Somalia, Ethiopia and Kenya has reduced numbers to perhaps fewer than 3000 individuals (Georgiadis, in Fennessy 2007). Smaller, managed populations have also declined: a decline in Lake Nakuru N. P. in Kenya has been attributed to dietary complications from highly concentrated tannin levels because of forced overconsumption of the park’s declining acacia trees, which may have compromised young Nubian/Rothschild’s Giraffe, making them easy and opportunistic prey for the park’s Lion population (Brenneman et al. 2009). Important safety havens include Waza N. P. and the hunting zones of North Province in Cameroon, Zakouma N. P. (Chad), Murchison Falls N. P. (Uganda), Boma N. P. (Sudan), Omo N. P. (Ethiopia), South Luangwa N. P. (Zambia) and, in southern Africa, Etosha N. P. (Namibia), Hwange N. P. (Zimbabwe) and Kruger N. P. (South Africa). Masai Giraffes occur widely both within and outside protected areas in Kenya and Tanzania, but represent a vestige of their former numbers and range. Small numbers of Nubian Giraffes have been introduced to smaller parks and game reserves in Kenya, including Lake Nakuru and Ruma National Parks, while both Angolan and Southern Giraffes have been widely reintroduced and introduced in southern Africa. Giraffes were also introduced to Akagera N. P. in Rwanda in 1986 (East 1999). In Niger, conservation development projects have facilitated the Niger Giraffe’s population increase in an area outside any formal protected park or reserve. None the less, poaching and habitat loss, fragmentation and degradation as a result of increased aridity, and expansion of human activities continue to impact on the Giraffe’s distribution. This small population survives only in the wild, since the Giraffes held in captivity in the Vincennes Zoo, France, which were long referred to as peralta, in fact belong to the subspecies antiquorum (Hassanin et al. 2007). The recently uncovered genetically distinct populations clearly represent evolutionarily significant units that are highly threatened and lack appropriate recognition in current management plans (Brown et al. 2007). 109

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Measurements Giraffa camelopardalis TL (??): 5.05 (4.86–5.27) m, n = 15 TL (//): 4.44 (4.16–4.75) m, n = 16 Sh. ht (??): 3.31 (3.13–3.47) m, n = 15 Sh. ht (//): 2.80 (2.72–2.92) m, n = 16 WT (??): 1191.8 (973–1395) kg, n = 18 WT (//): 828.4 (703–950) kg, n = 18 South Africa (from individuals older than eight; Hall-Martin 1975)

Mitchell et al. (2009a) reported an upper weight for // of 1049.2 kg and 1511.6 kg for ?? Key References  Ciofolo & Pendu 1998; Dagg 1971; Dagg & Foster 1982; Fennessy 2004; Hall-Martin 1974, 1975; Hall-Martin et al. 1975, 1978; Leuthold & Leuthold 1972, 1978a, b; Mitchell & Skinner 1993, 2003, 2009; Pellew 1983a, b, 1984a, b; Pratt & Anderson 1979, 1982. Isabelle Ciofolo & Yvonnick Le Pendu

Subfamily OKAPINAE – Okapi Okapinae Bohlin, 1926. Palaeontologia Sinica, ser. C, 4: 1–179.

Subfamily Okapinae is a monotypic subfamily, represented by a single species, the Okapi Okapia johnstoni.

Genus Okapia Okapi Okapia Lankester, 1901. Nature 64: 24.

The genus Okapia includes only the Okapi Okapia johnstoni, which is confined to lowland rainforest in DR Congo. The only confirmed fossil record for the species is an early Pliocene ossicone (Cooke & Coryndon 1970). The Okapi exhibits a number of primitive and conservative morphological and behavioural traits. It has been referred to as a living fossil by some authors (Joleaud 1937, Colbert 1938) and some have aligned it with the extinct Palaeotraginae (McKenna & Bell 1997).

Okapia differs from Giraffa in being much smaller, as little as onethird the mass, with proportions more like those of other ruminants of similar size; its cervical vertebrae are not elongated, there are five sacral vertebrae (cf. usually four in Giraffa) and neck and limbs are only moderate in length. Its ‘horns’ (ossicones) are hair-covered, situated above the eyes, in ?? only and there is no median ossicone. The skull is elongated, the long braincase in line with the facial part of skull and not flexed. Peter Grubb

Okapia johnstoni Okapi Fr. Okapi; Ger. Okapi Okapia johnstoni (Sclater, P. L., 1901). Proc. Zool. Soc. Lond. 1901 (1): 50. DR Congo, ‘in sylvis fluvio Semliki adjacentibus’ (= Semliki Forest, Mundala).

Taxonomy  Monotypic. The discovery and naming of this species is one of the great epics of natural history as well as a vivid illustration of the power of preconceived ideas.While the explorer H. M. Stanley was in the Ituri Forest in 1876 he saw pieces of striped skin, which his interpreters told him came from a type of ‘forest donkey’. This unknown animal caught the interest of his friend and admirer, the naturalist, Sir Harry Johnston, who, around the turn of the century, was Governor of Uganda. When the opportunity arose, Johnston visited Ituri and, for a time, thought he was on the trail of the extinct three-toed Hipparion. Disbelief in the Okapi’s two-toed tracks, together with an attack of malaria made Johnston abandon his quest, but he carried away two hide bandoliers cut from the haunches of Okapis. These were sent to the anti-Darwinian, P. L. Sclater, who provisionally assigned them to the genus Equus, naming the new species Equus johnstoni. In early 1901 Johnston was sent an entire skin and two skulls

of Okapis by a Swedish agent of the Congo Free State, Lieutenant Karl Eriksson, and immediately realized that the animal was an ally of the Giraffe Giraffa camelopardalis. He sent the skin, skulls and his own brilliant pictorial reconstructions of the living animals to Sir Edwin Ray Lankester, Director of the British Museum of Natural History. Latinizing the Mbuti pygmy name into Okapia, he described Okapia johnstoni in the journal Nature in the same year that Sclater’s Equus johnstoni came out in the slow-to-be-published Proceedings of the Zoological Society of London. Synonyms: erikssoni, kibalensis, liebrechtsi, tigrinum. In contrast to Giraffes, which have only 30 chromosomes, the chromosome number of the Okapi recorded from animals in captivity is either 2n = 46, 2n = 45 or 2n = 44 (Ulbrich & Schmitt 1969, Hösli & Lang 1970, Koulisher 1978, Petit et al. 1994,Vermeesch et al. 1996, Petric 2004a); the 2n  = 45 karyotype has been recorded from at least one wild-caught

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Okapi Okapia johnstoni.

? (Petit & de Meurichy 1986), and has occurred in 53 of 116 captive animals examined (K. Leus pers. comm.). This aneuploidy was likely established through a Robertson fusion.These animals appear normal and produced viable offspring (Lindsey et al. 1999). Description  A small forest giraffid that may, in some postures, appear strikingly deer-like, with a narrow, long-muzzled head, short, muscular, tapered neck and posteriorly sloping back. The Okapi’s bilobed lower canines and hair-covered horn formations originating from epidermal cartilaginous ossicones are features it shares uniquely with all other living and fossil giraffids. The Okapi shares with the Giraffe a distinctive lateral pacing gait in which the weight is momentarily shared by legs on the same side of the body as the animal progresses. The Okapi also differs in having cervical vertebrae that are not elongated; there are usually five sacral vertebrae in the Okapi (cf. four in the Giraffe), and only ??, generally, have ossicones. The Okapi is a distinctively coloured and patterned animal. Face pale grey, variably grizzled with dark hairs. Crown chestnut brown with furcovered, posteriorly oriented, bone-tipped ‘horns’ (see below), although the hair on the tips is sometimes rubbed to the bone. Eyes dark brown, set off by long lashes. Ears large, chestnut brown, with a black fringe of hairs. Neck and body overall dark chestnut brown with maroon overtones. Chest and upper abdomen chestnut brown, lightening and becoming nearly white in the perineal region. Hindquarters dark brown to black with 18–25 or more irregular white horizontal stripes, extending over the buttock and gaskin (tibia) in a pattern that is unique to each animal. Rear cannon or metatarsus white with a black band on the fetlock and white pastern. Similar, though reduced pattern, occurs on the upper forelegs. Fore cannon, or metacarpus, white, with a brown to blackish line running up the front, black hair on the knee and fetlock, white pastern.Tail chestnut brown with black tuft, reaching to the level of the hocks. Young Okapis have patterning similar to adults, but with overall more shaggy pelage, and a distinct blackish mane from the base of the head along the upper back, which gradually diminishes as the animal grows older and all but disappears by 12–14 months. Hooves

rounded and relatively small for the size of the animal. The pelage is short and lightly oiled from a reddish-brown secretion from the skin that accumulates abundantly in the ears, and renders the pelage somewhat water-resistant. Unlike Giraffes, both sexes have large pedal glands above all four hooves (larger in the fore hooves; Pocock 1936). These drain around the top of the hoof in a channel that leads down between the two toes. Adult // are distinctly larger than ??, averaging about 7 cm higher at the withers, and on average 8% larger body mass. Females have four inguinal nipples. Bodmer & Rabb (1992) and Lindsey et al. (1999) have provided a detailed description of the Okapi, including variability in coat patterns found in a sample of animals from the central Ituri that were brought into captivity at Epulu. The skull is elongated, the long braincase in line with the facial part of the skull, and not flexed. There is a large parietal region, and short diastema. The lacrimal fossae are large. The large auditory bullae, and the large frontal and palatine sinuses are unique to the Okapi and separates it from the Giraffe (Colbert 1938, Bodmer & Rabb 1992). The horns measure 6–15 cm in length in ??, but are absent (or rarely present in very reduced form) in //. These horns ossify and fuse with the frontal bones supraorbitally as the animal matures (Churcher 1990); horn development in ?? begins around one year of age (Bodmer & Rabb 1992).There is no median ossicone. The cheekteeth are low crowned; the deciduous and permanent canines are incisor-like, while the incisors form a semi-circle at the end of the jaw. Skull development and tooth eruption are discussed by Jaspers & de Vree (1978). Geographic Variation  There are no currently recognized subspecies. Okapi populations occurring west and south of the Congo R. are reported to be overall darker than those to the east and north (I. Liengola pers. comm.). While the Congo R. must be a major biogeographical barrier for a species not known to swim, the genetic relationships between populations on either side of this river await investigation.

Lateral, palatal and dorsal views of skull of Okapi Okapia johnstoni.

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Okapi Okapia johnstoni skeleton.

Similar Species  In local parlance, might be mistaken for Bongo Tragelaphus eurycerus (due to striping), and sometimes even referred to as a zebra (this harkening back to the original misconception of the identity of the species). In the forest, the dung and tracks are distinct from other ungulates, with Okapi tracks being more rounded than Bongo, and smaller than African Buffalo Syncerus caffer. Distribution  Endemic to Africa. The Okapi is confined to the forests of DR Congo, occurring between about 500 m and 1500 m elevation over a fairly large range, on both sides of the Congo R. The Okapi ranged occasionally into the adjoining Semliki forest of extreme W Uganda in the recent past (Kingdon 1979). In the forests east and north of the Congo R., the Okapi is known from about latitude 1° S, in a broad arc north through the Maiko and Ituri Forests, then west through the Rubi,Tele and Ebola river basins (irregularly, to the Ubangui R.). The Okapi is also present in central DR Congo, west and south of the Congo R. in the forests of the upper Tshuapa and Sankuru Districts, from the west bank of the Lomami between 1 and 2° S latitude, west to the upper Lomela and Tshuapa basins (confirmed by I. Liengola pers. comm.).The forest–savanna transition defines the eastern and northern limits of the Okapi’s range. In contrast, the southern limits east of the Congo R., and the western and southern limits of the species in C DR Congo, are poorly defined. In the west, the species does not appear to occur very far to the west of 24° E.The southern and western limits are not associated with any apparent biogeographical barriers, such as changes in vegetation type and elevation or occurrence of major river barriers (neither do overall forest composition and forest structure change over most of the apparent range boundaries). The Okapi appears to be absent from large areas of closed-canopy forest in E and C DR Congo. Incidental reports of Okapi occurrence (mapped in Kingdon 1979) and rumours of Okapi presence persist in areas well beyond their currently known range. Okapis are secretive and shy, and their occurrence, especially at low densities, can easily go undetected. The primary strongholds of the Okapi today are the Ituri/ Aruwimi and adjacent Nepoko basin forests, and the forests of the upper Lindi, Maiko and Tshopo basins.The species is also well known in the Rubi-Tele region in Bas Uele. Up until the last two decades of the twentieth century, the Okapi’s occurrence remained essentially the same as when it was discovered in the first decades of the century. However, since 1980, major expansions of human settlement

Okapia johnstoni

accompanied by deforestation and forest degradation, have eliminated important portions of their range, in particular in the southern and eastern Ituri Forest, where the species was at one time abundant.The type locality of the Okapi at Mundala has been overtaken by expanding settlement from the growing urban centre of Beni. Habitat  The Okapi is limited to closed-, high-canopy forests, occurring in a wide range of primary and older secondary forest types. It does not range out into gallery forests or into the forest islands on the savanna ecotone and it does not persist in the disturbed habitats surrounding larger forest settlements. Okapis will also forage in regenerating clearings abandoned by shifting cultivators when these are small (generally 6 years) ?? to receptive (>2 years) // is 1 : 12. Since there are barely enough bulls to go around among female groups (effectively one mature bull for every 2–3 female units), ?? rarely need to contest dominance through overt aggression or territorial exclusion. Five-year-old bulls even get mating opportunities when no older ? is present. This system was called ‘mating enhancement by attrition of rivals’ (Owen-Smith 1993b). The prime cause of the differential male mortality is not clear. It seems basically a consequence of larger-than-optimal body size, with larger size benefiting reproduction at the cost of survival. Costs probably arise through malnutrition as well as lessened agility in evading predators. Unlike many other ungulate species, Greater Kudu ?? show relatively little reduction in foraging time during the breeding period (Owen-Smith 1984). Greater Kudus are susceptible to mortality when cold wet spells occur at the end of the winter dry season in southern Africa (Wilson 1970, Simpson 1972a). In Kruger N. P., a maximum daily temperature of 14 °C associated with overcast conditions, rain and wind resulted in a doubling of adult mortality from 10% to 20%, especially among old //, and reduction in calf survival from 60% to 25%, despite prior good rainfall (Owen-Smith 2000). Predators, Parasites and Diseases  Greater Kudus, or at least // and young, are subject to predation by all the larger carnivores, including Lions Panthera leo, Spotted Hyaenas Crocuta crocuta, Leopards Panthera pardus and African Wild Dogs Lycaon pictus. They usually are not the major prey species for any of these predators, except in areas where abundant (e.g. see Pole et al. 2004). Even though predation is the direct cause of death of almost all animals, except during severe droughts, it is generally interactive with nutrition because of the strong correlation between the mortality rates of all except the prime age class and rainfall relative to population density (OwenSmith 1990). Greater Kudus are among the main hosts of tsetse flies, although they do not seem to suffer ill effects from the trypanosome microparasites transmitted by these flies. They seem especially susceptible to several wildlife diseases causing severe mortality during outbreaks. Greater Kudu were among the ungulate species decimated during the rinderpest panzootic that spread through Africa during the 1890s (Mack 1970, Rossiter 1994). Animals in E Kenya were adversely affected by a rinderpest epidemic in 1993–97 (Kock et al. 1999). The species seems to be centrally involved in spreading anthrax during outbreaks of this disease in Kruger N. P., as well as suffering severe mortality, resulting in a population decline by as much as 40% in infected areas (De Vos & Bryden 1996, Bengis et al. 2003). Greater Kudus pick up spores through feeding on leaves where blowflies have defecated after feeding on infected carcasses. However, in Etosha N. P. in Namibia,

where anthrax is endemic, Greater Kudus make up a very small proportion of the animal deaths (Lindeque & Turnbull 1994). Rabies caused severe mortality among Greater Kudus in Namibia in 1981, amounting to 16–20% of the estimated population per farm, and up to 50% losses in some areas (Hassel 1982). Transmission seems to occur via salivary contamination of browse, and the outbreak was associated with an abnormally high density of Greater Kudus at that time. Greater Kudus also suffer from bovine tuberculosis, and may play a role in maintaining the disease in the system (Bengis et al. 2003). They also appear to be very susceptible to Bovine Spongiform Encephalitis (Cunningham et al. 2004). Greater Kudus carry high tick loads, especially of Boophilus decoloratus, Amblyomma hebraeum and Rhipicephalus spp. (Horak et al. 1992b, Gallivan & Surgeoner 1995, Zieger et al. 1998b). The Redbilled Oxpeckers Buphagus erythorynchus that commonly clamber over them probably play an important role in controlling the tick burdens, along with the reciprocal grooming that Greater Kudus show. They are also subject to bites from swarms of tabanid flies as well as tsetse flies (Boomker et al. 1989b, N. Owen-Smith pers. obs.), probably on account of their comparatively sparse hair coat. The nematode roundworm burdens carried by Greater Kudus are somewhat lower than those found in grazing ungulates. Species commonly found include Haemonchus vegliai and Coopera neitzi, Coopera acutispiculatum, Cordophilus saggitus and Trichostrongylus deflexus (Boomker et al. 1988, 1989b). Lactating // may show substantially elevated worm loads, but no increase in nematode numbers was evident in a drought year. Conservation  IUCN Category: Least Concern. CITES: Not listed. Greater Kudus are much sought after by hunters, both for the magnificent horns of bulls and more generally for their high-quality meat. Populations seem to be quite resilient to hunting pressure, aided by their alert and reclusive habits and their apparent ability to persist in settled areas that have suitable cover.They are also a favoured gameranching species, because as browsers they do not compete with domestic livestock. In southern Africa, Greater Kudus are common on private farms and conservancies; East (1999) estimated that some 60% of the global population occurs on private land, and they seem to be expanding their distribution outside protected areas. In the southern and south-central parts of their range, Greater Kudus are generally well represented in protected areas, with important populations in Ruaha N. P. and Rungwa and Selous Game Reserves (Tanzania), Luangwa Valley and Kafue National Parks (Zambia), Etosha N. P. (Namibia), Chobe G. R. (Botswana), Hwange and Mana Pools National Parks (Zimbabwe), Niassa G. R. (Mozambique) and Kruger N. P. (South Africa). However, in the northern parts of the species’ range, the Greater Kudu seems to be in decline, with its status threatened further by the species’ fragmented distribution and susceptibility to diseases transmitted by cattle. Key populations in this part of the range include those in Zakouma N. P. (Chad), Nechisar, Omo, Mago and Awash National Parks (Ethiopia), Laikipia and Tsavo N. P. (Kenya) and Tarangire N. P. (N Tanzania) (East 1999). Measurements Tragelaphus strepsiceros HB (??): 2170 (2060–2490) mm, n = 33

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HB (//): 1970 (1520–2160) mm, n = 68 T (??): 450 (400–520) mm, n = 33 T (//): 410 (350–470) mm, n = 68 E (??): 260 (245–290) mm, n = 33 E (//): 240 (210–255) mm, n = 68 Sh. ht (??): 1340 (1220–1430) mm, n = 17 Sh. ht (//): 1240 (990–1390) mm, n = 48 WT (??): 249.0 (174.0–344.0) kg, n = 94 WT (//): 160.0 (112.0–210.0) kg, n = 97 Southern Africa (combined from Wilson 1965, 1970, Skinner &

Chimimba 2005, V. De Vos pers. comm. and J. D. Skinner pers. comm.) Record horn length is 187.6 cm for a pair of horns picked up near the Save R., Mozambique (Rowland Ward), the longest of any antelope recorded. Key References  Owen-Smith 1979, 1990, 1993a, b, c, 1994; Owen-Smith & Cooper 1989; Simpson 1968, 1972a. Norman Owen-Smith

Tragelaphus buxtoni  Mountain Nyala (Gedemsa) Fr. Nyale des montagnes; Ger. Bergnyala Tragelaphus buxtoni (Lydekker, 1910). Nature 84: 397. Ethiopia, Bak Prov., ‘Arusi plateau of Gallaland, in the Sahatu Mountains, and south-east of Lake Zwei [Zwai], at an estimated height of 9000 feet [2700 m] above sea level’.

flank and spiral horns with a strong posterior keel, but of stockier build, thicker neck and heavier horns. Adult ?? have a dun brown coat, which gets progressively darker with age, with white markings on face, throat, legs and poorly defined stripes on back and upper flanks. Coat texture varies, possibly by season, from sleek to shaggy; hair in back of thighs longer and stiffer. An erectile white crest or mane, 100 mm high, runs from neck to tail; the bushy tail reaches the heel. Females smaller and lighter in colour, resembling a Red Deer hind in size and proportions; dark greyish-brown coat, with paler undersides, white markings on legs and face and faint spots and stripes; erectile crest absent. Young cows bright rufous, with older cows as grey as a young bull. Ears large, lined with tracts of white hairs and a narrow black smear-like mark on the lower margin. Only ?? have horns, which are lyre shaped, and grow outwardly in a spiral with 1–2 turns, a distinctive back ridge, and sometimes having ivory tips. Females have two pairs of inguinal nipples. Geographic Variation  None recorded.

Mountain Nyala Tragelaphus buxtoni male.

Taxonomy  Monotypic. Described from a specimen brought to England in 1908 by Major Ivor Buxton, the Mountain Nyala is the last major large mammal to be discovered in Africa. Despite its common name, studies involving nuclear DNA reveal that the species forms a monophyletic clade with the Bongo T. eurycerus, Sitatunga T. spekii and Bushbuck T. scriptus, all species adapted to closed forest (WillowsMunro et al. 2005). Synonyms: none. Chromosome number: not known. Description  A large tragelaphine similar in appearance and size to the Greater Kudu T. strepsiceros, sharing a row of spots along the

Similar Species Tragelaphus strepsiceros. Drier wooded areas of Ethiopia, potentially sympatric on some mountain slopes. Taller, colour paler and more graceful; horns longer, with 2–3 spirals and tips more separated. T. angasii. Southern Africa only, in Malawi, Mozambique, Zimbabwe and South Africa. Smaller and readily distinguishable by an abundant fringe of long hair on throat and neck; horns similar, although slender and narrower. Distribution  Endemic to Ethiopia. Restricted to a few scattered populations alongside an arc of mountains south-east of the Rift Valley, between 6° N and 10° N. Formerly occurred from Gara Muleta in the east to Shashamane and north Sidamo in the south, but eliminated from a large proportion of its former range. Presently, up to half of the world population is found in the Bale Mountains and the eastern escarpments of the Bale massif (Shedem, Odo Bulu, Abasheba). Smaller relict populations occur in Chercher (= Amhar) Mts (Kuni-Muktar, Arba Guggu, Din Din), Arsi Mts (Chilalo, Galama, Mt Kaka, Munessa) and West Bale (Somkaro-Korduro ridge) (Hillman 1988a, East 1999, Malcolm & Evangelista 2005). 159

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Tragelaphus buxtoni

Habitat  Ideal habitat is provided by montane woodlands (3000– 3400 m), dominated by Juniper, Podocarpus and Olea in the lower parts and Hagenia, Juniper and Hypericum in the upper reaches. Mountain Nyalas frequent the fringes of montane grasslands (2800–3100 m) dominated by sage brush Artemesia afra, red-hot pokers Kniphofia foliosa and everlasting Hypericum spp. Highest densities (up to 21 ind/ km2) have been recorded in the montane grasslands of Gaysay, Bale, where there is a combination of browse and grass with woodland cover to retreat into during the day (Hillman & Hillman 1987). What were formerly large continuous blocks of suitable woodland and afroalpine habitat have now been reduced to a series of habitat islands in a sea of cultivated fields. It appears likely that Mountain Nyalas have been forced into even higher areas due to human increase and livestock grazing, with animals also found above 3400 m on heath forest and ericaceous heath lands (Erica and Phillippia spp. with Hypericum, Euphorbia and Helichrysum shrubs) and on afroalpine grasslands (Alchemilla spp., Festuca spp.) up to 4300 m. In the eastern extreme of its distribution, a relict population was recorded in forests as low as 1800 m (Bolton 1973); the Munessa population occurs at 2400 m (Malcolm & Evangelista 2005). Abundance  The world population of Mountain Nyalas was estimated at 7000–8000 (and perhaps even as high as 12,500) in the 1960s by Leslie Brown (Brown 1969a), and at between 2000 and 4000 individuals in the 1980s (Hillman 1988a). Global numbers have declined since. Habitat suitability projections indicate that there may be as little as 8333 km2 (Atickem et al. 2011) or up to 40,000 km2 (Evangelista et al. 2008) of forest habitat available to the species, but the latter estimate includes many areas heavily impacted by human activities. Mountain Nyalas are possibly extinct in the eastern and southern extremes of their distribution, but small numbers may still occur in Asba Tafari and in the border between Bale and Sidamo south of Kofele. The total population was calculated by East (1999) at 2650, but subsequent

Lateral and palatal views of skull of Mountain Nyala Tragelaphus buxtoni.

information indicates that this may be an overestimate. The main extant population occurs in and around the Gaysay grasslands in the northern extreme of the Bale Mts, where numbers have been monitored since 1983. In response to the creation of a national park in the 1970s, which provided protection from poaching and the exclusion of cattle grazing, the Mountain Nyala population in Gaysay increased to 1050 by the late 1980s (Hillman 1988a). Woldegebriel (1996) put the population prior to 1990 between 1500 and 1900. Unfortunately, as a result of the political unrest that followed the end of the war in 1991, most Mountain Nyala habitat in northern Bale was encroached by cattle and there was extensive shooting of nyalas. Consequently, the Gaysay population decreased to only a fraction of what it was, and park staff estimated it to be 150–260 by 1994 (Woldegebriel 1996). There have been signs of recovery since, and the population was tentatively estimated in 1997 at 530–1000 using transect counts (Stephens et al. 2001), although this result may be biased due to transect design, and a conservative population estimate should be put close to the lower figure. More recently, the Gaysay population was estimated at 550 by Refera & Bekele (2004) and Malcolm & Evangelista (2005). In addition to the population in Gaysay there may be as many as 80–120 Mountain Nyalas elsewhere in Bale Mountains N. P., less than 100 in adjacent hunting areas to the north of the park and 30–60 in Somkaro in W Bale (C. Sillero-Zubiri pers. obs.). Malcolm & Evangelista (2005) estimated as many as 500 Mountain Nyalas may occur in hunting blocks east of Bale. This would give a total population estimate for the Bale massif of 1000–1400. Small fragmented populations found in Arsi (Galama, Chilalo, Kaka and Munesa) and elsewhere (Kuni Muktar, Din Din, Arba Gugu) would total some 600 animals (Malcolm & Evangelista 2005). Although Atickem et al. (2011) estimated a population of over 3000 using faecal pellet counts, it is likely that only 1500–2000 Mountain Nyalas survive throughout the range. There are none currently kept in captivity (East 1999).

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move up into wooded areas and into the thick ericaceous heath. They move mostly at night to avoid human disturbance by day.

Mountain Nyala Tragelaphus buxtoni female.

Adaptations  Mountain Nyalas are shy, rather elusive antelope, and have adapted to life in the relatively rich highland habitats, resulting in a specialized diet and physiology; they have a limited diet of montane vegetation, and have adapted to extreme temperature variation. As a result, it is likely that they are outcompeted by Greater Kudus in the surrounding arid lowlands. The animal’s skeletal proportions, with relatively short distal limb segments, are characteristic of a mountain quadruped, which probably evolved in the relatively richer and varied periglacial habitats. In a richer environment there is a premium on early reproduction and more open conflict over //. This ecological role is correlated with a relatively neotenic horn morphology. Males have lyrate horns that provide an effective jabbing weapon for male–male contests, but are ineffective as defensive shields. Mountain Nyalas living in continuous forests at lower elevations have an extra second spiral, suggesting that populations living in more stable lower productivity habitat may be more similar to the Greater Kudu in life history parameters (J. Malcolm pers. comm.). Foraging and Food  Mountain Nyalas are mainly browsers, and were classified as such by Gagnon & Chew (2000) in their review of the dietary preferences of African bovids. They are selective feeders, feeding on low-growing herbs and sprigs of shrubs and bushes, and tree foliage, with more grasses eaten during the early wet season. Favoured species include Artemesia afra, Hypericum revolutum, Kniphofia foliosa, Solanum sessilistellatum and Hagenia abyssinica leaves, which they either pick from the ground or use their horns to bring branches within their reach. On the afroalpine grassland, Mountain Nyalas frequently eat Alchemilla rotti, Helichrysum splendidum and the lower leaves of Lobelia rynchopetalum. They sometimes feed on the site of a fresh burn or on the lush green of abandoned homesteads, and have been observed eating lichens, ferns and aquatic plants (Hillman 1986a). Mountain Nyalas spend the night in the forest fringes where they probably feed part of the time, coming out early in the morning and late afternoon to feed in the grassland fringes, using the woodland and heather thickets during the hotter and colder times of the day. However, some animals often feed briefly in the middle of the day, laying down between feeds. This pattern is probably determined by the weather, with animals preferring to stay in cover during frosts and midday heat and coming out when it is overcast or raining. In the dry season (Nov– Mar), when the grassland is in poor condition, many Mountain Nyalas

Social and Reproductive Behaviour  Mountain Nyalas have a highly cohesive basic social group consisting of an adult /, accompanied by her calf of the previous year, and a calf of the current year. Family units aggregate with others for short periods of time to form small herds of 4–5 (mean group size = 9) (Hillman 1986a). Apparently, herds have always been small, even when the species was more common, with herds of 4–12 reported by Maydon (1925). Larger more fluid groups of up to 100 animals are often seen in Bale, with family units moving in and out (Hillman 1988a). Size and longevity of these larger groups varies with season, habitat type and time of day. Female–young groups are often monitored by adult ??, depending on the presence of oestrous //.Young ?? leave the family herds at around two years of age and join other adult and young bulls to form bachelor groups of up to 13 animals (Hillman & Hillman 1987).

Circling dominance display by male Mountain Nyala Tragelaphus buxtoni.

Males are not territorial, and have home-ranges of 15–20 km2 in the wet season, with // and juveniles using smaller wet season homeranges of 5 km2 (Hillman 1986a). Dry season ranges are significantly larger for both sexes. A dominance hierarchy is apparent in male herds, maintained by pushing and tussling of horns in the young and ritualized displays in the older ??. Old bulls live alone, occasionally approaching a herd to check for receptive //. More ?? are associated in mixedsex groups during the period when most mating, pre-mating and follow-up behaviour takes place. Although intense following is occasionally observed throughout the year bulls are more likely to be seen alone during the dry season. Bulls follow receptive // persistently and test their vulvas, followed by flehmen behaviour. Often, 3–4 bulls follow in ritualized parallel walks. When two evenly matched bulls meet, they engage in a wary circling dance. They move very slowly, with strutting gait and head lowered, their spinal crests and fluffed tail raised. Sometimes the strut gives way to a stiff sideways shuffle, until eventually smaller ?? give way and drift away. Less often the ?? take the challenge with a brief heated clash of horns. Rutting ?? engaged in these ritualized walks often carry ‘decorations’ of soil, branches and grass tussocks hanging off their horns. 161

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Calves lie up for several weeks in cover, later rejoining their mother. Mountain Nyalas are mainly silent, but // may utter a low bark when alarmed or ‘cough’ when the threat is less serious (Brown 1969a). Reproduction and Population Structure  Females first mate at two years of age, giving birth to a single calf about 8–9 months later. In Bale, calving takes place throughout the year, with a peak of births (70%) from Sep to Nov, towards the end of the wet season (Hillman 1988a). There is a high calf to adult / ratio, suggesting a calving rate of at least 50%, and a high survival rate of calves (Hillman 1986a). Predators, Parasites and Diseases  Possible natural predators include Leopards Panthera pardus, Spotted Hyaenas Crocuta crocuta, Servals Leptailurus serval, Ethiopian Wolves Canis simensis and domestic dogs. The latter two species have been known to kill Mountain Nyala calves in and around the Gaysay grasslands in Bale (Sillero-Zubiri & Gottelli 1995, Woldegebriel 1996), but there are no records available for the other species. None is thought to have a serious effect on adult Mountain Nyalas (Brown 1969a). Lions Panthera leo could take adults, but they are not common in Mountain Nyala range. Calves and juveniles may be taken frequently by Leopards and Spotted Hyaenas at night but the effect on survival, if any, is unknown. Almost nothing is known regarding diseases in Mountain Nyalas, although they are susceptible to rinderpest (Fekadu Shiferaw pers. comm.). Females are often found dead during the wet season in Bale, showing symptoms compatible with those of poisoning from eating certain vegetation. Graber et al. (1980) recorded a fluke (Cotylophoron cotylophoron) and two nematodes (Oesophagostomum walkeri and Haemonchus vegliai). Conservation  IUCN Category: Endangered C1. CITES: Not listed. Threatened directly by illegal hunting and indirectly by disturbance and destruction of montane forest and heath lands, encroachment by cattle, harassment and hunting by dogs, highaltitude cultivation, and roads. In the past, Mountain Nyalas were often seen living alongside pastoralists when the latter were present at low densities (Maydon 1925). However, permanent occupation of suitable habitat as a result of increasing human pressure and livestock populations is exerting tremendous pressure on Mountain Nyala habitat throughout the range, with anecdotal evidence suggesting the animals actively avoid livestock (C. SilleroZubiri pers. obs.). This pressure is also apparent within Bale Mountains N. P., home to 60–80% of the world’s Mountain Nyala population, and which was created primarily to protect the Mountain Nyala (Brown 1969a). The impact of effective conservation measures in northern Bale during the 1970s and 1980s was phenomenal, and the Mountain Nyala population flourished as a result of the exclusion of domestic stock (Hillman 1986a). Unfortunately, a decline in (and at times collapse of) park management during the 1990s resulted in human encroachment, overgrazing, illegal hunting and Mountain Nyalas

moving out of optimal habitat. Additionally, the constant passage of people through their range disturbs their foraging habits. Although Mountain Nyalas are fully protected by law there has been a proliferation of hunting licences issued out to hunting concessions around Bale. The absence of enforcement and the general ignorance of protective legislation among the local people, and the failure of police and other officials to make any serious attempt to enforce the law, restrict effective protection to a small area of habitat (20 km²) in Gaysay and around Bale Mountains N. P. headquarters. Elsewhere, Mountain Nyalas are extensively hunted for meat and horns, the latter used for local medicine and to make teats for traditional milk bottles. Conservation action required in Bale Mountains N. P. and adjacent hunting areas includes the enforcement of patrolling to curb cattle grazing and the protection of the Mountain Nyala population from illegal hunting. The park should ideally be extended northward to include suitable habitat within hunting blocks. There is also a need to extend patrolling to the higher-altitude areas of the park, where uncontrolled livestock grazing needs to be checked. Elsewhere illegal hunting needs to be controlled throughout the Arsi Mts, where the recent establishment of an additional protected area may deliver some conservation benefit for the species. Although it has been argued that high licence fees may finance conservation measures and derive a benefit to local people the current quota of adult bulls made available for trophy hunting may be unsustainable in the long term. Hunting blocks in Arsi have been hunted out and concessions moved to Bale, with continued pressure by the industry for additional hunting blocks and larger quotas. While legal hunting is restricted to adult ??, trophy size needs to be monitored carefully. Absence of old bulls or a sex ratio imbalance may affect reproduction and population dynamics. Measurements Tragelaphus buxtoni HB (??): 2400–2600 mm HB (//): 1900–2000 mm T: 200–250 mm HF c.u.: 550–600 mm E: 230–250 mm Sh. ht (??): 1200–1350 mm Sh. ht (//): 900–1000 mm WT (??): 180–300 kg WT (//): 150–200 kg Chercher and Bale Mts, Ethiopia (Oboussier 1978, park records unpubl.); mean and sample number not given Record horn length is 100.3 cm for a pair of horns from the Chercher Mts (Rowland Ward) Key References  Atickem et al. 2011; Brown 1969a; East 1999; Evangelista et al. 2008; Hillman 1986a, 1988a; Hillman & Hillman 1987; Maydon 1925; Refera & Bekele 2004. Claudio Sillero-Zubiri

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Tragelaphus scriptus

Tragelaphus scriptus  Bushbuck Fr. Guib Harnaché (Antilope Harnaché); Ger. Schirrantilope Tragelaphus scriptus (Pallas, 1766). Misc. Zool. p. 8. No locality cited but based on ‘Le Guib’ of Buffon, from ‘Sénégal’.

ornatus, phaleratus, pictus, powelli, punctatus, reidae, roualeyni, roualeynei, sassae, signatus, simplex, sylvaticus, tjaderi, typicus, uellensis. Chromosome number: 2n  = 33 for ?, 2n  = 34 for / (Wallace 1977).

Senegal Bushbuck Tragelaphus scriptus (scriptus group).

Taxonomy  No fewer than 27 subspecies were recognized by Allen (1939) and this was reduced to 23 by Haltenorth (1963); only nine were accepted by Ansell (1972). Kingdon’s (1997) treatment largely echoed that of Ansell (1972), except that he did not recognize powelli (which Haltenorth [1963] had included in decula), and accepted two subspecies found in the montane regions of E Uganda (barkeri and heterochrous, which Ansell [1972] included in bor) for a total of ten subspecies. Grubb (1985) recognized six subspecies in East Africa, and Meester et al. (1986) three from southern Africa. Most of the variation within the species has been analysed based on coat colour, patterns of black/dark brown and white markings on the legs, lower shoulder, belly line and haunches, and patterns of spotting and stripes. It might be that some of the characteristics are controlled by hormones or diet given the large variation that can occur even at a single site (Kingdon 1982). Recently, continent-wide variation in the Bushbuck has been investigated using mitochondrial markers (control region, cytochrome b) revealing that the Bushbuck comprises two genetically divergent lineages: a scriptus group from the north-western half of the continent, and a sylvaticus group from the south-east (Moodley & Bruford 2007). Further molecular analysis (Moodley et al. 2009; and see Hassanin et al. 2012) suggests that these may represent two distinct species as suggested by early taxonomists (e.g. Sclater & Thomas 1899/1900). The subspecific taxonomy adopted here is a combination of several taxonomies, principally those suggested by Grubb (1985, 2005) and Meester et al. (1986), with modifications from Kingdon (1997) and the study by Moodley & Bruford (2007). Synonyms: barkeri, bor, brunneus, cottoni, dama, decula, delamerei, dianae, dodingae, eldomae, fasciatus, fulvoochraceus, gratus, haywoodi, heterochrous, insularis, johannae, knutsoni, laticeps, locorinae, makalae, massaicus, meneliki, meridionalis, meruensis, multicolor, nec, nigrinotatus, obscurus, olivaceus,

Description  A medium-sized antelope with a great variation in coat colour and patterning across its wide geographical range. Head red or brown with white spots or flashes on the cheeks below the eye. Ears large with white around the rim and black flash inside the ear towards the base contrasting with the pink above it. Females and young are generally red whereas adult ?? become progressively darker with age, sometimes to a dark brown and almost black. All age/sex groups have a white belly extending back to a broad, woolly tail that is red/brown above and white below. A dorsal crest of lighter coloured hair is present down the back of the animal. Legs are long and thin, with white fetlocks above black hooves. Hindquarters are more developed than the forequarters. The back is rounded due to shorter forelimbs (metacarpus is 20% shorter than metatarsus; Hofmann 1973). Western forms have a ‘harness’ with vertical and horizontal white body stripes whilst the eastern and southern ‘sylvan’ populations are plainer with fewer light streaks or spots on the flanks or haunches. Some montane forms have longer hair around the neck and face. Young are similar in colour to adult // but stripes are more marked. Males are larger in body size than //, sometimes up to 50% larger (Wilson & Child 1964). Females have two pairs of inguinal nipples. Only ?? bear horns, which are smooth and have one twist in the spiral and can be straight or slightly kinked. Simpson (1973), who discussed tooth replacement and wear in T. s. ornatus, recorded cases where skulls had supernumerary teeth (one specimen had 16 premolars, four of which probably developed from the twinning of tooth buds). Geographic Variation  Based on molecular data, Bushbucks can be grouped into two main groups: a scriptus group that includes most of the Bushbucks found in West and central Africa, and which tend to be more clearly striped and spotted, and a sylvaticus group that is found in East and southern Africa and is generally plainer in coat patterns. Between these two groups several forms occur that show some intermediate phenotypic characteristics, but which can be clearly assigned, at least on genetic analysis, to one of the two groups (Moodley & Bruford 2007, Moodley et al. 2009). Grubb (1985) identified four main groups, including the scriptus and sylvaticus groups, and then grouped the intermediate subspecies into a further two groups, namely decula and fasciatus (but later lumped fasciatus with sylvaticus; see Grubb 2005). Scriptus group (Harnessed Bushbucks): T. s. scriptus (incl. gratus, typicus and obscurus) (Senegal Bushbuck):West Africa, from Senegal to Liberia. Males rich dark rufous with black suffusions, with vertical stripes down the back and horizontal 163

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Bushbuck Tragelaphus scriptus polymorphism (see key opposite).

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A   A dark morph of West African scriptus male B   West African scriptus male C   West African scriptus female D   A light morph of West African scriptus male E F G

Three males from a single locality (Kasulu, Tanzania) to show variation in pattern and colour (dama)

H   Pale male from Ujiji, Tanzania (dama) I    Ethiopian Mts male (meneliki) J    Pale male from Moyale, North Kenya K  (K2) L M

Four animals from a single high-altitude locality (Mt Elgon) (K + K2: female and her young) (heterochrous)

N    Male from Fort Portal, Uganda (dama)

stripes with spots on the shoulders, flanks and hindquarters; // chestnut with fewer white stripes; this subspecies has the largest number of white markings on the body. T. s. phaleratus (incl. knutsoni and johannae) (Kabinda Bushbuck): Cameroon, Gabon, Equatorial Guinea, W Congo R., W DR Congo. A montane form (knutsoni) confined to the region above the forest zone on Mt Cameroon is sometimes recognized as a separate subspecies (e.g. Grubb 2005). Tragelaphus s. phaleratus lies within the phenotypic range of T. s. scriptus (indeed, Grubb [2005] included phaleratus as synonymous with the nominate form), but without any blackish suffusions except on the withers and upper legs. A longitudinal band is often absent, particularly in //. T. s. bor (incl. cottoni, dodingae, meridionalis, pictus, punctatus, signatus and uellensis) (Nile Bushbuck): NE Nigeria, Lake Chad, E Cameroon, Central African Republic, N DR Congo, S Sudan and NW Uganda, Ethiopian lowlands. Males and // yellowish-brown in colour with // yellower (in ?? this contrasts strongly with black markings on belly); ?? have relatively short and straight horns. Molecular data evidence intergradation with the Niger basin form (see below), with T. s. phaleratus in Cameroon and T. s. dama in NE DR Congo and NW Uganda. T. s. decula (Abyssinian Bushbuck): N, S and W Ethiopia and parts of Eritrea. Small in size, with short horns in ?; ? is dull sandy-ochre with grizzled hairs, giving appearance of dark brown or blackishbrown coat; / light sandy-ochre with dark brown dorsal stripe

expanded into broad blackish saddle; few white markings on coat except an upper dorsal white stripe (Grubb 1985). Here attributed to the scriptus group following Moodley & Bruford (2007). Moodley & Bruford (2007) distinguished three further distinct forms geographically intermediate between T. s. scriptus and T. s. phaleratus: two forms from the Upper Volta and Lower Volta, and a third form ranging from Togo to about the Cross R. in E Nigeria. Sylvaticus group (Sylvan Bushbucks): T. s. sylvaticus (Cape Bushbuck): southern and eastern coast of South Africa, from the Western Cape to S KwaZulu–Natal. Male dark brown, with hair at base of neck often lost in old ??; little or no trace of white stripes on body (typical of T. s. ornatus), but some spots on sides of belly and hindquarters; / fawn brown with some spotting as in ? (darker in colour than T. s. ornatus). Intergrades with T. s. delamerei and T. s. roualeyni in NE South Africa. T. s. roualeynei (Limpopo Bushbuck): south-east Africa (NE South Africa, S Mozambique, E Swaziland and S Zimbabwe, E Botswana). Phenotypically intermediate between T. s. ornatus and T. s. sylvaticus. Males dark red and // darker than T. s. sylvaticus; both sexes lack the vertical white stripes on the coat and are less spotted than T. s. ornatus. Treated as distinct by Meester et al. (1986), but considered synonymous with sylvaticus by Grubb (2005). 165

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T. s. ornatus (Chobe or Zambezi Bushbuck). South-central Africa (N Zimbabwe and N Botswana, NE Namibia, S DR Congo, C and E Zambia, and ranging into Tanzania). Males dark red with distinct vertical stripes (up to eight) on coat and extensive white throat, and // chestnut with fewer white stripes; both sexes have extensive white spots on shoulders and particularly the hindquarters. Due to its pronounced harness pattern, initially classified with the scriptus group, but more appropriately included in the sylvaticus group (Grubb 2005, Moodley & Bruford 2007). Treated as a distinct subspecies by both Grubb (1985) and Meester et al. (1986), but considered synonymous with sylvaticus by Grubb (2005). T. s. dama (incl. barkeri, dianae, heterochrous, locorinae, sassae and simplex) (Uganda or Kavirondo Bushbuck): Uganda, including Mt Elgon (heterochrous),WTanzania, the Albertine Rift (dianae) and the Imatong Hills in SE Sudan (barkeri). Male yellowish-chestnut to chestnut on haunches; belly and upper limbs dark grey to black; / chestnut above and buff below; flank and rump spots occur with some vertical striping; large in size with large horns in ??.The montane forms (barkeri and heterochrous) are considerably darker, and were considered distinct subspecies by Kingdon (1982, 1997). Intergrades with T. s. bor in the north: the montane forms occur in the same area as T. s. bor, but are separated altitudinally. Intergrades with T. s. ornatus in the south-west and T. s. delamerei in the east.This subspecies is considered synonymous with sylvaticus by Grubb (2005). T. s. delamerei (incl. eldomae, haywoodi, meruensis, olivaceus and massaicus) (Maasai Bushbuck): C and E Tanzania, C and S Kenya, Mozambique, Malawi, E Zambia, E Zimbabwe and E South Africa. Large size, with large horns in ??. Male dark brown, lighter at rump; belly and upper limbs dark brown; / reddish-chestnut with yellowish cheeks contrasting with white throat. The montane form (haywoodi) from Mt Kenya, Kilimanjaro, Meru, Hanang and the Gregory Rift is phenotypically distinct, and is given subspecies status by Moodley & Bruford (2007). Intergrades with T. s. dama, but with less rump and flank spots. Treated as distinct by Grubb (1985), but later considered synonymous with sylvaticus by Grubb (2005). T. s. fasciatus (Somali Bushbuck): NE Tanzania, E Kenya, E Somalia and presumably along the Juba R. into E Ethiopia. Adult ?? usually greyish-ochre, sometimes brownish-grey; // yellowochre; both sexes have greyish necks; relatively large in body size with large, widespread horns in the ?; dorsal neck hairs relatively short in the ?. Grubb (1985) regarded it as a very distinctive subspecies, though he later included it in sylvaticus (Grubb 2005). T. s. meneliki (incl. powelli) (Menelik’s Bushbuck): Ethiopian Highlands. Adult ? dark chocolate brown (though some are paler), with only white spots on rump as coat markings; / dark russet brown with no dorsal stripe and 1–3 buffy spots on haunches; horns in ? relatively narrow and straight. Here attributed to the sylvaticus group following Moodley & Bruford (2007).

Tanzania and Zambia. Larger with longer hooves; shaggier // are most similar but ?? are darker grey/brown with long sharply keeled horns; generally found in waterlogged or swampy habitats. T. angasii. Thicket and forest habitats of Malawi, Mozambique, Zimbabwe, South Africa and Swaziland. Larger; in ?? horns longer (typically >560 mm), well-developed dorsal crest, plus fringe of long dark hair along throat and ventral surface to between hindlegs, and rufous ‘socks’ (carpal and tarsal) contrast with dark upper legs; in // vertical stripes more well developed than southern forms of Bushbuck, and tail longer. T. eurycerus. In West Africa, from Sierra Leone to Benin, and from the Sanaga R. in Cameroon along the lowland forest zone to E DR Congo; isolated montane populations exist on Mt Kenya, Mau and Eburu forests, and the Aberdares. Larger and stockier; dark russet red colour, which tends to darken on the forequarters and limbs with age; black legs with patches of white distinguish this species from the Bushbuck, which has more red/brown and white legs. Distribution  Endemic to Africa. The Bushbuck is among the most widespread of African antelopes, occurring in some 40 African countries, more than any other antelope species (East 1999). The species ranges widely from Mauritania, Senegal and Guinea-Bissau across West Africa, south of the Sahara, around the forests of the central Congo Basin to north-east Africa (SW Eritrea, W and C Ethiopia) then southwards throughout East Africa and the more mesic areas of southern Africa to around Bredasdorp in the Western Cape of South Africa (East 1999, Skinner & Chimimba 2005). It also occurs on some small islands off the African mainland including Orango N. P. in the Bijagos Archipelago off the coast of Guinea-Bissau (East 1999). The only sub-Saharan country from which it has not recently been recorded, and where it may formerly have occurred, is Lesotho (Lynch 1994). Habitat  Bushbucks occur in a wide variety of habitat types across Africa, being absent only from arid and semi-arid regions and dense

Moodley & Bruford (2007) distinguished three further forms: two in the Luangwa Valley and the middle Zambezi Valley, and a third from Angola; Angolan animals were referred to the subspecies T. s. ornatus by Crawford-Cabral & Veríssimo (2005). Similar Species Tragelaphus spekii. Swamp and swamp forest-dwelling species from central Africa and, marginally, from West Africa to Uganda, W

Tragelaphus scriptus

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closed-canopy forest (as typical of parts of DR Congo). They range from low altitudes near the coast in southern, East and West Africa (Keep & Broker 1986, Coates & Downs 2006) up to 4000 m on the mountains of East Africa (Plumptre 1991, East 1999). Bushbucks require some cover and are most often associated with woodland, scrub and forest edges. Bushbucks appear reluctant to utilize areas away from surface water, which may be explained by them exhibiting high evaporative heat loss response, mainly due to the production of strongly diluted urine (Schoen 1971). In some areas, Bushbucks exhibit seasonal movements; in the Zambezi Valley, Simpson (1974a, b) recorded them dispersing from the riverine underbush in the warm, wet summer months (Sep–Mar), to the thickets, where water is temporarily available, returning to the riverine associations during the drier periods. In Mole N. P. in Ghana, Bushbucks preferred marshy habitat over riverine and open savanna (Dankwa-Wiredu & Euler 2002). Bushbucks are often able to survive in humandominated landscapes such as on large farms as well as on coffee and timber (eucalypt and pine) plantations (Odendaal & Bigalke 1979, Odendaal et al. 1980, Schmidt 1983) and may cause severe damage to seedlings (de Zwaan 1977, Schutz et al. 1978). The species has disappeared from some of its former range, particularly in drier areas where habitat destruction and desertification have occurred, but is expanding where equatorial forest is being opened up by agriculture and logging, creating more forest edge and bush (East 1999).

Lateral, palatal and dorsal views of skull of Bushbuck Tragelaphus scriptus.

Abundance  Owing to the cryptic nature of this species, abundance is generally underestimated by aerial flights and gives densities of 0.01–0.5/km2. Ground surveys (Plumptre 1991, Plumptre & Harris 1995, Mizutani 1999) and individual recognition (Jacobsen 1974, Waser 1975a, Allsopp 1978) provide very much higher, more accurate, estimates. In grassland with thickets on the Mweya Peninsula in Queen Elizabeth N. P., W Uganda, Bushbucks occurred at densities of about 26/km2 (Waser 1975a), whilst in mixed-forest, bushland and grass in Nairobi N. P., Kenya, density was estimated to be similar at 30.1/km2 (Allsopp 1978). In montane forest of the Virunga Mts, Rwanda, estimated densities (using faecal counts) were 11.1–44.2/km2 in different habitats with highest values in or at the edge of woodland (Plumptre & Harris 1995). In riverine forest along the Sengwa R., Zimbabwe, Jacobsen (1974) recorded the highest-known densities of up to 67/km2. In ground surveys that counted several species at the same time in central and western Africa, lower estimates from 0.08 to 2.0/km2 have been obtained (East 1999, Fischer & Linsenmair 2001a, Brugière et al. 2005). For the Ethiopian Highlands (Denkoro Forest), a mean population density of 11.8/km2 was reported (Yazezew et al. 2011). East (1999) estimated total population size in Africa at about 1.34 million individuals, which is probably an underestimate.

recorded associating with Bushbucks as possibly a strategy to avoid predation by Leopards Panthera pardus (Allsopp 1971, Elder & Elder 1971, Wronski 1996). It is unclear why Bushbucks bark, given that they are principally a solitary species. It is possible that the bark indicates to a predator that it has been spotted, which may reduce the predator’s likelihood of attacking. Possibly, other Bushbucks in the vicinity tend to be more related (e.g. matrilineal clan members; Wronski & Apio 2006, Apio et al. 2010) so their genes may be passed on more successfully as a result of warning them. Similarly, the white flashes of the underside of the tail when fleeing may also serve as a warning to others. Kingdon (1982, 1997) suggested that barks may function as indicators of rank, status, identity and physical movement within a local community of Bushbucks. Bushbucks are the smallest of the tragelaphines, and yet have been the most successful in terms of distribution across Africa. Their small size and cryptic colouring and behaviour have probably allowed them to survive in smaller habitat patches and around human settlements better than the larger species. Cover seems to be the most limiting resource for Bushbucks, while food, competition and predation had no effect on female home-range size in Queen Elizabeth N. P. (Wronski et al. 2006e). Unlike other tragelaphine antelopes, Bushbuck horns are small and tend to spiral much less. This could be because this species lives in dense habitat and relies on flight into dense habitat to avoid predation (similar to strategies duikers use). It has been suggested that the horns of larger tragelaphines have evolved due to sexual selection rather than for defence (Walther 1964a) and it is possible that the Bushbuck’s small size and greater vulnerability to predation has selected for smaller and straighter horns.

Adaptations  The widespread distribution of the Bushbuck across several habitat types implies an exceptionally adaptable physiology. The cryptic coat patterning provides camouflage in a wide variety of habitats. They apparently are excellent swimmers, taking to water to feed and also to flee from predators. Their large ears are probably an adaptation to detect predators and are particularly large in calves relative to body size. Bushbucks typically give a warning bark when danger threatens and other species use this as a warning too; baboons (Papio spp.) and Vervet Monkeys Chlorocebus pygerythrus have been

Foraging and Food  Bushbucks are primarily browsers, tending to prefer herbs selected from between grasses, or young leaves of shrubs and trees; they also select young green leaves of grasses (Wilson & Child 1964, Jacobsen 1974, Simpson 1974c, Odendaal 1983, AllenRowlandson 1986, Plumptre 1995, Seymour 2002, Apio & Wronski 2005,Yazezew et al. 2011; and see Gagnon & Chew 2000, Cerling et al. 2003, Sponheimer et al. 2003b). In a study of food preferences in South Africa, Bushbuck showed a strong selection for plants with high digestibility and low fibre content (Haschick & Kerley 1997a), but a 167

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Bushbuck Tragelaphus scriptus adult female.

Bushbuck Tragelaphus scriptus young adult male.

relatively small bite size and bite rate (Haschick & Kerley 1997b). The preferred feeding height was determined as 52.5 ± 3.8 cm from the ground (Haschick & Kerley 1996). In sites with strong seasonal variation, the diet seems to consist of herbs and young grass leaves in the wet season, replaced by the leaves of shrubs and trees in the dry season (Wilson & Child 1964, Jacobsen 1974, Okiria 1980, Smits 1986, MacLeod et al. 1996). Where there is little variation between seasons in a montane environment, herbs and grasses are selected throughout the year, particularly the grasses Festuca schimperi/Agrostis spp. and herbs, Impatiens and Solenostemon sylvaticum (Plumptre 1991, 1995). In drier areas such as Mole N. P., Bushbuck primarily consume herbs and rarely take grass unless young and green; Raindia macaritha, R. captiatum and Eurene lobata were the dominant foods in Mole N. P. (Dankwa-Wiredu & Euler 2002). Dichrostachys cinerea, Pavetta albertina and Indigofera spp. were preferred foods in Queen Elizabeth N. P. (Apio & Wronski 2005). In South Africa (Woody Cape N. R.), Bushbuck also fed on dicotyledonous herbs, in particular Lycium afrum and Schotia afra (MacLeod et al. 1996). Staple foods consumed throughout the year in Zimbabwe included leaves of three tree/shrub species, namely Combretum mosambicense, Grewia flavescens and Trichilia emetica (Jacobsen 1974), while in a study in Mpumalanga, South Africa, Acacia nigrescens, Combretum heroense, Ficus spp. and Ziziphus mucronata comprised up to 20% of the diet in all seasons (Seymour 2002). Where available, the pods of Acacia species and other fallen fruit are consumed, including Ficus, Ricinodendron, Cordia, Balanites and Diospyros (Wilson & Child 1964, Jacobsen 1974). Seymour (2002) has demonstrated how Bushbucks may be out-competed, during times of nutritional stress, by Nyalas Tragelaphus angasii, since the latter species is a mixed feeder with a greater browse height and

therefore has a competitive advantage in the late dry season when available browse is limited. Mycophagy may be an important mechanism by which Bushbucks are able to exist in areas such as forests and plantations in which trace elements, such as copper, are not readily available (Odendaal 1983). Geophagia also has been recorded (Jacobsen 1974). Bushbucks may enter agricultural fields at night and in the early morning to eat crops and are considered a pest in some places. For example, around Parc National des Volcans in Rwanda they consume the leaves of potatoes, maize, cabbages, beans, peas, pyrethrum, wheat and tomato plants (Plumptre & Bizumuremyi 1996). Where undisturbed, Bushbucks feed throughout the day and night (Waser 1975b,Wronski et al. 2006b) in regular cycles.Where there is a likelihood of detection and predation during the day they will feed mainly at night but may rest in dense cover, occasionally feeding during the day. Their movements are often associated with dawn and dusk. In Queen Elizabeth N. P., Bushbucks moved towards the thickets along the river at dawn, where it is presumably cooler during the heat of the day, and then back to the more open ridge at night (Waser 1975b). In the Virunga Mts, Bushbucks moved towards open meadows at dusk and back into the denser woodland and nettles during the day around the Karisoke Research Station (A. Plumptre pers. obs.) and they fed throughout the period they were in dense nettles (Plumptre 1991). Social and Reproductive Behaviour  Bushbuck are generally solitary or in pairs, with single ??, single // and male–female pairings accounting for most observations in the areas where the species has been studied: 91% in N Zimbabwe (Wilson & Child 1964), 79% in N Botswana (Elder & Elder 1971), 74% in W Uganda (Wronski 2004) and 71% in Knysna, South Africa (Odendaal & Bigalke 1979).

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Other small associations usually consist of adult //, adult // with young or subadults, or two or more adult ?? (though such male groupings are both unusual and ephemeral). Larger groups of 9–12 have been recorded (Simpson 1974a, Simbotwe & Sichone 1989, Brugière et al. 2005, A. Plumptre pers. obs.). In the Virunga Mts, Bushbucks were generally solitary during the day when in dense vegetation but when they moved to clearings in the forest to feed in the evening it was not uncommon to see several individuals together (A. Plumptre pers. obs.). In this case the animals were not in the strict sense forming a group but were associating with each other only because of a common attraction to the same resources. They did not move and function as a coordinated group.Wronski et al. (2009b) compared group sizes among six different populations and found no significant difference in relative frequencies of group size categories. A study in Uganda showed that // are organized in matrilineal clans, sharing a common home-range and jointly defending that area against non-related // (Wronski & Apio 2006, Apio et al. 2010). Home-range overlap between related // was significantly higher than that between non-related individuals. Agonistic interactions between // appeared significantly more often between non-related // but rarely between kin (Wronski & Apio 2006). Moreover, // keep close social bonds (e.g. reciprocal grooming, also between adult related //; Walther 1964a, Wronski et al. 2006d) and individuals communicate through localized defecation sites (mostly females to males and females to females of the same clan; Wronski 2006b). Localized defecations function neither in terms of parasite avoidance nor as a consequence of allelomimetic behaviour (Apio et al. 2006a). Individual home-ranges overlap (Jacobsen 1974,Allsopp 1978,AllenRowlandson 1986, Coates & Downs 2005, Wronski 2005, Wronski & Apio 2006); some studies have shown that there is little defence of territories (Waser 1975a, Allsopp 1978), while others have recorded pronounced territoriality (Verheyen 1955, Walther 1964a, Jacobsen 1974). Owen-Smith (1975) discusses territoriality in Bushbuck in light of social patterns that have evolved in tragelaphine antelopes. Recent

studies have revealed further insights into the question of territoriality in Bushbuck. In a study in KwaZulu–Natal, South Africa, Bushbucks utilized a core area within their home-ranges in which 50% of their time was spent in approximately 17.0% and 11.7% of their total home-range for ?? and //, respectively, and there was substantial overlap in core areas (Coates & Downs 2005). However, in Uganda, a considerable defence of 50–70% of home-range cores (territories) was observed (Wronski 2005, Wronski et al. 2006, 2007b, Wronski & Plath 2006). Home-range cores of adult ?? did not overlap and were therefore considered exclusive areas. Moreover, adult territorial ?? perform intense scent-marking (i.e. vegetation horning, front rubbing; Wronski et al. 2006d, 2007b) and these marked areas correspond with exclusive core areas (Wronski et al. 2006a). A dominance hierarchy as suggested by former studies (e.g. Jacobsen 1974) could explicitly be excluded (Wronski 2004, Wronski et al. 2009a). Instead, male Bushbuck exhibited two age-dependent mating tactics: adult territorial ??, and young adult non-territorial ??, which frequently attempt to mate with // although they indulge in less pre-mating behaviour, herding and monopolization of // (Apio et al. 2007). Males and // advertise their presence and position by standing on termite mounds or close to vertical structures (Wronski et al. 2007a). Home-range sizes vary between sites. In Nairobi N. P., small ranges of 0.25–2.0 ha were recorded, the larger ranges being occupied by juvenile animals (Allsopp 1978), whereas the homerange of a dominant ? in Sengwa Valley, Zimbabwe, was 5 ha (Jacobsen 1974). In an area of valley bushveld in KwaZulu–Natal, total home-range size for ??, using minimum convex polygons, was estimated at 33.9 ha and for // at 12.0 ha (Coates & Downs 2005). In Queen Elizabeth N. P., Waser (1975a) recorded homeranges of adults varying from 6.3 to 26.5 ha for adults with one subadult ? having a range of 35.2 ha. In contrast to these recorded home-range sizes, average home-range size in Knysna Forest (South Africa, which is a more temperate climate) was 102 ha (Odendaal & Bigalke 1979), while Allen-Rowlandson (1986) recorded a mean

Bushbuck Tragelaphus scriptus display.

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home-range size of 120 ha for ?? and 60 ha for // in afromontane forest in KwaZulu–Natal. Home-range size apparently increases with decreasing density across the range of the species, and density appears to increase with an increase in rainfall during the growing season (Odendaal & Bigalke 1979). In Queen Elizabeth N. P., animals spent 38% of their time feeding, 50% resting, immobile or ruminating and 12% moving (Waser 1975b). In the montane habitats of the Virunga Mts a similar amount of time was devoted to these activities: 50% feeding, 44% resting or ruminating, 6% walking or grooming (Plumptre 1991). Similar patterns were described for Bushbuck from Denkoro Forest in the Ethiopian Highlands (Yazezew et al. 2011). Wronski et al. (2006c) investigated differences in activity patterns in relation to time of day, season, radiation, moonlight, age and sex. Bushbucks were found to show peak activities around sunrise and sunset. No difference in the mean activity rates was found between the dry and wet season. Daytime activity was not predicted by differences in sun radiation, nor was night-time activity predicted by the presence or absence of moonlight. The activity of adult territorial ?? was strongly positively correlated with that of //, whereas that of nonterritorial ?? was not correlated with the activity of //. As noted already, Bushbucks often associate with baboons (Papio spp.) at various sites across the continent and it is thought that both species benefit from the association with the Bushbuck’s sense of smell and the baboon’s ability to spot predators from trees (Elder & Elder 1971, Allsopp 1978, Wronski 1996). Bushbucks were accompanied by baboons in about 35% of sightings in Chobe N. P., Botswana and Nairobi N. P. (Elder & Elder 1971). Males threaten other ?? by a stiff-legged walk and lateral presentation, with the head up (Walther 1964a, Kingdon 1982, Estes 1991a, Wronski et al. 2006d). They raise the hairs on the dorsal crest, arch the back and raise the tail to expose the white underside and may circle each other. In Uganda, territorial ?? were observed escorting intruders to the edge of their territories (Wronski et al. 2006d). Occasionally, a ? will feint attack and lower his horns and horn at the ground. If closely matched, the ?? may raise the intensity of combat with head-on clashes and locking horns, and attempts made to gore the flanks of the opponent (Walther 1964a, Allsopp 1978,Wronski et al. 2006d). Defensively, a ? keeps the head low, turns away and licks the aggressor. Females also show agonistic interactions, but on a much lower scale (Wronski & Apio 2006, Wronski et al. 2006e). When approached by a predator, ?? or // freeze, sinking to the ground with the neck and head stretched out along the ground. If the danger is close, they sometimes bark and flee with their tails raised, exposing the white underside in a bright flash. The young have a bleat not unlike a goat’s call when lost from the mother, but adults do not vocalize much other then barking. When courting, ?? nuzzle the female’s genitals, lick her, rub cheeks on her rump, press the neck or head on her rump and test urine by licking the stream as it falls to the ground. In the urine test, the ?? approach with neck extended and head low, with no dorsal crest showing and the / responds by crouching, a position typically used in urinating with the hindlegs bent and the tail raised. Males tend the / while she is receptive, chasing off other ?? unless supplanted by another ? (Walther 1964a, Estes 1991a). Calves are born in areas of dense cover and lie still whenever the / is away feeding. They may spend up to four months like this

before venturing into more open habitat (Estes 1991a). At about 6 months of age young ?? leave their mother’s home-range and join a bachelor group. From this pool they challenge existing territorial adult ?? until they take over a territory (Wronski 2005). Females will stay with their mothers (and other female kin) throughout life, forming matrilines (Wronski & Apio 2006, Apio et al. 2010). Reproduction and Population Structure  Calves are born throughout the year in the wild (and in captivity, see Mentis 1972, Skinner et al. 2002), although there are certainly some seasons where more calves are dropped than others (Ansell 1960a, Allsopp 1971, Simpson 1974a, Plumptre 1991, Apio et al. 2009). In Kenya, there were peaks in births in Feb and Sep, just prior to the two main wet seasons when more nutritious forage was available (Allsopp 1971). In the Virunga Mts, births were more clumped from Jun–Sep, a drier period of the year in an environment where the cold rain can lead to mortality (Plumptre 1991). In Zambia, calves were recorded throughout the year (Wilson & Child 1964). Gestation length is around 24–35 weeks with the average being around 25–26 weeks (Allsopp 1971, Dittrich 1972, Mentis 1972) and calves dropped weigh 3–4 kg. Usually one calf is born although occasionally twins occur. Calving intervals measured in captive animals are around 35–36 weeks and a / can produce another calf within 48–54 weeks after the previous birth (Mentis 1972, Dittrich 1974, von Ketelhodt 1976a). Male calves develop horns about seven months after birth (von Ketelhodt 1976a) and ?? develop the secondary sexual characteristics of a dorsal crest and darker pelage about two-and-a-half years after birth (Kingdon 1982). Males become sexually mature from about ten months of age, but do not take part in breeding until they are at least two years old (Mentis 1972). Females become sexually mature at 14–19 months (Dittrich 1972, Mentis 1972, Simpson 1974a, Allen-Rowlandson 1986). In Zimbabwe, the mean age at maturity was about 11 months in both sexes, coinciding (in ??) with a combined testes weigth of about 15 g (Morris & Hanks 1974). Sex ratios at birth are equal, but adult ratios have varied from 45–115 ?? to 100 // in different sites (Wilson & Child 1964, Mentis 1972,Waser 1975a). Adult sex ratio obtained from a tsetse fly control area in Zambia was reported to be 80 ?? to 100 // (or 1 ?: 1.48 /; Wilson & Child 1964). Longevity in the wild has been reported at about 12–13 years (Mentis 1972); in captivity, maximum longevity has been recorded as 16 years and nine months for T. s. sylvaticus (Jones 1993). Predators, Parasites and Diseases  Leopards (Bushbuck are a significantly preferred prey, Hayward et al. 2006a), Lions Panthera leo, Spotted Hyaenas Crocuta crocuta, Cheetahs Acinonyx jubatus, African Wild Dogs Lycaon pictus, Nile Crocodiles Crocodylus niloticus and African Rock Pythons Python sebae have all been recorded killing adults, and Robust Chimpanzees Pan troglodytes, baboons, Caracals Caracal caracal and several eagle species have been reported taking calves. African Golden Cats Profelis aurata also are probable predators (Kingdon 1982). Zieger et al. (1998c) recorded a Bushbuck found dead on a ranch in Zambia with multiple superficial abscesses in the neck region, extensive granulomatous lesions in the lung, the bronchial and

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mediastinal lymph nodes and several nodular lesions in the spleen. Histologically the lesions resembled those of tuberculosis, but mycobacteria could not be isolated. In Queen Elizabeth N. P., one reported case of rabies, a viral disease (Lyssavirus) that causes acute encephalitis was confirmed for Bushbuck (A. Apio & O. Bwangamoi pers. comm.). Round (1968) provides a checklist of helminth parasites of the Bushbuck from all over Africa. The internal parasites of the Bushbuck have been relatively well investigated, particularly in southern Africa (e.g. Boomker & Kingsley 1984, Boomker et al. 1984, 1986), and Boomker et al. (1987) provided an updated list of all helminth parasites recorded from Bushbuck in South Africa at the time, including two species of cestodes (Cysticercus spp. and larvae of Taenia spp.) and 24 species of nematodes. Zieger et al. (1998a) recorded Stilesia hepatica from an animal in Zambia. Bwangamoi (1968), Woodford & Sachs (1973), Pullan et al. (1971) and Apio et al. (2006b) report on helminth parasites from Bushbuck in Uganda. Strongyle eggs recorded in faecal samples from Queen Elizabeth N. P. include Oesophagostomum spp., Haemonchus spp., Bunostomum spp. and the eggs of the cestodes Moniezia benedini and M. expansa (Apio 2003, Apio et al. 2006b). Generally, the prevalence of gastrointestinal tract parasites in Bushbuck from Queen Elizabeth N. P. is low. Apio et al. (2006c) tested the risk of infection with gastrointestinal parasites at different foraging height levels, and suggested that high browsing levels are responsible for low parasite infestations in Bushbuck. In the same park, Woodford (1976) reported on muscular and aberrant cysticercosis, which are conditions caused by larval stages of tapeworms infesting the musculature and connective tissues of their host. Sarcosporidiosis and larval pentastomidosis were also regularly found in the tissues of Bushbucks. Amongst the herbivores of Queen Elizabeth N. P., Bushbucks were the most commonly infested species (Woodford 1976). A filarial worm (Setaria sp.) was detected in blood samples of Bushbucks in the same study area. Apio & Wronski (2004) reported further on post-parturient changes in faecal helminth egg and coccidian oocyst counts of female Bushbucks in Queen Elizabeth N. P. Eimeria spp. was the most common coccidian infection, but did not lead to any clinical or pathological signs in the study population (Apio 2003). Apio et al. (2006b) investigated also seasonal, sexual and age-related variation in helminth egg and coccidian oocyst counts in the same population. The prevalence of Moniezia spp. and strongyle eggs was significantly higher during the wet season than during the dry season, and peak counts were recorded either during or soon after a peak rainfall. The same was true for Eimeria sp. Sexual and age-related differences in the prevalence of either parasite types were not found. Other parasites recorded infesting Bushbucks include Babesia sp. (Bigalke et al. 1972) and the tremetode Schistosoma leiperi (Malek & Ongom 1984). Trypanosomiasis, the nagana pest of cattle, is a condition commonly found in Bushbucks (Ferrante & Allison 1983, Moloo et al. 1995); the parasites (e.g. Trypanosoma brucei, T. rhodesiae) are transmitted by several Glossina spp. (tsetse flies). In some areas, Bushbucks suffer heavy tick loads consisting predominantly of hard ticks (Ixodidae, De Castro & Newson 1984). The ixodid ticks of Bushbucks have been reported on by various authors (e.g. Dinnik et al. 1963,Woodford 1976,Terenius et al. 2000, Ntiamoa-Baidu et al. 2005), and Horak and co-workers (Horak et al. 1983c, 1989) determined actual tick and lice burdens of the species

in South Africa. Bushbucks have been infected with some tickborne diseases like bovine petechial fever (Davies 1993a), a rickettsial disease of cattle for which the Bushbuck is the reservoir host, as well as East Coast Fever, caused by a protozoan parasite regularly found in several tragelaphine species (Grootenhuis 1991). Conservation  IUCN Category: Least Concern. CITES: Not listed. Due to its widespread distribution, presence in numerous protected areas across its range, and its ability to survive around anthropogenically disturbed habitats, this species is not thought be threatened. The Bushbuck’s tendency to hide in dense cover has enabled it to survive in areas where hunting pressure is not too high (e.g. Parc National des Volcans in Rwanda; Plumptre et al. 1997). However, after larger-bodied species have been extirpated from an area, this species probably succumbs prior to the smaller-bodied species such as the duikers. Indeed, there are reports of Bushbuck being replaced by species such as Common Duiker Sylvicapra grimmia when human impacts (e.g. frequent burning, cattle grazing) are extensive (Kumanenge 1980). In areas where hunting pressure is severe, such as the Ankole Ranching Scheme around Lake Mburo N. P. in Uganda, the reduced competition from species that became less abundant outside the protected area (such as Bushbuck) may actually benefit other species, such as Common Duiker. Indeed, Common Duiker exhibited a significant change in habitat use around L. Mburo encroaching on the vegetation type otherwise normally used by Bushbuck (Averbeck et al. 2009a). Most bushmeat research has been carried out in dense tropical forests where this species either does not occur or is not very common. Data from Rwanda show that Bushbuck meat is significantly lower in price than domestic meat and was the bushmeat most commonly purchased (Plumptre & Bizumuremyi 1996, Plumptre et al. 1997). In Sierra Leone (Kenema & Lalehun markets, respectively), Bushbuck meat accounts for about 2.5– 6.6% of all traded animals and 3.1–23.5% of total biomass (Davies et al. 2007). The price for 1 kg of meat on a Ghanaian bushmeat market was reported as US$2.63 and the distance travelled until it reached the consumer was 65 km (Cowlishaw et al. 2007). More research is needed on the savanna/forest-edge bushmeat trade in East and southern Africa to better understand how this species is affected by the trade. Measurements Tragelaphus scriptus HB (??): 1270 (1170–1420) mm, n = 8 HB (//): 1200 (1140–1320) mm, n = 13 T (??): 210 (190–240) mm, n = 8 T (//): 200 (190–220) mm, n = 13 HF c.u. (??): 300 (290–320) mm, n = 8 HF c.u. (//): 280 (270–300) mm, n = 13 E (??): 135 (121–152) mm, n = 8 E (//): 136 (127–152) mm, n = 13 Sh. ht (??): 700 (640–740) mm, n = 8 Sh. ht (//): 640 (610–670) mm, n = 13 WT (??): 42.0 (29.0–54.0) kg, n = 15 WT (//): 28.0 (24.0–34.0) kg, n = 16 Zambia (Wilson & Child 1964) 171

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T. s. bor* HB (??): 1220–1310 mm, n = 6 HB (//): 1180–1260 mm, n = 3 Sh. ht (??): 780, 780 mm, n = 2 Sh. ht (//): 700, 720 mm, n = 2 T. s. decula* HB (//):1090, 1090 mm, n = 2 Sh. ht (//): 650, 670 mm, n = 2

T. s. delamerei* HB (??): 1250–1540 mm, n = 8 HB (//): 1070–1380 mm, n = 3 Sh. ht (??): 770–910 mm, n = 8 Sh. ht (//): 620–760 mm, n = 3 *Eastern Africa (Grubb 1985) The longest pair of horns of Bushbuck on record is a pair from Tolwe, North West Province, South Africa, which measured 51.4 cm (Rowland Ward)

T. s. fasciatus* HB (??): 1440, 1500 mm, n = 2 HB (//): 1080, 1520 mm, n = 2 Sh. ht (??): 860, 870 mm, n = 2 Sh. ht (//): 550, 710 mm, n = 2

Key References  Allen-Rowlandson 1986; Allsopp 1970, 1971, 1978; Apio et al. 2007, 2009, 2010; Grubb 1985, 2005; Jacobsen 1974; Kingdon 1982; Plumptre 1991, 1995; Simpson 1974a, b, c; Walther 1964a;Waser 1975a, b; Wilson & Child 1964; Wronski 2005; Wronski & Apio 2006;Wronski et al. 2006a, b, c, d, e. Andrew J. Plumptre & Torsten Wronski

Tragelaphus spekii Sitatunga Fr. Sitatunga; Ger. Sumpfantilope Tragelaphus spekii Speke, 1863. Journal of the Discovery of the Source of the Nile, p. 223. Tanzania, Karagwe, E of L. Victoria, at a lake named ‘Little Windermere’ by Speke; identified as Bukoba district, L. Lwelo, 2° S, 30° 57’ E by Moreau et al. (1946: 441).

Sitatunga Tragelaphus spekii gratus male.

Sitatunga Tragelaphus spekii gratus female.

Selous (1881) found the peoples of the lower Chobe and central Zambezi Rivers using the name ‘situtunga’ (sic), the latter confirmed by Ansell (1978) who remarks that the Lozi and Tonga of Zambia use the name ‘sitatunga’.

description, and as this is clearly Speke, he is the authority and spekii (the genitive of the Latinized Spekius) is then the original and correct spelling (Grubb 2004). As many as ten subspecies have been described, mainly based on hair texture, pelage colour and absence or presence of body stripes and spots. However, hair texture probably varies with climate, and pelage colour and presence of stripes and spots are highly variable even within the same population (some individuals are born with stripes and spots and others without). Furthermore, pelage colour darkens with age in some individuals, especially in older ??, and stripes and spots fade with age, again especially in ?? (E. Stokes pers. comm.). Sitatunga are probably monotypic, but until verified, at best only three subspecies, centred on different drainage systems, are provisionally retained (Kingdon 1982; and see Ansell [1972] who listed five). However, a

Taxonomy  The scientific name has been given variously as spekei or spekii and the authority as Sclater or Speke. J. H. Speke described a young nzoé (the Kiswahili name for Sitatunga) in his Journal published in 1863 and stated in a footnote that the animal had been named by P. L. Sclater as Tragelaphus spekii. A year later Sclater published a description in a scientific journal under the heading Tragelaphus spekei sp. nov. Although Sclater’s contemporaries clearly regarded him to be the authority, the rules of Zoological Nomenclature acknowledge the person who provided the first 172

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Lateral and palatal views of skull of Sitatunga Tragelaphus spekii.

comparison of recent field descriptions spanning these basins (E. Stokes pers. comm., C. Thouless pers. obs., J. May & R. Lindholm pers. obs.) reveals that these three subspecies cannot be reliably distinguished on the oft-cited characteristics of pelage colour and pattern (see Geographic Variation). Synonyms: albonotatus, anderessoni, baumii, gratus, inornatus, larkenii, selousi, speckei, spekei, sylvestris, typicus, ugallae, wilhelmi. Chromosome number: 2n = 30 (Wurster & Benirschke 1968). Sitatungas have unusually large sex chromosomes, the X chromosome comprising 13% and the Y 7.3% of the haploid chromosomal complement in contrast to the mammalian norm of 5% (Wurster et al. 1968). Only one other member of the Bovidae, the Blackbuck Antilope cervicapra, has this peculiarity. Description  A medium-sized, swamp-dwelling antelope, exhibiting quite marked sexual dimorphism. The widely splayed, very long and narrow hooves (more than half the length of the foot) are unique. Adult ?? have long pelage (4–7 cm), which is coarse to silky, dense or sparse, and uniform dark-chocolate brown, greyish-brown to grey. Top of the head and ears dark grey to dark brown, with a white chevron forward of the eyes; cheeks paler with one or two white spots, front of upper lips, chin and eyebrows white, inner ears with broad white margins. There is a white band across the throat and another across the chest. Body stripes and spots conspicuous, faint or absent. An inconspicuous, narrow, white stripe along the mid-line of the back is occasionally present. Tail dark brown at centre with white tip above, all white below. Legs same colour as body, sometimes darker at knee joint. Underside of feet naked. Adult // similar to ??, but smaller and less robust especially about the neck (see Measurements) and shoulders. The pelage is uniform bright to pale reddish-brown (usually), drab brown or rarely brownish-grey. Vertical white stripes across back, lateral stripe along flanks and white spots on thighs either conspicuous, faint or absent. Dark brown stripe usually along mid-line of back. White chevron forward of eyes usually less

Tragelaphus spekii

distinct than in ? and sometimes absent. Calves woolly coated, more vibrant, but otherwise similar to // in pelage colour and body stripes and spots. Females have two pairs of inguinal nipples. Horns, present in ?? only, are long and smooth, with spirals of up to two twists, almost triangular in cross-section, tips ivorycoloured, spread highly variable within populations. Geographic Variation T. s. spekii (Eastern or Northern Sitatunga) (includes larkenii, ugallae and wilhelmi): L. Victoria basin. Only faint shadow stripes on body of both sexes. T. s. gratus (Western Sitatunga) (includes albonotatus): Congo Basin, West Africa and Sudan. Bold body stripes and spots in both sexes, but fading in adolescent ??. T. s. selousi (Southern or Zambezi Sitatunga) (includes inornatus): Bangweulu, Zambezi and Okavango basins. Body stripes and spots often present in // and young, but absent in ??. Meinertzhagen (1916) erected the name sylvestris for Sitatungas on Nkose I. in the north-west corner of L. Victoria, which had stouter and stronger hooves than a population on a larger island nearby. Nkose I. is 50 km. They do not usually frequent large expanses of swamp habitat. Bongos generally do not wallow and only cautiously enter deep water. However, during a widespread Stomoxys fly outbreak that caused a major die-off in N Congo in 1997, they did wallow in muddy pools along roads and deep water in forest clearings (Elkan et al. 2009). Abundance  The Mountain Bongo has been reduced to less than a few hundred individuals (East 1999). Pointing out that the wet season dispersal of Bongos in Aberdares N. P. in Kenya depended on low-level secondary growth on abandoned cultivation sites, Kingdon (1982) thought that the long-term survival of this confined population would be compromised if trees in and around these sites were permitted to reach climax and shade out low-level herbage. The formerly cultivated areas are now dominated by woody shrubs (such as Toddalia asiatica) that may be somewhat depauperate in the forbs that Bongos prefer (or may not offer the type of cover that Bongos like) (L. Estes pers. comm.). The status of the Lowland Bongo in West Africa is uncertain or rare with populations fragmented and declining in many areas (East 1999). In contrast, thousands of Lowland Bongos probably still exist in central Africa. However, they are rare over large areas of closedcanopy forests of N Congo and in the Ituri Forest, DR Congo (Elkan 2003, S. Blake pers. comm., J.A. Hart pers. comm.). The highest recorded density of Bongos occurs in the forest–savanna transition zones in S Sudan (Hillman 1986b). After Sudan, some of the highest concentrations of Lowland Bongos are found discontinuously in Cameroon, Central African Republic and Congo. Detailed ecological surveys in the Ndoki-Likouala lowland forests of N Congo demonstrated that Bongos occur in highly clumped local distributions, reaching the highest abundance near forest clearings and areas of high herbaceous growth. Although Bongos occur in unlogged primary forest such as Nouabale-Ndoki N. P., they are found in much higher abundance in adjacent logging concessions, which have secondary growth and a high density of forest clearings (Elkan 2003, S. Blake pers. comm.). There are also poorly understood differences in abundance. For example, in both logged and unlogged forests in the Lobeke area of Cameroon there was higher Bongo abundance than for any areas surveyed in N Congo (Elkan 1995, 2003).

Bongo Tragelaphus eurycerus male skeleton.

Dense forest habitat, patchy distributions, wide-ranging patterns, retiring behaviour and crepuscular/nocturnal activity patterns hinder estimation of Bongo densities. Hillman (1986b) estimated 1.2/km2 in S Sudan based on group-size observations and minerallick distribution. Remote camera trapping identified 65 ?? and 109 // visiting the Mombongo forest-clearing complex in the Kabo forest concession in N Congo over a six-year period. Using mark-recapture techniques, Elkan (2003) estimated 139 (± 33) Bongos in the Mombongo area during a one-month period in 1996. East (1999) estimated 28,000 Bongos remaining, with Mountain Bongo populations in the few hundreds, and populations of Lowland Bongo in the order of a few thousands in the west, and tens of thousands in the central African forest zone. However, these estimates must be considered with caution and a large error margin. Patchy distribution, population declines irrespective of habitat availability, rapid ecological change in unstable habitats, and wide ranging behaviour call for extreme caution in extrapolating local abundance estimates to density calculations for large areas (Elkan 2003). Adaptations  With the high point of the body in the lumbar region and head frequently lowered, short legs compared with other tragelaphines (Ralls 1978), and horns that can be laid against its back, a Bongo can run full speed through dense forest. Although they prefer to go under or through objects, rather than over them, they are (like other tragelaphines) capable jumpers. Horns of both sexes, but particularly in the /, assist in defending young against predators (Kingdon 1982), and are sometimes employed to break high branches. They are also able to rear up on their hindlegs to reach leaves and twigs at heights of 2.5 m above ground. Although they lack or have very rudimentary external glands, Bongos secrete strong scents from their skin. This odour is hypothesized to assist in locating other individuals despite the dense habitat in which they live. The Bongo’s white stripes and its reddish-maroon colouring blend in to the forest’s filtered light making detection difficult. Kingdon (1982) noted that although a full frontal view of a Bongo can be quite conspicuous, animals often offset this by turning their striped cryptic backs on a disturbance. By keeping very still in the 181

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Bongo Tragelaphus eurycerus myology.

face of any potential threat, Bongos tend to leave the option of flight to the last moment. Bongos are most active in the evening, night and early morning when, between bouts of feeding and using mineral licks, they often walk long distances at several kilometres/hour. There is one cameratrap record of an old / (and occasionally her calf) spending several hours at a salt-lick in broad daylight (L. Estes pers. comm.). Bongos typically ruminate and rest in the hotter parts of the day (Hillman 1986b, Elkan 2003); Gilbertiodendron forest habitat with relatively open understorey as well as dense vine areas were frequently used for resting in N Congo (Elkan 2003) while bamboo habitat provides refuge to Mountain Bongos (Kingdon 1982). Foraging and Food  Diet varies greatly (107 species of food plants recorded in Sudan, 100 in Central African Republic, >80 in N Congo), but consists primarily of browsing on dicotyledonous plants, with some seasonal grazing on grasses (Hillman & Gwynne 1987, Klaus-Hugi et al. 1999, Elkan 2003; and see Gagnon & Chew 2000). These observations are borne out also by studies involving carbon isotope analyses (at least, on Mountain Bongos) (Cerling et al. 2003). Bongos prefer young leaves and new growth in recently burned or perturbed areas (i.e. post-timber exploitation, tree falls). In central Africa, just after early rains, Bongos concentrated feeding on seasonally palatable grasses, the new leaves of the highly aromatic invasive herb Chromoleana ordorata, and Ipomoea involucrata, Macaranga barteri, Triumfetta cordifolia, Sida alba, Paspalum conjugatum, Alchornea cordifolia, Adenia cissampeloides, Cissus dinklagei, Manniophyton fulvum, Diodia sp., Costus sp. found along roadsides and forest openings (KlausHugi et al. 1999, Elkan 2003). During the dry season, they moved further into forest as forage quality and water decreased in ecotone areas. Important food plants included Thomandersia laurifolia, Palisota ambigua,Whitfieldia elongata, Rinorea sp. and Barteria sp. in closed-canopy forest. Gilbertiodendron dewevrei was also occasionally an important food source during rare, intermittent masting periods when Bongos would feed for days on this super-abundant food source (Elkan 2003). Bongos submerged their heads for tens of seconds to feed on algae (Spirogyra sp.) at a pond in N Congo. They also seasonally entered cleared fields near villages for manioc leaves and shoots of corn (Elkan 2003). Mountain Bongos inhabit a very different and botanically less diverse habitat with fewer food plants available. Edmond-Blanc (1960)

identified Parochetus communis and Senicio biafrae as important food items on Mount Kenya. Impatiens spp., Sericostachys scandens, Rubus, Asplenium and Mimulopsis spp. are common in the diet at high altitudes in the Aberdares with Cassia didymobotrya, Vernonia auriculifera and Crotalaria agatiflora readily consumed at lower altitudes (Kingdon 1982, L. Estes pers. comm.). Other species commonly encountered as browse/ graze plants include: Cynodon dactylon, Pennisetum clandestinum, Hypoestes forskalei, Plectranthus spp., Achyranthes aspera, Cyathula spp., Vernonia amygdalina, Impatiens fischeri and I. hoehnelii (L. Estes pers. comm.). Bongos visit mineral licks where they consume water and soil high in sodium, other mineral salts and trace elements (Hillman 1986b, Elkan 1995, 2003, Klaus-Hugi et al. 1999). Their drive for salt was demonstrated by a concentration of activity around artificial saltlicks in Congo (Elkan 2003). Ingestion of soil with high clay content is thought to buffer the rumen and assist in the digestion of high levels of secondary compounds and alkaloids found in much of its browse (Klaus-Hugi et al. 1999, Elkan 2003). When licks become baked, hardpan activity decreased. Social and Reproductive Behaviour  The Bongo is a highly social antelope, and, next to the Common Eland Tragelaphus oryx, forms the largest herds of any tragelaphine (Estes 1991a). Aggregations of up to 50 animals have been recorded in the forest mosaic of S Sudan (mean = 9.1, n = 42) (Hillman 1986b), although, in dense lowland Ituri Forest, group size is often only a few individuals (range 2–8) (Hart 2001). Mean group size in N Congo was 5.7 (range 2–23; n = 93) (Elkan 2003) and in SW Central African Republic, 13 (range 2–28; n = 78) (Turkalo & Klaus-Hugi 1999). In the Aberdares, group size ranges from 2 to 21 (Kingdon 1982). Females and young of both sexes including calves accompanied by an adult ? were the most common and largest social grouping in N Congo (Elkan 2003). In contrast to //, adult ?? are often solitary; 75% (n = 114) of solitary individuals observed in N Congo (Elkan 2003) and 78% (n = 41) in S Central African Republic (Turkalo & Klaus-Hugi 1999) were ??. However, seasonally, adult ?? form temporary associations with female groups. When not associating with //, single ?? overlap in home-range, and show little or no territoriality. More than 34 adult ?? visited the Mombongo forest-clearing complex over a six-month period in 1996 in N Congo. These activity levels coincided with high levels of female group visitation of the area. Resighting of individuals in N Congo has shown that Bongos range over distances >50 km and with broad herd movements that suggest seasonal habitat shifts (Elkan 2003). Home-range sizes on Mt Kenya and the Aberdares show a distinct dry/wet season shift with, tentatively, an annual home-range of 100–300 km2 (Kingdon 1982). Fighting is rare and probably dangerous (Estes 1991a). Aggressive male–male interaction has been observed; however, ?? were not aggressive or threatening towards // during courtship (Ralls et al. 1985). Mutual tolerance in groups, lack of aggressive behaviour, and larger body size in dominant ?? indicate establishment of dominance hierarchies (Elkan 2003). No fighting was observed in N Congo and other agonistic behaviour was rare; however, ?? did displace other ?? for access to licks and young ?? were observed locking horns and strength testing. Bongos exhibit dominance through lateral presentation with head up and a slow stiff-legged walk (Hamaan 1979, Estes 1991a).

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Bongo Tragelaphus eurycerus frontal view of signal patterns.

Bongo Tragelaphus eurycerus adult female.

Bongos have a keen sense of smell. Males have been observed wandering through clearings following the scent of a group that had passed hours earlier. They also have been observed to horn ground and rub in soil leaving a strong odour (Elkan 2003). There is some evidence that their body secretions are soluble in water and run off the body in raindrops (Kingdon 1982). This would assist the scentmarking of trails and resting places. Adult ?? frequently mark small trees by rubbing their horns to scrape the bark or breaking them over at 1.25–1.5 m. Although marking of presence seems to be common, no evidence of defence of territories has been observed (Elkan 2003). Males in the wild followed potentially receptive // with a low body posture, neck outstretched and lips curled back exposing the upper gums to test for oestrus (flehmen) (Elkan 2003). Males did not herd // but tended to follow behind the group when entering and leaving forest clearings. Ralls et al. (1985) categorized Bongo male behaviour towards oestrous // in captivity as similar to other tragelaphines and most like that of the Sitatunga Tragelaphus spekii. Males regularly test // by sniffing and licking the vulva and prodding to provoke urination (Ralls et al. 1985). Flehmen behaviour followed sniffing urine, faeces, or a place on the ground where a / had recently lain. Males assumed a ‘low stretch’ posture laying their head against the female’s side often with a soft vocal ‘clicking’ sound. Mounts were preceded by ?? assuming ‘frozen’ or ‘facing away’ posture, a behaviour that may be distinctive to this species (Ralls et al. 1985). This type of behaviour was also observed in an attempted copulation observed in the wild in N Congo. Oestrous // are more restless and urinate and defecate more often than non-oestrous // (Ralls et al. 1985). Oestrous // engaged in mutual licking with ??. Copulation was preceded by circling away and then back to the ? with the / then often sniffing or licking the male’s genitals. Females assumed a ‘mating face’, head and neck extended in a concave curve, raised nose, and mouth open. Females with new calves lie out for a period prior to joining groups (Kingdon 1982, Elkan 2003). Calves stay close to the mother during the early months and then form close associations with other young Bongos. Bongos are relatively quiet animals and make few vocalizations. Males utter a loud bellow when alarmed or when entering a social group.

Reproduction and Population Structure  Females are thought to begin breeding at 2.5 years old and ?? probably at 3–4 years (Ralls 1978). There appear to be bi-annual calving peaks in both Lowland and Mountain Bongos. Peaks in Lowland Bongo birthing occur from May to Jun and Sep to Nov in the short and long wet seasons, respectively (Turkalo & Klaus-Hugi 1999, Elkan 2003). As the calves are dropping in May and Sep at the beginning of the wet seasons, forage quality would be high (P. Elkan pers. obs.). Mountain Bongos calve in Jan and Sep (Kingdon 1982). Gestation periods of 282–287 days (Ralls et al. 1985) have been observed in captive Bongos. Oestrous cycle is 21–22 days and oestrus lasts three days (Ralls et al. 1985). Captive // conceived at 2.3 and 2.7 years of age and inter-birth intervals were 466 and 525 days. A single calf weighing an average of 20 kg is born (Forthman et al. 1993), although twins have been recorded in captivity (Schürer 1999, Ibler 2001). Females with new calves lie out for a period prior to joining groups (Kingdon 1982, Elkan 2003). Calves stay close to the mother during the early months and then form close associations with other young Bongos. In captivity, calves were able to eat vegetation during the first week (Forthman et al. 1993). Horns are visible after 2–3 months, growing rapidly to 100 mm after six months and 200 mm by about one year. Observations of young in captivity suggest that Bongo calves exhibit less aggressive behaviour and slower horn development than do Common Elands. The sex ratio estimates from individually recognized Bongos was 0.6 adult ?? to 1 / (Elkan 2003). Mortality and birth rates for Bongos are not known and are likely to be site-specific. Differences in adult to calf/juvenile and adult to subadult ratio suggest higher mortality in young, probably due in part to Leopard Panthera pardus predation (Turkalo & Klaus-Hugi 1999, Elkan 2003). Adult ?? had the highest mortality rate of any age/sex class during a 1997 die-off of Bongos in N Congo (Elkan et al. 2009). They live to 25 years in captivity, but are unlikely to attain that age in the wild. Predators, Parasites and Diseases  Young Mountain Bongos are vulnerable to African Rock Pythons Python sebae, Leopards and Spotted Hyaenas Crocuta crocuta during the lying up period (Verschuren 1975), as well as to predation from Lions Panthera leo 183

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(D. Western pers. comm.). In Central Africa, all ages, but especially young, are subject to predation by Leopards (Elkan 2003). Percival (1928) reported that rinderpest drastically reduced the populations of the Mountain Bongo in 1890 and 1896 and populations are thought to have suffered greatly in later epidemics in the early 1900s. Unusual cases of widespread mortality have been registered in both Lowland and Mountain Bongos. A case involving the mortality of a large number of Bongos was reported from the Mau Forest, Kenya in 1960 (Simon 1962). Local hunters indicated that foraging on poisonous second-year growth of an irregular Mimulopsis sp. cycle (after the extensive flowering and dieoff of bamboo Arundinaria alpina) caused extensive mortality not only in Bongos, but also in Forest Hogs Hylochoerus meinertzhageni and domestic cattle. Although symptoms of death did not resemble those of rinderpest, Davies (1993b) later argued that this artiodactyl disease was a more likely explanation for the deaths. Resolution of this controversy is of practical importance for the conservation of this species in Kenya. Theileria may also be an important disease affecting wild Kenyan populations (P. Reillo pers. comm.). Parasitological investigation of several Bongos in Congo found evidence of Elaeophora sagitta and nematode parasites (Huchzermeyer et al. 2001). In Apr and May 1997 a Stomoxys fly outbreak contributed to the mortality of large numbers of Bongos and other ungulates over a broad area of the tri-national region of Cameroon–Congo– Central African Republic. A prolonged severe dry season followed by exceptionally heavy rains created ideal conditions for several highly abundant generations of Stomoxys flies. Clouds of biting flies attacked large mammals in the region leading to mortality of Bongos, Sitatungas and Yellow-backed Duiker Cephalophus silvicultor. Mortality rates were highest in adult male Bongos. Elkan et al. (2009) hypothesized that the multifactorial mortality resulted largely from disruption of foraging patterns and the extreme fatigue of attempting to fight off and escape the biting flies. Conservation  IUCN Category: Near Threatened (T. e. isaaci – Critically Endangered C2a(i); T. e. eurycerus – Near Threatened). CITES: Appendix III (Ghana). Mountain Bongos are now rare on Mt Kenya and have been substantially reduced in Aberdares N. P. and its surrounding forest reserves. There is evidence of a remnant population in the Mau and Eburu forests, which are both subject to high levels of illegal logging and hunting. The Eburu population, located as it is in a small, isolated forest, is under especially intense pressure (L. Estes pers. comm.). The decline of Mountain Bongo populations in the Aberdares in recent years has been attributed to increased hunting by local people and habitat loss and transformation (Hillman 1986b, T. Butynski pers. comm.), and even to the increased numbers of Lions in the area (D. Western pers. comm.). Although these factors have surely played a role in the decline of Mountain Bongo, the impact of disease has probably been underestimated (L. Estes pers. comm.).The continued grazing of cattle in the forest reserves of Mt Kenya and the Aberdares may have greater implications for Bongo conservation than hunting pressure in terms of disease transmission; cattle have been found all the way up in Hagenia forest on the Aberdares plateau in the dry season – in an area where they certainly overlap with Bongos (L. Estes pers. comm.). The negative impact of forest succession as a result of total protection needs further investigation (Kingdon 1982)

and ongoing research at the time of going to press might shed some light on this issue (L. Estes pers. comm.). Two conservation initiatives are currently in progress on Mountain Bongos, including: (1) a multi-phased international programme to reintroduce captive Bongos from North America to Mt Kenya; and (2) the Bongo Surveillance Programme (BSP), which protects and investigates the status of the remaining wild Bongos in Kenya. The first phase of the Bongo reintroduction programme, spearheaded by the Rare Species Conservatory Foundation (RSCF), began in 2004, when 18 animals were flown from North American zoos to a captivebreeding facility at Mt Kenya Game Ranch, on the north-western slopes of the mountain. The second phase began in 2005, with the commencement of a research programme into the Mountain Bongo’s ecology. The research project, currently ongoing, is investigating the configuration of Bongo habitat on both the Aberdares and Mt Kenya, using recently collected field and remotely sensed data. Since 2004, the BSP has been surveying potential Bongo habitat, focusing primarily on the Aberdares, but also Mt Kenya and the Mau and Eburu forests; plans to survey the Cherangani Hills are also afoot. The BSP was created, and is run entirely, by Kenyans local to the Aberdares and Mt Kenya, and is largely responsible for the current state of knowledge about the remaining populations (L. Estes pers. comm.). Direct threats to Lowland Bongo conservation include snare hunting associated with expanding commercial forestry exploitation, high demand for Bongo trophies, and changing land use (Elkan 1995, 2003). Traditionally, due to taboos against eating of the meat, Bongos

Bongo Tragelaphus eurycerus.

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are not considered a preferred game species by local people in many parts of Africa, such as N Congo, S Sudan, SW Central African Republic and SE Cameroon (Hillman 1986b, Elkan 1995, 2003). However, there is still heavy loss due to snaring that indiscriminately kills young and wounds adults. Eroding traditional beliefs and expanding commercial hunting are creating new pressures on populations and its meat is sometimes smoked and sold as ‘buffalo’. Bongos are also the primary target of tourist safari hunting in the forests of central Africa. Demand for Bongo hunting safaris has been increasing greatly over the past decade and the use of dogs and inadequate regulation has resulted in over-hunting in several areas (Elkan 1995, 2003). H. Planton and A. DeGeorges (pers. comm.) reported that hunter success and trophy quality were declining in Cameroon. Hunters have argued that ‘trophy’ ?? taken are ‘solitaries’ and are too old and no longer involved in mating. While this may sometimes be the case, age estimation data indicate that many of these ?? observed as solitary are likely to be reproductively active (P. Elkan pers. obs.). Although the distribution and numbers of Lowland Bongos have declined over large parts of their former range, and particularly in West Africa, they remain patchily distributed, with localized concentrations in areas of favourable habitat (East 1999). Populations of Lowland Bongos in Central Africa receive protection in DzangaNdoki N. P. and Bangassou areas of Central African Republic, Lobeke N. P. (Cameroon) and in Nouabale-Ndoki and Odzala National Parks (Congo) (East 1999). In Gabon, they are reliably recorded from Mwagne, Minkebe and Ivindo National Parks (Maisels et al. 2004, P. Henschel pers. comm.). In Congo, the highest known Bongo densities occur in the Kabo and Pokola logging concessions (Elkan 2003). In West Africa, where declines have been more severe, the Lowland Bongo remains common in only a few areas, such as Taï N. P. (Côte d’Ivoire), Sapo N. P. (Liberia) and Kakum N. P. (Ghana) (East 1999). East (1999) estimated that perhaps 60% of Bongo numbers were confined to protected areas. Because the highest known abundances of Bongos in central Africa occur in logging concessions not protected areas, an approach is needed that incorporates both protected areas and logging concessions. The goal is to devise a strategy that reduces indiscriminate killing of Bongos by snares, strictly regulates and monitors safari hunting outside of reserves, and incorporates a landscape analysis of the effects of different land ownership, land use practices and resource needs of local people. This approach is being pioneered in Congo by establishment of community-supported subsistence hunting regulations in logging concessions that aim to ensure a sustainable supply of wildlife meat without illegal killing of Bongos and other protected species. Forest blocks are zoned as non-hunting or subsistence hunting areas. Separate hunting zones are designated for each village and logging camp. Transport and

commercial sale of meat across these zone boundaries is prohibited and regulations are enforced by a staff of both government officers and ecoguards (Elkan 2003). Rational allocation of hunting rights, promotion of alternative meat sources for logging company employees, and wildlife law enforcement are developed and combined with an intensive education programme that emphasizes both general conservation and the utilitarian practicality of nonhunting areas adjacent to hunting zones. Current Bongo distribution patterns are patchy and not well understood. It is important to begin developing and testing an ecological model of their distribution and conservation based on data on abundance and distribution in relation to forest cover and condition, critical forest clearing habitat and current and historical logging and hunting. The model can then be used to explore how the Bongo will respond to different logging and other anthropogenic factors. Measurements Tragelaphus eurycerus T. e. eurycerus HB (??): 2043 (1860–2190) mm, n = 12 HB (/): 2010 mm, n = 1 T (??): 561 (530–610) mm, n = 13 Sh. ht (??): 1210 (1100–1260) mm, n = 13 Sh. ht (/): 1220 mm, n = 1 Congo (P. Elkan pers. obs.) Record horn length is 89.2 cm for a pair of horns from the Kerre R., Central African Republic (Rowland Ward) T. e. isaaci HB: 1700–2500 mm T: 450–650 mm Sh. ht: 1100–1250 mm WT (??): 362 (335–400) kg, n = 3 WT (//): 229 (182–276) kg, n = 7 Kenya Body measurements: Haltenorth 1963 (mean and sample number not given) Weight: L. F. Bosely (pers. comm.) Record horn length is 87.9 cm for a pair of horns from Mt Kenya, Kenya (Rowland Ward) Key References  Elkan 1995, 2003; Hillman 1986b; Kingdon 1982; Klaus-Hugi et al. 1999; Ralls 1978; Turkalo & Klaus-Hugi 1999. Paul W. Elkan & James L. D. Smith

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Tragelaphus derbianus Giant Eland (Lord Derby’s Eland) Fr. Éland de Derby; Ger. Reisenelenantilope Tragelaphus derbianus (Gray, 1847). Ann. Mag. Nat. Hist. (1) 20: 286. Gambia, Western Africa.

Giant Eland Tragelaphus derbianus male. Foreleg/metacarpus proportions compared in Greater Kudu T. strepsicer­os (left), Giant Eland T. derbianus (centre) and Common Eland T. oryx (right). left:

above:

Taxonomy  First described by Dr J. E. Gray, keeper of the Department of Zoology at the British Museum in London from 1840 to 1874, from horns, then skins, sent from Senegambia (type specimen). The species was originally placed in the genus Boselaphus, then Oreas, and then Antilope. Taurotragus, referring to the size and shape of the largest antelope, rather massive and more or less oxlike, has been used since the late 1800s, but is here included in Tragelaphus (see genus profile). The species has been treated as conspecific with the Common Eland Tragelaphus oryx (Haltenorth 1963), but is considered a distinct species by most authorities. Four Giant Eland ‘races’ have been recognized in the past (Clark 1931): T. derbianus derbianus from Senegambia; T. d. cameroonensis from Cameroon; T. d. congolanus from Congo; and T. d. gigas, from Sudan. Only two subspecies are now recognized: T. d. derbianus is dedicated to the 13th Earl of Derby who employed the collector of the first horns sent to England, while T. d. gigas was described by von Heuglin from a pair of horns collected during travels to the White Nile region in 1863 (von Heuglin 1864). Synonyms: cameroonensis, colini, congolanus, derbii, gigas, typicus. Chromosome number: 2n = 31 in ? (translocatedY) and 32 in /, the same as the Common Eland T. Oryx (Nguyen et al. 2008). Description  The largest of all antelopes, massive, and often said to show a ‘bovine’ appearance (though more finely shaped) and sometimes exceeding the size of the African Buffalo Syncerus caffer. Long muzzle, but much narrower than that of an ox, with a black mark starting below the eyes and extending down to the nostrils (narrow on / and young, wide on old bulls); there is a white chevron between/below the eyes, except on old dark bulls. Ears broad and expanded, rufous outside, whitish with black patches on the hind margins. There is a mat of dark hair on the forehead of adult ??; a prominent dewlap commences below the chin. Coat hair short,

general colour ruddy fawn, sides (shoulder to rump) of body marked with usually 10–18 long vertical white stripes. A mostly black stripe runs along neck and back, with white hair on top of stripes. There is a short, darkish mane on the bull’s neck and withers; neck covered with rufous grey to blackish hairs, bordered at its rear end by a paler (??) or even white (//) collar. Dark marks are present behind the knees of the forelegs, and surrounding all four pasterns and fetlocks, with white spots often marking the front face of pasterns. Internal side of legs paler than body, or whitish. Hooves grey. Tail long, reaching hocks, with terminal black tuft 100–150 mm below the hocks, always moving (the most prominent detail when seen from a distance in dense bush). Females more lightly built and smaller, with no mat of hair on the forehead. Variation in size and shape of dark and white marks as well as horns permit individual identification. Females have two pairs inguinal nipples. Horns, present in both sexes, are thick-set, very large and massive, especially in ?? (smaller in //), diverging from the base and almost straight, tightly spiralled, with anterior ridge more pronounced than the posterior. Geographic Variation T. d. derbianus (Western Giant Eland): Senegal, Mali, Guinea and possibly E Guinea-Bissau. Bright rufous ground colour. T. d. gigas (Eastern Giant Eland): Cameroon, Central African Republic, Chad and Sudan. Somewhat larger (the name Giant Eland is most applicable to this form), with a sandy ground colour. The number of white stripes has often been noted as a distinctive feature between these taxa, with the western subspecies (derbianus) said to have more stripes (approximately 15) than the eastern (gigas; 12). However, individuals observed in Central Africa may have as many as 10–18 stripes (mean = 14) (H. Planton pers. obs.), and individuals of the western subspecies 12–16 stripes (mean = 14) (M. Antonínová & P. Hejcmanová pers. comm.). Similar Species Tragelaphus oryx. Allopatric species from open or lightly wooded savannas of eastern and southern Africa. General appearance

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the late 1990s (P. Chardonnet pers. comm.). They may still occur widely in SW Sudan (and indeed were recently recorded in Southern N. P.; Fay et al. 2007), and perhaps still occasionally visit NE DR Congo and NW Uganda (East 1999). Habitat  Giant Elands inhabit woodlands and forested Soudanian to Guinean savannas (1000–1500 mm rainfall in 5–7 months/year), and are never far from hilly/rocky landscapes or from water. They are frequently found in or near Isoberlinia doka woodlands (Kingdon 1997), although their range includes Terminalia–Combretum–Afzelia woodland, where Isoberlinia is not found (Bro-Jørgensen 1997). The western distribution corresponds to Combretum glutinosum–Annona senegalensis woodland and a mosaic of grass and woody savanna dominated by Combretaceae (Nežerková et al. 2004). Giant Elands move large distances daily (5–20 km/day) and all year round, and usually return to the same places during the same season. Wet season habitat is not clearly known, though it is widely believed that they spend the wettest months on rocky hills.

Tragelaphus derbianus

similar, though body slightly smaller, colour fawn or tawny, turning to greyish or bluish-grey with age, except for lower parts of legs; sides almost uniform or lightly striped (4–9 stripes); mane only on back of neck; dewlap commencing on throat; chevron often not visible between eyes; horns shorter, thinner; ears narrower, more pointed, without black marks. Distribution  Endemic to Africa.The historical range of the Giant Eland is not well known, but they probably occurred throughout the relatively narrow belt of savanna woodland extending across West and central Africa from Senegal to the Nile (East 1999). The distribution of the Western Giant Eland is centred on and around SE Senegal, with the Eastern Giant Eland occurring in the central African region. The Western Giant Eland has been formerly reported from Senegal to Togo (Bigourdan & Prunier 1937, Baudenon 1952, East 1990), though its occurrence in Togo might have been a mistaken confusion with the Bongo Tragelaphus eurycerus (Grubb et al. 1998). The distribution has been reduced since the 1950s to small, scattered populations (Jeannin 1951). The subspecies still occurs in SE Senegal (Chardonnet 1997, G. Mauvais pers. comm.), the far north of Guinea (Sournia et al. 1990, Chardonnet 1997, Darroze 2004), probably SW Mali (Chardonnet 1997, Darroze 2004) and possibly E Guinea-Bissau (Chardonnet & Limoges 1990, East 1999). The Eastern Giant Eland was formerly distributed from NE Nigeria to NW Uganda; it is now apparently extinct in both of the aforementioned countries, surviving in the former in the northern part of Gashaka-Gumti N. P. until the 1970s (Anadu & Green 1990), and being exterminated in the Uganda part of its range during military operations in 1970 (Kingdon 1982).The species now occurs mainly in NE Central African Republic, and in N Cameroon, with herds crossing the Chad border to the east; occasional vagrants may enter Nigerian territory (East 1999). There has been some recolonization by Giant Elands of hunting blocks of SE Chad since

Abundance  Densities in the order of 0.15–0.30/km² have been recorded, with great variations as herds in Cameroon move a lot within home-ranges of some 1000–1500 km² (H. Planton pers. obs.); they can be locally abundant during a few weeks/months and then totally absent for months. Thus, the same herd can be observed in very distant places at different times and densities are easily overestimated. A single study in Central African Republic reports an average home-range of 223 km² (Graziani & d’Alessio 2004). Although no specific count has been carried out, it is estimated that some 200 or less individuals of the Western Giant Eland survive in Senegal (G. Mauvais pers. comm.). According to East (1999), there are probably more than 15,000 Eastern Giant Elands remaining, with over 12,500 in the Central African Republic. Since the early 1990s, numbers have tended to increase, at least for Cameroon (Planton et al. 1995), Chad and the Central African Republic (Blom et al. 1995). There is no information on numbers surviving in Sudan, although Fay et al. (2007) estimated a population of 165 for Southern N. P. Adaptations  Giant Elands drink daily when water is available, but even youngsters will trot or walk tirelessly all day long if disturbed, without stopping at any water point. They browse day and night, and do not rest except, when undisturbed, during the heat of the day. They use their strong horns to knock down or break branches and gain access to leaves of trees. Giant Elands have an excellent sense of smell and hearing, as well as excellent sight even at long distances. The visual impact of the black-and-white markings on legs and ears is greater than in the Common Eland. This difference could be related to their less open habitat, to a greater role for ear and foot signals or to carry-overs from an ancestral condition. Ears are substantially larger than in the Common Eland and this, too, could be due to evolutionary inertia, but is more likely to represent selective pressure to maximize acuity of hearing in wooded environments. Prior to a well-marked breeding season there are colour (and, perhaps, physiological and olfactory) changes that are exclusive to the foreheads and necks of breeding bulls. This overall darkening of the bulls’ forequarters might signify a visuo-olfactory advertisement of the males’ reproductive condition. A dense brush of hair on the 187

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forehead of adult ?? is often employed in ‘marking’ behaviour. Supposed territories are marked by adult bulls urinating on a patch of moist soil, and they then rub their forehead and horns to pick up the odour. Bulls anoint themselves on their sides and back and thereby more effectively proclaim their presence. The thick mat of woolly dark hair on the forehead acts as a paintbrush and allows the bull to leave samples of his scent on tree trunks at just the right level to be noticed by other elands (Ruggiero 1990). Foraging and Food  Giant Elands feed mostly on leaves, shoots, herbs and fruits (but occasionally on grasses), and are classified as browsers by Gagnon & Chew (2000) in their review of the dietary preferences of African bovids. Movements are to a certain extent determined by the presence of certain trees or shrubs, the young leaves of which form their favourite diet. Feeding habits have been observed in several places, but systematically studied only in N Cameroon (Bro-Jørgensen 1997, Michaux 1998), where more than 80 plant species were eaten (the ingestion of 16 of these having been directly observed). Shortly after bushfires, a lot of dried leaves and dry fruits, and ashes, are eaten. During the driest season (Jan–Mar), they feed mainly on Terminalia spp. and Combretum spp. leaves, which remain important until the rains (Jun). Where Adenodolichos sp. occurs, herds will actively look for sweet young shoots of this plant after fires and until the rains. Isoberlinia spp. leaves are eaten during the dry season, but never exceed 20% of the diet. When rains fall, Giant Elands start looking for Tephrosia spp. and Acacia spp. leaves and pods. Legumes form by far the major part of their diet during the wet season (40– 85%). Grasses can be eaten all year round, but never exceed 4–5% of the diet. Salt-licks are visited every day when possible. Social and Reproductive Behaviour  Herds comprise // and young of both sexes, plus young adult ?? and sometimes adult breeding bulls. The latter regularly wander alone or in small male groups outside breeding herds. Herd size varies a lot according to season. Most often herds include about 20–30 individuals; large herds of 150 individuals and more are observed early in the dry season (Dec and Jan: Cameroon), before the mating season, and stay together almost until the rains (Jun). In such herds, an adult / often calls continuously, which has the effect of keeping youngsters together near her (H. Planton pers. obs.). Before and during the wet season, herds tend to split into smaller groups of less than ten, often a few // and one ? plus young. The same herd can be observed in various places of the home-range over which elands move continuously. They are locally abundant for a few weeks/month and then move to another part of the range. Relations between individuals within the group consist mainly of smelling or licking or rubbing nose, mouth, forehead, horns, ears, belly, tail and anogenital region (Altmann & Scheel 1976). Close relationship is sometimes observed between two specific individuals for a couple of days, during which they often rest their heads on each other’s back. Males, mainly young, seem to establish their status in the group’s hierarchy by simulating threats and fights, often kneeling, near muddy places or salt-licks. The dominant action is to charge the other, head down, and/or snort noisily; the weaker usually gives up and walks away. However, serious fights are sometimes observed or evidenced by scars or broken horns. These often occur early in the wet season, but also during the mating season. Fighting ?? engage

Giant Eland Tragelaphus derbianus head in profile.

their horns and wrench their heads sideways in an effort to twist each other’s neck and unbalance their opponent. Marking behaviour by adult ?? has been mentioned earlier. Before mating, the bull approaches the / with neck and head stretched out. The / jumps aside then pushes the ? away. The ? tirelessly tries again to approach, turns around her, smells her, rests his head on her rump, until she is ready to mate. Pregnant // tend to stay together with young individuals, while breeding bulls wander in search of other // on heat.Young often stay together and playfully chase each other among and around the adults (Ruggiero 1990). Females accompany their offspring for about one year. Giant Elands have often been said to be silent. Actually, when undisturbed and browsing, they often sniff and growl discreetly; the / keeps contact with her youngster by a high-pitched call, to which the latter answers. Though they can often be close to other species, Giant Elands very seldom mix with them. Like Common Elands, they are easily tamed when captured as young, and can behave then like domestic stock. Reproduction and Population Structure  In Cameroon, mating mainly takes place in the first half of the dry season (Dec–Feb). Prior to this, changes in the breeding bull’s forehead and neck have been mentioned and are quite noticeable in the field (H. Planton & I. Michaux pers. obs.). Mean gestation length is 39 weeks. Calving has yet to be observed in the wild, but occurs between Sep and Jan, i.e. before (peak) and shortly after the last rains. Under captive conditions in Senegal, calving takes place between Nov and Feb, in the early dry season (Al-Ogoumrabe 2002, M. Antonínová & P. Hejcmanová, pers. comm.). Usually a / gives birth to a single calf, about 25–35 kg, which stays hidden and out of the herd for one week. The calf starts browsing from one week old, and is weaned after six months. Maturity is reached between 15 and 24 months, and animals are full size by 4–5 years. Bulls do not usually succeed in mating before they are seven

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Giant Eland Tragelaphus derbianus subadult male.

years old. Sex ratio among calves is 1 : 1, but adult // outnumber adult ?? (Altmann & Scheel 1976, H. Planton pers. obs.). Predators, Parasites and Diseases  Little information is available regarding predators, which include Lions Panthera leo and African Wild Dogs Lycaon pictus, while Leopards Panthera pardus, Spotted Hyaenas Crocuta crocuta and Cheetahs Acinonyx jubatus also prey on calves, which // try to protect. Major epizootic diseases cause great variations in numbers. In particular, Giant Elands have suffered heavy mortality from rinderpest, e.g. in the early 1900s (Blancou 1960), to which it is said to be more susceptible than any other antelope. Its demise in Gambia has been attributed primarily to the devastating effects of this disease (Camara 1990). Populations in the central African region crashed by 60–80% during and after the 1983–84 rinderpest outbreak, but have almost recovered now facilitated by the bush encroachment that accompanied the severe reduction in elephant numbers by poachers during the 1980s and the prevalence of uncontrolled fires (East 1999). Studies on parasites have been carried out in Cameroon through helminthological necropsies and year-round faeces collection/analysis

(Michaux 1998). Cestodes, trematodes and nematodes have been found, but in low numbers compared with neighbouring wild grazers (e.g. Kobs Kobus kob) and in very low numbers and species diversity compared with domestic stock. Parasitic load was almost nil in the dry season.The Giant Eland is a new host for five parasite species (Moniezia monardi, Haemonchus vegliai, Cooperia yoshidaï, Cooperia sp. and Ostertagia angusdunni), and two other species found might represent undescribed new species of Cooperia (Michaux 1998). Conservation  IUCN Category: Least Concern (T. d. derbianus – Critically Endangered C2a(ii); T. d. gigas – Least Concern). CITES: Not listed. Uncontrolled hunting (notably assisted by fire) and loss of habitat have contributed greatly to the reduction of Giant Eland numbers, particularly in West Africa where they have been exterminated throughout most of their former range. Traditionally, the Fulani people did not hunt them, as they are believed to transmit diseases and cast spells. Their current distribution range includes a number of sizeable protected areas, including Niokolo-Koba N. P. and Falémé Hunting Zone in Senegal (derbianus); Faro, Bénoué and Bouba Ndjida 189

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National Parks and most of the 27 surrounding hunting concessions in N Cameroon, Bamingui–Bangoran and Manovo–Gounda–St Floris National Parks and most of the hunting blocks in N and NE Central African Republic, and Southern N. P. in Sudan (gigas). Further development and maintenance of sustainable trophy hunting and improved protection and management of national parks within its range states will be essential to ensure this antelope’s long-term survival (East 1999). Attempts to capture Giant Elands and establish herds of both subspecies outside the species’ natural distribution area have been made on at least ten occasions for purely commercial purposes. The Eastern Giant Eland is held in captivity in two countries outside their distribution range: South Africa and the USA. Historical captures were undertaken in Sudan (1938–46), Cameroon (1958) and Chad (1967–70). Of an unknown number of Giant Elands handled, 31 were delivered to six European or American zoos, none of which has any representation in the current captive populations. More wild animals were caught in 1985 in the Central African Republic. The survivors (3 ??, 5 //) were the initial founders of the current North American captive population (Romo 2001). Animals in captivity derive from the population established in the USA, initially at Cincinnati and Los Angeles zoos, from nine wild-caught animals imported from the Central African Republic in 1986. As of 23 December 2011, the North American population stood at 49 individuals (20 ??, 29 //) in five institutions: San Francisco Zoological Gardens, White Oak Conservation Center, San Diego Zoo Safari Park, Miami Metrozoo and Houston Zoo.This suggests a 30% decline compared to the number of 69 reported in March 2004 (ISIS records). In mid-2000, a private South African capture team caught 23 Giant Elands in Central African Republic, of which two died and 21 were flown to Togo. IUCN’s Antelope Specialist Group has no information on the fate of the ten captive Giant Elands that are believed to have remained in Togo (East 2001), but 11 of the 21 were later shipped to John Hume’s Mauricedale Game Ranch (Mpumalanga, South Africa), with a three-month temporary import permit. Four more died there, and the seven survivors were moved to Johannesburg Zoo in December of the same year following an outbreak of footand-mouth disease (East 2001, Estes 2003), shortly after which another one died. The two remaining ?? died during blood transfusion procedures, and since then there have been additional age-related deaths (D. Moss pers. comm.). The survivors, two // only, are held at Johannesburg Zoological Gardens. No births have been reported in RSA since October 2001 (ISIS records). No Western Giant Elands are known to have been held in captivity until recently. Captures made by San Diego Zoo Safari Park in Falémé Hunting Zone, Senegal, in 1979, resulted in all nine animals dying during or shortly after capture (S. Diouf pers. comm.). The next captures were conducted in Niokolo-Koba N. P., Senegal, in 2000

by a South African team for the Ministère de l’Environnement et de la Protection de la Nature and private Société pour la Protection de l’Environnement et de la Faune au Sénégal. Some 126 animals were removed alive from the park, including nine Giant Elands (eight // and one ?). Some of these animals were shipped to South Africa, but none of the elands. Three of the elands died in quarantine, the six survivors (one ?, five //) were kept in a 25 ha enclosure (enlarged to 50 ha in 2004) of unsuitable habitat in Bandia G. R., Senegal, just next to exotic (South African) species including Common Elands, with which they could potentially have hybridized (East 2000).There were four births at Bandia in 2002 (Nežerková et al. 2004), and 30 in total between 2000 and 2006 (M. Antonínová & P. Hejcmanová pers. comm.). A second enclosure was built in Fathala Reserve, to which a male-only group (9) and a breeding nucleus (1, 3) were translocated in mid-2006 (Antonínová et al. 2006). In 2009, the semi-captive population comprised 54 individuals (26 ??, 28 //), with an annual population growth of 1.4% (Koláčková et al. 2011). Further translocation of Giant Elands to areas such as private hunting reserves outside the species’ natural range could potentially have serious consequences in terms of spreading diseases and destroying trophy hunting activities in West and central Africa, which are currently the main support and reason for wildlife conservation in these regions. Measurements Tragelaphus derbianus T. d. gigas HB: 2620–2930 mm, n = 30 T: 600–700 mm, n = 30 HF c.u.: 100–110 mm, n = 30 E: 230–280 mm, n = 30 Sh. ht: 1650–1780 m, n = 30 N Cameroon (H. Planton & I. Michaux pers. obs.) Depierre & Vivien (1992) recorded a maximum HB of 3200 mm Average estimated weight is around 450 kg for // and 900 kg for ?? (H. Planton pers. obs.); Haltenorth & Diller (1980) recorded a mass of 300 kg for // and 1000 kg for ?? Record horn length for T. d. gigas is 142.9 cm for a pair of horns from Ouandjia R., Central African Republic, and for T. d. derbianus the record is 115.6 cm for a pair of horns picked up from an undefined locality (Rowland Ward) Key References  Akakpo et al. 2004; Altmann & Scheel 1976; East 1999; Jeannin 1951; Michaux 1998; Nežerková et al. 2004; Ruggiero 1990. Hubert P. Planton & Isabelle G. Michaux

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Tragelaphus oryx Common Eland Fr. Éland de cap; Ger. Elanantilope Tragelaphus oryx (Pallas, 1766). Misc. Zool. p. 9. Known to the Dutch ‘ad Promontorium B. Spei’, restricted to South Africa, Western Cape Prov., near Cape Town by Shortridge (1934: 607).

Common Eland Tragelaphus oryx male.

Common Eland Tragelaphus oryx female.

Taxonomy  The Common Eland Tragelaphus oryx and Giant Eland Tragelaphus derbianus have been considered conspecific by some authors (e.g. Haltenorth 1963), but are usually treated as full species. Three subspecies are generally recognized (Ansell 1972, Kingdon 1997), although their validity requires confirmation. Lorenzen et al. (2010) recorded a significant regional divide between mtDNA lineages in Common Eland in East and southern Africa, and posited a more recent origin of the East African population that could result from colonization following extinction from the region. Synonyms: alces, barbatus, billingae, canna, kaufmanni, livingstonei, livingstonii, niediecki, oreas, pattersonianus, selousi, triangularis, typicus. Chromosome number: 2n = 31 in ?, and 2n = 32 in /, the difference being due to a Y-autosome translocation; the X chromosome is large and acrocentric (Wurster & Benirschke 1968, Wurster 1972, Buckland & Evans 1978, Robinson et al. 1997). There are two well-documented records of hybrid male offspring between a Common Eland and Greater Kudu Tragelaphus strepsiceros, one known to be sterile, the other unknown (Jorge et al. 1976, Van Gelder 1977a).

forehead has a strong smell due to a secretion from a glandular area in the skin (and frequent rubbing in urine). Females are smaller than ??, with the dewlap much smaller and thicker, a terminal tuft of black hair, and no facial hair tufts. Females have two pairs of inguinal nipples. Horns, present in both sexes, are nearly straight, slightly diverging, with a heavy spiral ridge at the base. Horns of oldest ?? are shorter than those of younger animals. The horns of adult // are less heavy and thinner in diameter and more uneven, and without the heavy spiral ridge at the base. Age determination based on horn growth and teeth eruption is discussed by several authors (Kerr & Roth 1970, Attwell & Jeffery 1981, Jeffery & Hanks 1981). Horn buds are present at birth, and the horns grow rapidly until about seven months of age, and thereafter more slowly. At about 18 months of age, the horns of ?? clearly show signs of the developing spiral ridge at the horn base. In the dentition, molars 1, 2 and 3 erupt at 6–8, 14–20 and 24–27 months, respectively, and permanent premolars erupt at 36–37 months; full adult dentition is attained in the fourth year of life.

Description  Common Elands are large, cow-like bovines, with straight spiral horns and a dewlap. The pelage is tawny with short hairs, becoming blue-grey with age, especially in ??. There are variable white stripes on the back and flanks, which usually become less distinct with age.The pelage is whitish on the belly, inside the legs and ears and above the hooves. A dark mark is present above the knee on the back of the forelegs. Dark hair on ridge along back and on tufted tail tip. The dewlap is long, thin, pendulous, with a terminal tuft that is lost in older ??. Neck thick and muscular, darker than body; there is no mane, but pale curly hairs on the upper surface. Tufts of hair on the nose and forehead vary in size and colour between individuals and over time within individuals. The mat of hair on the

Geographic Variation T. o. oryx: Southern Africa, with its range extending north to S Botswana and N Namibia. T. o. livingstonii: East-central African woodland areas. T. o. pattersonianus: Tanzania northwards. Common Elands in the southern part of the range (T. o. oryx) are dull fawn in colour, the white stripes on the body not particularly distinct, and having a dark brown mark on the back upper region of the forelegs. Common Elands further north have distinct body stripes, but a less distinct dark mark on the back of the forelegs; body colour tends to be richer in Common Elands from northern areas. However, there is a large degree of intergradation and also considerable variation within 191

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Lateral, palatal and dorsal views of skull of Common Eland Tragelaphus oryx.

Tragelaphus oryx

regions. For example, animals in Zimbabwe show traits characteristic of both T. o. oryx and T. o. livingstonii. Likewise, Common Elands from Angola have been referred to the subspecies T. o. livingstonii, although Ansell (1972) noted that Angolan Common Elands occur in an area of intergradation between the nominate form and livingstonii. Crawford-Cabral & Veríssimo (2005) note that Angolan animals are more comparable to the nominate form, and thought that at least animals from the west of the country were representative of T. o. oryx. Similar Species Tragelaphus derbianus. Savanna of West Africa and central Africa in N Cameroon, Central African Republic to SW Sudan (west of the Nile R.). Larger, with longer horns; horns in ?? are longer, more widely splayed, and have a looser spiral; neck of ? darker, with dewlap hanging from jaws and neck only, rather than continuing between legs; ears broader, and more conspicuously marked, as are hocks. Distribution  Endemic to Africa, being widespread in savanna and woodland areas in eastern and southern Africa from SE Sudan and SW Ethiopia southwards to South Africa. Historical Distribution From the Cape Peninsula (South Africa) to forest margins in the Congo Forest basin, the Nile flood-plain and arid N Kenya. Interestingly, a fossil specimen attributed to the Common Eland (and not to the older, now extinct T. arkelli) has been found at Ternifine in Algeria (Arambourg 1962). Current Distribution Still present in much of its former range, except in Burundi, where the species is considered extinct, and Uganda and Rwanda, where it is close to extirpation (East 1999). In Sudan, Common Elands are confined to the south-east, where they still occur in Boma N. P. (Fay et al. 2007); in Ethiopia they are known only from the Omo region (Hillman 1988a, East 1999). In Angola, where it was formerly widespread, the species is also believed to be

close to extinction, although there is no recent information on its status across much of the country. In South Africa, natural surviving populations are confined to the Northern Cape near Botswana (and where populations move into the area from S Botswana), parts of the North West and Limpopo provinces, and in the KwaZulu–Natal Drakensberg mountains (East 1999). In Lesotho, Common Elands occur only in Sehlabathebe N. P. (Lynch 1994). Within the rest of its range, it has been increasingly restricted to protected areas and regions with low human populations. Common Elands have been widely reintroduced in southern Africa (e.g. to Zimbabwe, many parts of South Africa, and Swaziland). Habitat  Common Elands occupy a wide variety of habitats, from semi-desert scrub in the Kalahari, to open Brachystegia woodland, to alpine moorlands on the slopes of Mt Kenya, Mt Kilimanjaro, the Nyika Plateau and the KwaZulu–Natal Drakensberg. In the KwaZulu–Natal Drakensberg, they occur up to 2400–2700 m, favouring slopes of 20°  or less (Rowe-Rowe 1983, 1994), while in East Africa they have been recorded to 4750–4890 m on Mt Kilimanjaro (Grimshaw et al. 1995; and see Kingdon 1982). They are not found in deep forest, in true deserts, or in completely open grassland, though they do occur in grassland with good herb cover or which is interspersed with browsing habitat along drainage lines. Although they no longer occur in coastal regions, they do still occur at relatively low altitudes throughout their range. Abundance  Estimated to number more than 130,000 in the wild (correcting for undercounting biases in aerial surveys), with local density estimates (based on aerial counts) ranging from 0.05 to 0.4/ km2 (e.g. Laikipia Ranchlands in Kenya, Selous in Tanzania, Etosha N. P. in Namibia and Kgalagadi Transfrontier Park). Higher density estimates (0.6–1.0/km2) have been obtained in areas such as Omo N. P. in Ethiopia and Nyika N. P. in Malawi; ground surveys reveal similar estimates in areas such as Lake Nakuru N. P. (Kenya) and De Hoop N. R. (South Africa) (East 1999). The largest populations are in Namibia (ca. 32,000, where the majority are on private land), Tanzania (ca. 24,000), Zimbabwe (ca. 14,000), Botswana (ca. 14,000) and Kenya (ca. 13,000).

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Common Eland Tragelaphus oryx adult male myology.

Common Eland Tragelaphus oryx skeleton

Adaptations  Common Elands are unusual among browsing bovines since they move long distances in search of ephemeral food sources. They have other attributes that are more typical of grazers such as an open social system. This life-style allows Common Elands to exploit a resource that is not available to other browsers – bushy vegetation and herbs sparsely scattered in grassland. In the Kalahari, the only other large browsers (Greater Kudus) are confined to patches of thicker vegetation, while Common Elands move freely through the system, and make use of dune areas, where the only bushy vegetation consists of dwarf shrubs. Adaptations to a mobile life include the ability to go without water for prolonged periods. While they will drink when water is available, they are also able to obtain sufficient moisture from their food. Common Elands followed on a daily basis using satellite technology in the southwestern Kalahari in Botswana never went close to a water source throughout the entire dry season (C. Thouless pers. obs.). Common Elands have a number of physiological adaptations that allow them to survive without drinking.Their body temperature can vary substantially. Under experimental conditions, body temperature increased from 33.9 to 41.2 °C while ambient temperature was kept at 40 °C, although body temperature generally remained below ambient temperature once it had exceeded 40 °C. The excess heat accumulated during the day is dissipated at night through conduction and radiation, and in order to reduce water loss at night Common Elands breathe more slowly and deeply (Taylor 1969). Recent studies have shown that selective brain cooling does not occur in Common Elands, and brain temperature remains constant at around 0.4 °C above carotid blood temperature (Fuller et al. 1999). Common Elands have a very high metabolic rate considering their size. Compared with Hereford cattle, their metabolism was 30% higher within the thermoneutral zone, and they excreted a far greater amount of urea in their urine (Taylor & Lyman 1967). Activity patterns in Common Eland tend to vary depending on environmental factors and food availability (see, for example, Lewis 1978). In hot climes, such as Tsavo N. P., they may rest up in the shade all day feeding at night (Hillman 1979). On Loskop Dam N. R. in South Africa, Common Elands exhibited four main periods of activity during the day in winter, while in summer they moved and

fed in the morning, rested around mid-day, and fed again intensively in the late afternoon (Underwood 1975). Although Common Elands, with their considerable bulk, usually move deliberately, they are capable of moving at great speed. When disturbed, they gallop, and jump high into the air, sometimes leaping over the backs of other animals.The gallop generally develops into a fast trot, which can be sustained for a considerable time.They are remarkably good at jumping, and can easily clear a two-metre-high obstacle. There are a number of displays carried out by Common Eland ??, involving the horns and the frontal brush of hair.These include vigorously rubbing the face in mud, particularly mud pats created by urine – either their own or from another animal. Adult ?? also thrash vegetation with their horns, and break branches, and also rub their frontal brushes on tree stumps and broken branches. The effect of these behaviours is to change the appearance of the animal, with mud and/or vegetation being attached to the frontal brush and horns, and to accentuate the animal’s smell. They appear to prefer ‘smelly’ substances for rubbing, such as aromatic shrubs and wet ash after grass fires (Hillman 1979). Foraging and Food  Common Elands are primarily browsers, though the degree to which grass forms an important component of the diet is debated. They have been classed as intermediate mixed feeders (Hofmann & Stewart 1972, Hofmann 1973), and Gagnon & Chew (2000), in their review of the diets of African bovids, considered them as browser–grazer intermediates taking as much as 50% grass. Analysis of rumen contents of shot animals in Kenya indicated that they were feeding on up to 60% grass in the wet season, but less than 10% during the dry season (Hillman 1979). Other studies (Lamprey 1963, Underwood 1975) have also indicated high proportions of grass in the diet, although Watson & Owen-Smith (2000) have highlighted the unreliability of these studies. Grass appears to be eaten in quantity only during the wet, summer months, and most stomachs examined from other times of the year contained only a small percentage of grass (Wilson 1969, Kerr et al. 1970, Scotcher 1982, Buys 1990, Watson & Owen-Smith 2000). In the study by Watson & Owen-Smith (2000), grass formed 45% of the diet in early Dec, but only 6–10% of the diet in other months of the early wet season. Studies involving 193

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stable carbon isotope analyses of animals in both East Africa (including the Athi Plains region) and southern Africa indicate that although Common Elands may consume some grass, they have a significant component of browse in their diet and that few populations consume as much as 50% grass (Cerling et al. 2003, Sponheimer et al. 2003b; and see Wallington et al. 2007). Diet usage has been associated with habitat use: Common Elands show a preference for bush habitat for the dry part of the year and for grassland in wet periods, although ?? tend to use thicker bush areas than // (Buys 1990, Fabricius & Mentis 1990). Common Elands introduced to Mountain Zebra N. P. in the Eastern Cape of South Africa showed similar habitat preferences, but habitat selection was unrelated to the availability of palatable grasses in the habitat (Watson & Owen-Smith 2000). Common Elands in Mountain Zebra N. P. have a diet comprising as much as 94% browse (Watson & Owen-Smith 2000), and have fared very well since their introduction. Watson & Owen-Smith (2002) have suggested this is because Common Elands have been able to select a diet sufficiently low in fibre content by eating young shoots of most woody species encountered and by consuming large proportions of palatable woody species with low fibre content.

In the wet season in Shinyanga, Tanzania, Common Elands were observed to eat herbs, particularly Ipomaea spp. and Commellina africana (Harrison 1936). During the dry winter months in southern Africa, they also eat dry fallen leaves. They also are known to eat fruits, such as those of Sclerocarya caffra and Ximenia caffra, and in the dry season have been known to eat the fleshy leaves of aloes (Harrison 1936, Wilson 1969). Feeding takes place from ground level up to about 2 m. They use their lips, rather than tongue, to grasp food. Common Elands may strip the leaves off twigs by drawing them through their lips. The use of horns to break down high branches has been reported from some populations. Common Elands insert their horns on either side of a branch, and then twist until it breaks off or hangs down, when it can be reached (Skinner & Chimimba 2005).This behaviour was not observed in the Kitengela in Kenya or the Nyika Plateau in Malawi (Hillman 1979), but was seen in Common Elands confined in an enclosure with Camphor Bush Tarchonanthus camphoratus and in a wild population in E Zambia (Wilson 1969).

Common Eland Tragelaphus oryx.

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Social and Reproductive Behaviour  Common Elands have a very fluid social structure, and there appear to be no stable long-term relationships between individuals. The strong bonds between calves, and the ecological differences between ?? and //, have resulted in an unusual social structure. Only one study has been carried out on the social organization of a large, naturally occurring, free-ranging Common Eland population – in the Kitengela Plains of Kenya (Hillman 1979, 1987); some aspects of this study differed from those of smaller enclosed populations such as that studied by Underwood (1973, 1975). Group sizes are very variable, both between and within populations. In many areas the groups are usually small, but in some places and times groups in excess of 500 individuals may form. These have been reported from Hwange N. P., Zimbabwe (Wilson 1975), and from the south-western Kalahari, Botswana (C. Thouless pers. obs.). Groups show a high degree of flux, and adult Common Elands appear to move freely between groups. Association between individual animals is slight, rarely lasting more than a few days (Hillman 1987). Common Elands are very seldom seen alone, and solitary animals are almost always the older ‘grey’ ??.The most frequently observed herds are small groups of adult ??, groups of // (typically with less than ten individuals), or slightly larger mixed adult groups (usually of 10–20 individuals). However, groups containing juveniles in addition to adults are much larger, and those with calves larger still. Thus, these groups often include a large proportion of the total population in an area even though they are not the most frequently observed groups (Hillman 1979). Male Common Elands appear to be more typical tragelaphines than the //. Males are most often seen in smaller, adult-only groups, particularly in all-male groups. Adult // are mainly in calf-containing groups, and to a lesser extent in mixed all-adult groups. As a subadult animal matures it spends more time away from mixed groups, and with the small adult groups. Females then return to the nursery groups as they begin to calve, while ?? continue to gravitate towards the very small all-male groups. These differences between the sexes are affected by their different feeding and ranging behaviour (see later). Within large groups, young Common Elands are more closely associated with each other than with their mothers, and it is believed that this ‘within-age-group’ bond is the main reason why these large groups form. Most animals within a group are well spread out, with the exception of calves, which tend to form a tight ‘creche’. Nursery groups are larger when environmental conditions are good, increasing to several hundred animals in grassland areas in times of high rainfall and good vegetation condition. This may be an antipredation measure only possible during times of good food supply. Common Elands in the KwaZulu–Natal Drakensberg display an unusual social organization, whereby they form large, mixed herds of up to 200 animals during summer (Dec/Jan), during which time breeding // are mated by dominant ??. From autumn (Mar) the large herds break up and they disperse widely in small groups of 4–10 animals, which may be of varying combinations of sexes and age classes. Shortly after the calves are born (Sep and Oct), the small groups start to join to form larger herds again (Scotcher 1982, Rowe-Rowe 1994). There is no evidence of territoriality in Common Elands (Underwood 1973, 1975, Hillman 1979), and no evidence that faeces are used for marking. Urine and secretions from the frontal tuft may be used in some kind of marking. However, there is a dominance hierarchy within

Common Eland Tragelaphus oryx development of male colouring.

a population. A range of threats is used between individuals, including a threatening look, wiping the horns across the animal’s own back, pointing the horns forward, and lunging with the horns (Hillman 1979). These threats usually result in the other animal moving away, and seldom escalate into more serious conflict. Sparring between the younger classes of ?? is common. It involves the deliberate lowering of the heads, and tangling of the horns, followed by a pushing contest; this is often followed by the subordinate ? mounting its sparring partner or another nearby animal (Hillman 1979). Fighting is much less common, and results only when two old ‘grey’ ?? are competing over // in a group. Instead of deliberate tangling of the horns at close quarters, this involves charging together from a distance of 1–2 m, followed by pushing and manoeuvring to throw the other animal off balance.This can result in serious injuries.Apart from displays involving the males’ frontal brush (described above under Adaptations), displays are in general not particularly ‘ritualized’ and are not unique to the Common Eland (Kiley-Worthington 1978). Self-grooming is carried out regularly and involves scratching their bodies and faces on trees and other objects (or the front part of the body with a hindleg), and grooming with lips, teeth and tongues. Mutual grooming usually occurs between standing animals while chewing the cud, and involves the head and neck – parts of the body that cannot be reached by the animal itself. Mutual grooming only occurs between young or adult //. Common Elands have a reputation as great wanderers; in some places this is thought to take the form of regular migrations, while in others there seem to be irregular movements, and some populations are believed to be more or less sedentary (such as those in SE Zimbabwe). The movements of free-ranging animals have only been studied in detail in the Athi-Kapiti area of Kenya, and in the south-west Kalahari of Botswana. Many populations are now confined within fenced areas, restricting what natural tendency there may be to move further. Hillman (1988b) studied the movements of Common Elands by repeated sightings of individually recognized individuals. Adult ?? had home-ranges of approximately 30 km2 while adult // and juveniles had home-ranges of approximately 200 km2. For both sexes most of these areas were used in the wet season; dry season ranges were considerably smaller. Male home-ranges were centred on areas of bush along gorges and drainage lines, or on forest edges. Females 195

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Common Eland Tragelaphus oryx serial outlines of leap (from film).

and juveniles were found in similar areas in dry periods, but used areas of open grassland for longer periods in the wet seasons. Common Eland movements have been studied in the south-western Kalahari, where they are largely confined to Kgalagadi Transfrontier Park, using radio-telemetry and satellite technology over a number of years (R. Brett, M. Knight, S. Makhabu, N. Nagafela & C.Thouless unpubl.). Aerial surveys in the park show that there is a tendency for the main concentrations of Common Elands to be in the bushy northcentral part of the park in the wet season, with a surprising shift towards the drier sand dune areas of the south-west during the dry season (Verlinden 1998 ). In some dry years, such as 1985, this movement is particularly pronounced, and very large numbers of Common Elands may congregate in the south-west, including the fossil valley of the Nossob. Since animals fitted with collars were known never to go close to artificial watering holes, there must be some presently unknown ecological reason (and not a need for water) for this movement down the rainfall gradient in the dry season. Individual Common Elands in this area have very large home-ranges, averaging 8449 km2 for ten // tracked for at least one year, with a range of 1691–19,761 km2 (minimum convex polygon). Unlike other species, which tend to use a particular area for some time before moving on, Common Elands moved considerable distances on a daily basis. Typically they would be about 20 km distant from where they were 24 hours previously, and straight-line movements of up to 55 km were recorded over a 24-hour period. This pattern of movement was not dramatically affected by the onset of the rains. However, they do seek areas that have ‘greened up’ in response to the rains (R. Brett pers. comm.). In addition to migratory movements, Common Elands also make vertical movements in relation to food availability. In the KwaZulu–Natal Drakensberg, Common Elands spend most of their time in open grassland areas, particularly during the late spring and summer when the grass is green and nutritious. There is a vertical movement away from the central grassland area at the onset of the dry and cold winter, when green grass becomes unavailable. Animals then move into the montane forests and turn to browsing. The amplitude of movement ranges from between 5 and 40 km (Keep et al. 1972, Scotcher 1982, Rowe-Rowe 1983). Flehmen plays an important part in courtship and is exhibited by all Common Elands from about three months old. However, it is most prominent in adult bulls, and they actively solicit urine from // by pushing at the vulva. Females in oestrus are closely followed by the dominant bull in the herd, who chases and attacks less dominant ??, and often lays his head on the female’s rump or close to her side. Eventually she stands still to allow mounting, and intromission is brief. Most births take place between 04:00h and 08:00h, peaking just at sunrise (Underwood 1979).The / becomes restless then lies down on her side to give birth; calves, when born, are licked carefully before the afterbirth is eaten by the /. The calves struggle to a standing

position very soon after birth and can run within three or four hours. Calves lie up away from the herd for a few days. However, they soon join a herd and the great majority of calves are found in large nursery groups with other calves and juveniles.Within these groups the calves keep very close to each other in a calf crèche, and are often in close physical contact. Most of their activities take place in this group, and they leave it only to suck. Females visit the crèche to suckle their calves, calling as they approach. Usually the calf will run a short distance from the others to suck. Females do not suckle the calves of others, and sometimes even drive strange calves off. There is no evidence of a bond with the calf lasting beyond the time of weaning (Hillman 1979). Common Elands are not very vocal.They make a deep gruff ‘alarm’ bark similar to that of the Bushbuck Tragelaphus scriptus. Females communicate with their calves using a series of repeated clicks; or by mooing and grunting or a combination of these (Underwood 1973, 1975). The calves respond by whimpering or moaning. At closer proximity the / and calf exchange a weak bleat, and a similar noise is heard from adult ?? during courtship (Hillman 1979). Adult bulls bellow in expressing their dominance or emit a belching grunt in repelling others from food. The larger adult ‘grey’ ?? make a sharp clicking noise while walking as they lift each forefoot, which is audible up to several hundred metres away. Various sources have suggested the sound emanates from the hooves clicking together when the animal walks or from the carpal bones, but Posselt (1963), who worked with domesticated animals for many years, indicated it came from the knees, likely produced when a tendon slips over a carpal bone (Estes 1991a). Hillman (1979) suggested that the sound may act as an indication of the size and dominance of the animal. Subsequently, Bro-Jørgensen & Dabelsteen (2008) have shown that the dominant frequency of the knee-clicks in Common Eland is an honest signal of body size, reflecting both inter-individual variation and intra-individual changes over time. Reproduction and Population Structure  Common Elands are not strongly seasonal breeders, but there is a peak in calving during the early summer months of Aug–Nov in summer rainfall areas (Skinner & Van Zyl 1969, Skinner et al. 1974, Jeffery 1979,

Common Eland Tragelaphus oryx profile of male frontal ‘brush’.

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do not breed under natural conditions at less than three or four years of age. In the KwaZulu–Natal Drakensberg, sex ratio at birth is 1 : 1 (Stainthorpe 1972, Scotcher 1982); among adults, sex ratio has been given as 1 ? to 1.6–2.0 // (Rowe-Rowe 1994).The sex ratio among adult Common Elands in the Athi-Kapiti population studied by Hillman (1979) was strongly female-based, with 1.9 // to every ?. Part of the reason for this may have been the tendency for adult ?? to occur in smaller groups and in thicker vegetation, making them more vulnerable to predators. Rudnai (1974) found a male-to-female ratio of 9 : 1 in Lion Panthera leo kills in Nairobi N. P., Kenya. Over a five-year period in KwaZulu–Natal, mean sex and age structure comprised 15% adult ??, 47% adult //, 12% yearlings and 26% calves (Scotcher 1982). Maximum longevity for wild Common Elands is 14– 17 years, and captives have lived 25–26 years (Hillman 1979, Weigl 2005). Annual calving rates in the Athi-Kapiti population were estimated to be 67% in 1972 and 68% in 1973 (Hillman 1979); those in southern Africa are higher at up to 90% (Buys & Dott 1991) to 95% (Scotcher 1982).

Common Eland Tragelaphus oryx frontal view of male ‘brush’.

Underwood 1979, Scotcher 1982, Buys & Dott 1991), although in the south-western Kalahari the peak appeared to be in Nov (R. Brett pers. comm.). In Kenya, births occurred throughout the year, with a possible peak in Sep–Dec (Hillman 1979). Oestrus occurs at 21–26 day intervals and lasts three days (Posselt 1963). Females typically have a postpartum oestrus (usually about two weeks following parturition; Posselt 1963) and this results in a shorter inter-calving interval. Jeffery (1979) reported the annual inter-calving interval for Common Elands in KwaZulu–Natal as ranging between 281 and 532 days (n = 120; mean = 373); the gestation period was around 273 days (n  = 53; also recorded by Stainthorpe 1972). Usually only a single calf is born, with twins produced in about 2% of births in captivity (Zuckerman 1953). Recorded average birth weights of captive T. o. oryx calves include: 27 kg for ?? (n = 6) and 25 kg for // (n = 9) (Jeffery & Hanks 1981), and 30 kg (n = 20) and 26 kg (n = 30), respectively (Skinner & Van Zyl 1969; see also Stainthorpe 1972).Weaning takes place at about 4–5 months of age.The age at first conception is between 18 and 39 months (Hillman 1979), while ??

Predators, Parasites and Diseases  Lions are the main predators of Common Elands throughout their shared range in Africa, but Common Elands seldom exceed 3% of their prey. This is probably because of the relatively low density of Common Elands compared with other prey species. Wright (1960) stated that Common Elands formed about 2% of Lion kills in S Kenya and C Tanzania, and Rudnai (1974) calculated that Lions killed approximately 2% of the Common Eland population annually in Nairobi N. P. Mitchell et al. (1965) found that Common Elands comprised 3% of Lion kills and 1% of African Wild Dog Lycaon pictus kills in Kafue N. P., Zambia. In a total of 817 African Wild Dog hunts recorded by Creel & Creel (2002) in Selous G. R., Common Elands were hunted on less than ten occasions. Young animals may rarely be taken by Cheetahs Acinonyx jubatus. Starvation caused by drought can be a major cause of death, but Common Elands can sometimes escape its worst effects by moving long distances (Hillman & Hillman 1977). In the 1985 drought in the south-western Kalahari, it was estimated that 35% of the population died (Knight 1995a). In the KwaZulu–Natal Drakensberg, mortality was highest during Aug–Oct following the long, harsh winter, especially among calves and yearlings (Scotcher 1982). Common Elands are described as bovine antelopes and it is reasonable to assume that the species, in its natural habitat, will contract similar viral, bacterial or parasitic infections as described for the cow in Africa, but the severity, symptoms and pathology will vary. For example, the species is susceptible to rinderpest (Thomas & Reid 1944, Robson et al. 1959), and, in an outbreak in Nairobi N. P. in 1996, there was an infection rate of 20% and mortality rate of 10% in Common Elands (Kock et al. 1999). Other important viruses such as foot and mouth disease have minimal impact on the species and they appear to be insignificant as carriers. They show no tendency to contract malignant catarrhal fever virus and their status as carriers of Rift Valley Fever and Lumpy Skin Disease is unknown. Bacterial diseases such as anthrax and parasitic diseases such as anaplasmosis have been reported in the species, and under certain conditions ticks and tick-borne diseases (e.g. theilerosis, transmitted by ticks of the 197

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Family Bovidae

genus Rhipicephalus) can be significant; Rowe-Rowe (1994) notes that some introductions of Drakensberg animals to lowveld areas in KwaZulu–Natal failed because the animals succumbed to tickborne diseases. The bacterium Theileria taurotragi is pathogenic and has caused deaths (Grootenhuis et al. 1980). Their resistance to trypanosomosis (which is transmitted by tsetse flies) is similar to other antelope. Except for the highly infectious diseases such as rinderpest and foot and mouth disease, the behaviour of the species (living in relatively small herds, timid and highly mobile) would reduce the chance of inter-specific infection. Once an individual in a herd contracts an infection, such as tuberculosis or anthrax, a high incidence is likely, since intra-specific dynamics are similar to cattle and buffalo. Common Elands host a variety of ticks.Two animals taken in Central Province, Zambia, were host to seven species: Amblyomma variegatum, Boophilus decoloratus, Rhipicephalus appendiculatus, R. evertsi, R. lunulatus, R. simus and R. supertritus (Zieger et al. 1998b). Common Elands in Kruger N. P. harboured large numbers of Amblyomma hebraeum, but were also host to B. decoloratus, Rhipicephalus appendiculatus, R. zambeziensis, R. evertsi and R. simus (Horak et al. 1983c). In the Karoo of South Africa, Common Elands are good hosts for Ixodes rubicundus (Horak et al. 1987). Another animal examined post mortem was host to seven species of tick (Amblyomma gemma, A. variegatum, Boophilus decoloratus, Rhipicephalus appendiculatus, R. evertsi, R. pulchellus and R. pravus; Grootenhuis et al. 1980). Helminths recorded from Common Elands include three species of nematodes from Central Province of Zambia, Cooperia rotundispiculum, Haemonchus contortus and Oesophagostomum sp. (H. contortus is a bloodsucking parasite that can be pathogenic at even low levels of infection); the tapeworm Moniezia benedeni was also recorded (Zieger et al. 1998a). In Mountain Zebra N. P. and West Coast N. P., Boomker et al. (2000) recorded Haemonchus mitchelli, Nematodirus spathiger, C. rotundispiculum and Bronchonema magna. Conservation  IUCN Category: Least Concern. CITES: Not listed. Common Elands still occur over a large area of southern and eastern Africa, but their numbers and range have declined considerably. Their meat is highly prized and each animal provides a large quantity of meat, so they are particular targets of illegal hunters. In Botswana, their range has contracted over the last 20 years, so that now they are almost entirely confined to protected areas and to private lands.The Common Eland’s habit of wandering over large areas may affect its future in ways that cannot be fully predicted. As available habitat declines, animals’ ability to range widely is being reduced. It is possible that this will make populations that already tend to occur at low densities more vulnerable to environmental disturbances, such as drought and disease. However, Common Elands have been reintroduced to a number of game ranches and private ranchland in southern Africa (particularly South Africa), and this has done much to bolster numbers. In addition, animals have been introduced widely outside of their natural range, primarily to private ranches. For example, although their natural range in Namibia is restricted to the north-eastern parts, they now occur widely on game ranches in the southern and central parts (East 1999). Important naturally occurring populations are protected in Omo N. P. (Ethiopia), Boma N. P. (Sudan), Serengeti N. P. (Tanzania), Kafue and North Luangwa National Parks (Zambia), Nyika N. P. (Malawi),

Etosha N. P. (Namibia), Kgalagadi Transfrontier Park (Botswana/South Africa) and Ukhahlamba-Drakensberg Park (South Africa). According to East (1999), approximately half the total population occurs in protected areas and 30% on private land. At present, the Common Eland is not in any immediate threat of extinction, and this situation is likely to continue as long as there are sufficient protected areas and private ranches to sustain viable populations. Much has been written and published on the topic of domestication of Common Elands, which have been domesticated in Russia, Ukraine, England, Zimbabwe, South Africa and Kenya, for a number of perceived advantages: they provide a high yield of nutritious, ‘long life’, antibacterial milk, which has a high fat content; they can be readily tamed and herded; they can survive in arid regions unsuitable for cattle due to low water requirements; they have a long life expectancy in captivity; and they have a varied diet (Uspenskii & Saglanskii 1952, Posselt 1963, Skinner 1967, Lightfoot 1977, Lightfoot & Posselt 1977a, b). However, although captive animals reproduce successfully, management practices such as high food supplementation costs, confining them at night and herding them during the day are likely to negate their advantages over cattle in many environments (Hillman 1979). Measurements Tragelaphus oryx HB (??): 2510 (2390–2630) mm, n = 5 HB (//): 2270 (2200–2330) mm, n = 5 T (??): 620 (570–720) mm, n = 5 T (//): 530 (500–550) mm, n = 5 E (??): 220 (210–240) mm, n = 5 E (//): 210 (190–230) mm, n = 5 Sh. ht (??): 1630 (1580–1790) mm, n = 7 (on the curve) Sh. ht (//): 1423 (1250–1530) mm, n = 7 (on the curve) Sh. ht (??): 1422 (1355–1500) mm, n = 6* Sh. ht (//): 1296 (1245–1330) mm, n = 6* WT (??): 494 (450–540) kg, n = 5 WT (//): 344 (317–370) kg, n = 5 Athi-Kapiti Plains, Kenya (Hillman 1979) *Serengeti, Tanzania (Sachs 1967), taken between the pegs Common Elands in southern Africa generally are larger than the figures given here. Wilson (1969) gave the mean mass of five mature bulls as 604 kg (range 530–690), and ten // had a mean weight of 445 kg (range 385–470). Skinner (1967) gave the mean mass of 19 mature bulls from bushveld areas as 650 kg (range 425–840). However, wild Common Elands from the KwaZulu–Natal Drakensberg are lighter, with mean adult masses of 453 kg (n = 17) for ?? and 305 kg (n = 61) for // (Scotcher 1982; and see Keep 1972).While weights of ?? in Serengeti N. P. were similar to those of the Athi-Kapiti plains, weights of // were considerably lower, with a range of 277– 321 kg, and horns were shorter in both sexes (Sachs 1967) Maximum recorded horn length is 118.4 cm for a pair of horns from Grootfontein, Namibia (Rowland Ward) Key References  Hillman 1979, 1987, 1988b; Posselt 1963; Scotcher 1982; Taylor 1969; Underwood 1973, 1975, 1979. Chris R. Thouless

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Family Bovidae

Subfamily ANTILOPINAE – Antelopes, Sheep, Goats Antilopinae Gray, 1821. London Med. Repos. 15: 307. Tribe Neotragini  Neotragus (2 species)  Nesotragus (1 species) Tribe Cephalophini  Philantomba (2 species)  Sylvicapra (1 species)  Cephalophus (16 species) Tribe Raphicerini   Raphicerus (3 species)   Dorcatragus (1 species) Tribe Madoquini   Madoqua (7 species)* Tribe Antilopini  Gazella (4 species)  Eudorcas (4 species)  Nanger (5 species)*  Ammodorcas (1 species)  Litocranius (1 species)  Antidorcas (1 species) Tribe Ourebiini  Ourebia (1 species) Tribe Reduncini  Pelea (1 species)  Redunca (3 species)  Kobus (5 species) Tribe Oreotragini   Oreotragus (1 species) Tribe Aepycerotini  Aepyceros (1 species) Tribe Alcelaphini  Beatragus (1 species)  Damaliscus (2 species)  Alcelaphus (1 species)  Connochaetes (2 species) Tribe Hippotragini  Hippotragus (2 species)  Addax (1 species)  Oryx (3 species) Tribe Caprini  Ammotragus (1 species)  Capra (2 species)

Dwarf Antelopes Suni

p. 207 p. 213

Blue Duikers Common Duiker Forest Duikers

p. 223 p. 235 p. 244

Grysboks, Steenbok Beira

p. 303 p. 315

Dik-diks

p. 320

Slender Gazelles Ring-horned Gazelles Greater Gazelles Dibatag Gerenuk Springbok

p. 339 p. 356 p. 372 p. 387 p. 390 p. 398

Oribi

p. 405

Grey Rhebok Reedbucks Kobs

p. 416 p. 421 p. 437

Klipspringer

p. 469

Impala

p. 479

Hirola Damalisks Hartebeest Wildebeests

p. 490 p. 495 p. 510 p. 527

Roan Antelope, Sable Antelope Addax Oryxes

p. 547 p. 566 p. 571

Aoudad Ibexes

p. 594 p. 599

*including species-groups

The particular selection of antelope taxa that have been allocated to this subfamily has varied greatly over the years as has the relative ranking of Antilopinae within the family Bovidae: both remain subject to debate. In the treatment presented here, the Antilopinae embraces all the non-Bovinae bovids and these, colloquially referred to as antelopes, sheep and goats are held to share a common ancestry. A basal split between bovine and non-bovine taxa has long been recognized on morphological, physiological, ecological and behavioural criteria. Furthermore, a number of recent molecular studies have confirmed the reality of this most fundamental division of the family Bovidae (Hassanin & Douzery 1999, 2003, Matthee

Thomson’s Gazelle Eudorcas thomsonii (frontal view).

& Robinson 1999a, Matthee & Davis 2001, Hassanin et al. 2012). Antilopinae, being the prior sub-familial name for all non-bovine antelopes, is therefore used in that broader sense, to the exclusion of nomenclatures that would restrict this rank to a smaller subset of gazelle-like species (Eisenberg 1981, Rebholz & Harley 1999, Hernández Fernández & Vrba 2005 and others), which are herein classed as the tribe Antilopini. In distinguishing Antilopinae from Bovinae there are physiological differences that have a much broader, adaptive and biogeographic significance. Antilopinae regulate their body temperature by the advanced water-saving mechanism of nasal panting (although many of the Reduncini cool their bodies by sweating, this is almost certainly the secondary reversion of antelopes that have become tied to wellwatered habitats). The thermoregulatory mechanism employed by all Bovini is the older, typically mammalian system of sweating, which is dependent upon continuous access to water and can be presumed to be the original condition of the main stem or founding bovid stock (for example, Gentry [1978] regarded the Asiatic boselaphines as the closest living descendants of early bovids and these bovines are also ‘sweaters’). The development of ‘nasal bellows’ and nasal panting is possibly linked in some way with nasal ‘whistling’ calls that are uttered by all but the largest antelopes: the nasal areas of Bovinae are not modified for sound and they never utter any such calls. Antilopinae frequently have pedal glands (although individual species or groups have secondarily suppressed them), whereas all Bovinae lack these glands. Antilopinae also usually have facial glands (or their putative ancestors had typical glandular fossae on their skulls), although they are lacking in some species (e.g. Aepycerotini). Bovinae, on the other hand, lack facial 199

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Family Bovidae

Bright’s Gazelle Nanger (granti) notata black and white design on female udder.

glands (although some have glandular areas with a different histology). Antilopinae often have a single pair of nipples, although a number of tribes retain the older pattern of two inguinal pairs; all Bovinae have two pairs of nipples. Most Antilopinae have annulated horns (although a few conservative species have short, smooth spikes). Bovinae, by contrast, have smooth horns with a circular section or spiral ones (generated by differential growth rates at the base of the horn sheath). The differences listed above (and more detailed physiological study will undoubtedly uncover more) are merely morphological markers for a fundamental divergence among early ancestral bovids. The more conservative species of Antilopinae are generally small or very small and are limited to small and stable home-ranges: the founding antilopine lineage would appear to have become adapted to hot and relatively dry conditions at an early date. Bovines, instead, generally have large or very large body sizes, often live nomadic lives in unstable habitats, and are adapted to wetter and generally cooler conditions. The original divergence between Bovinae and Antilopinae probably involved a continental separation between the two ancestral stocks with the majority of some 24 species of Bovinae (sensu Grubb 2005) incontestably of Eurasian origin, whereas two-thirds (and by far the most diverse array) of about 120 Antilopinae are African. This separation broadly corresponds to adaptation by Bovinae to more temperate climates in the northern continents whereas stem Antilopinae adapted to the generally tropical climates of Africa. Recent molecular studies have shown that some small and very conservative species of African antelopes are basal Antilopinae, and their ecological and genetic distinctness implies speciation taking place very early on in the radiation of this group. The existence of small-bodied folivores among Australian macropods and Eurasian deer demonstrates that there are niches for small-bodied folivores in a wide range of habitats in all continents. Once such niches are occupied (and small African Antilopinae evolved more than 20 mya, probably at the beginning of the Miocene), the main selective pressure on them is to become better adapted to the specifics of their particular habitats, such as hot, arid thickets, cooler scrubs, equatorial forest or rocky hillsides. In particular, there is much to suggest that the smaller Antilopinae have made progressive improvements in physiological adaptation to local climates and environments. On the other hand, there has been much less pressure to alter the basic anatomical specifications incumbent on small size because selection has continued to favour a narrow range of body-sizes no less in Africa than other taxa in other continents. Although the common ancestor of all Antilopinae was likely to have

been a small animal, molecular evidence suggests that enlargement took place at a very early stage in antelope evolution (implied, for example, by the demonstration of basal status for the Impala Aepyceros; e.g. Georgiadis et al. 1990, Hassanin et al. 2012). The evolution of bigger bodies involves a chain of other adaptive changes and it seems significant that a majority of the more highly divergent, larger-bodied antelopes appear to derive from the same large-bodied basal lineage as the Impala. The survival of several very conservative basal lineages needs to be understood in this context of very narrow parameters for anatomical change in small antelopes, but much greater potential for elaboration in the larger lineages. Taking account of dated antilopine fossils and the probable time needed to achieve today’s very diverse radiation, Kingdon (1982) estimated a divergence date between Bovinae and Antilopinae of about 23 mya (Early Miocene). Using molecular clock techniques, Hassanin et al. (2012) have estimated that the split took place around 19.7 mya. Soon after arrival in Africa, antilopines began to adapt to local extremes of habitat and climate and sometime in the early or middle Miocene an early antilopine found its way back into Eurasia and this lineage gave rise to the predominantly Asiatic Caprini (Hassanin & Douzery 2003, Hernández Fernández & Vrba 2005). Subdivisions within the Antilopinae remain problematic, but seven major groups have long been recognized: the duikers, Cephalophini; the dwarf antelopes, Neotragini; the kobs or waterbucks, Reduncini; the gazelline antelopes, Antilopini; sheep and goats, Caprini; horselike antelopes, Hippotragini; and the alcelaphines, Alcelaphini. Each of these groups has radiated (in the case of caprines, into as many as 32 species). However, with the advent of molecular trees and clocks, a much more complex radiation has become apparent. The ‘Neotragini’, a grouping that formerly embraced some 13 or 14 species, and long suspected to represent contemporary survivors of a 1

5

4

3

9

2

10

8 6

6 a

7

b

a. Evolution of the brow buffer illustrated by a gradient between upright, smooth stabbing horns and slanted corrugated horns in some Antilopinae. 1. Grey Rhebuck Pelea capreolus; 2. Steenbok Raphicerus campestris; 3. Klipspringer Oreotragus oreotragus; 4. Beira Dorcatragus megalotis; 5. Oribi Ourebia ourebi; 6. dik-diks Madoqua and Suni Nesotragus moschatus. b. Progressive elevation of the corrugated anterior surface in the horns of Antilopinae: 6. primitive condition, as in dik-diks Madoqua and dwarf antelope Neotragus; 7. initial elevation primarily due to thickening of horn (tip upturns, as in some Reduncini); 8. elevation, lengthening and thickening of horn in primitive Antilopini, as in Dibatag Ammodorcas clarkei; 9. elevation of longer, thicker horn with increased convex curvature, as in some gazelles Gazella spp. and Gerenuk Litocranius walleri; 10. horn arched high above orbit as in some advanced Antilopini, Caprini and Hippotragini.

200

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Family Bovidae

25

20

15

10

5

0 mya Bovinae

Reduncini

Ourebiini

Antilopini

Raphicerini

Madoquini

Oreotragini ? Cephalophini ? Neotragini

Aepycerotini

Alcelaphini

Hippotragini

Caprini

Tentative phylogenetic tree for Bovidae (modified from Hassanin & Douzery 2003, Hernández Fernández & Vrba 2005 and Hassanin et al. 2012).

basal type from which all the Antilopinae evolved (Kingdon 1982), has now been confirmed to be paraphyletic (Georgiadis et al. 1990, Gentry 1992, Gatesy et al. 1997, Matthee & Robinson 1999a, Rebholz & Harley 1999). Recognition that several living species occupy basal positions, and do not belong to any of the seven traditional subgroups of antilopinae, has suggested a bold biogeographic pattern that accords with much broader patterns of adaptation by Eurasian immigrant biota into Africa. New outlines for the primary and secondary radiations of Antilopinae are beginning to emerge and, in spite of numerous discrepancies between published trees, some convergent data (and opinions) are emerging, and these accord reasonably well with the following reconstruction of the antilopine adaptive radiation, based on a best fit between ecology, behaviour, palaeontology, anatomy and genetics. In spite of having been assembled before the advent of molecular trees or clocks, the first detailed effort to reconstruct this radiation (Kingdon 1982) conformed, to a surprising degree, with contemporary molecular findings. Some mammalian immigrants entering Africa were able to do so because they were already very widespread, cosmopolitan species, but others, perhaps the majority, would have entered with readymade ecological preferences compatible with the environments of (the usually rather narrow) corridor that permitted intercontinental exchange. Subject to the particular climatic cycle of their incursion,

whether cool or hot, the majority were probably adapted to dry but seasonally variable habitats. In the 27–23 million years since Africa regained contact with Eurasia there has been no true tropical forest connection and, barring some bat species, even the most adaptable mammals would have taken time to adapt to equatorial habitats. It is, therefore, significant that the three or four basal lineages that have emerged from molecular studies imply a very early adaptation by the first antelopes to three or four major adaptive niches. The closest living approximation to the primary adaptive niche and to the relatively generalized anatomy of the earliest Antilopinae is the Cape Grysbok Raphicerus melanotis. Supposing then that the ancestral immigrant antilopine was a small, thicket-dwelling ruminant living under seasonally hot, but relatively dry, climates, a primary challenge for any such incoming group was to adapt to the more equatorial habitats of Africa, where diseases and all manner of environmental challenges awaited any immigrant. For small herbivores living on the forest floor, wetter climates meant higher, darker canopies and less ground-level foliage. Smaller bodies for an already small-sized antelope were the most likely outcome and the primitive dwarf antelope tribe Neotragini conforms with just such a prediction in terms of primarily equatorial distributions, sizes, habitats and feeding ecology. Genetically, neotragines are basal antelopes close to the primary radiation of all Antilopinae (Gatesy et al. 1997, Rebholz & Harley 1999, Hassanin et al. 2012), but they must have made their accommodation to tropical forest well after the first Antilopinae differentiated from ancestral Bovinae. Because a very mountainous region lies immediately south of the main connection between Africa and Eurasia, many mammal immigrant groups can be shown to have made initial adaptations to the mountains of north-eastern Africa. Here, too, molecular trees agree in suggesting a basal position for the Klipspringer Oreotragus oreotragus, the only representative of the Oreotragini (and an archetypically Ethiopian species, although now widespread down the south-eastern half of Africa). Another primary innovation would have been exploitation of more open, thicket-edge habitats. Here larger-bodied, longer-legged populations can be predicted because resources are more dispersed and predators at more of an advantage. Here, too, there is good conformity with molecular trees with a very early split accounting for the Impala as the only representative of a basal lineage, the Aepycerotini. All molecular trees imply that the major antilopine lineages have each developed lineage-specific and identifiable adaptations to particular habitats, but none is consistent in distinguishing primary from secondary radiations. The primary ones suggested here are: 1 Small-bodied, thicket-dwelling lineage under dry climates typified by an early and conservative derivative, Raphicerus. 2 Dwarfed, forest-dwelling lineage typified by Neotragus. 3 Smallish, mountain- and rock-dwelling lineage typified by Oreotragus. 4 Larger, ecotone-dwelling lineage typified by Aepyceros (which might share common roots with either one of the two previous lineages or the common ancestor of all three). It is suggested here that all other major antilopine tribes are likely derivative from one or other of the four primary lineages listed above, a conclusion that is broadly consistent with most recent molecular studies. 201

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Family Bovidae

Reduncini (+Pelea) Antilopini and subsequent derivatives

Ourebia

Oreotragus Raphicerus campestris Dorcatragus Raphicerus melanotis Cephalophini Madoqua kirkii Madoqua guentheri Madoqua saltiana

Raphicerus sharpei Nesotragus moschatus Neotragus batesi Neotragus pygmeus

Body size and climate as major parameters in the radiation of antilopines. Some derivative lineages are indicated outside the 20 kg box.

Gazelline antelopes (Antilopini), as well as dik-diks (Madoquini) and kobs (Reduncini), can plausibly be derived from the earliest antilopine ancestral stock. The Beira Dorcatragus megalotis shows both morphological and molecular characteristics that suggest genetic links between Raphicerus and the gazelles and their kin.The dik-diks also relate to the same lineage on both counts. All these lineages show progressive physiological adaptation to heat-tolerance and water-conservation, but remain conservative in relation to a browsing, or ‘concentrate-selector’ diet (Hofmann 1973, 1989). The gazelles span a wide range of sizes and degrees of sociality, whereas dik-diks retain residential, territorial habits, small size and a highly selective fresh leaf diet. The Reduncini have been proposed as a basal group (Hassanin & Douzery 2003), but they have a still unresolved relationship with two other antelopes. One, the Grey Rhebok Pelea capreolus, seems to represent the localized derivative from a very early reduncine. The other, the successful and widely distributed Oribi Ourebia ourebi, shares an odd mix of characteristics with the Steenbok Raphicerus campestris and members of the Reduncini. The Oribi’s basal relationship to Raphicerus has been confirmed (Hernández Fernández & Vrba 2005), but its link to the Reduncini has been less widely accepted. However, its resemblances with reduncines are too numerous and too detailed to be easily explained by convergence. In direct opposition to the previous thicket-dwelling or arid-adapted Antilopini, reduncines and Oribi have become open valley grazers, relying, to a greater or lesser degree, on a very abundant but unstable food supply and continuous access to water.

This has involved very substantial physiological changes compared with most other Antilopinae, notably a water-dependent mode of cooling, by sweating, but also remarkable resistance to direct radiation from the sun. A conservative reliance on hiding, whether in dense grass, in swamps and water-logged areas or in large, dense herds, betrays the fact that Reduncini have very poor stamina and are, in that respect, conservative antilopines. They are most likely to be an early but secondary offshoot of Raphicerus-like Antilopini. The duikers (Cephalophini) were once assumed to be among the most primitive of bovids (Estes 1974). In an early genetic analysis, Georgiadis et al. (1990) clustered duikers with the non-Bovinae but were unable to associate them with any other antilopine group (an outcome that could have suggested basal status). Duiker origins have proved remarkably opaque to molecular analysis and suggestions as to their closest affinities include Bovinae (Gentry 1992), somewhere in between Bovinae and Antilopinae (Jansen van Vuuren & Robinson 2001), Reduncini (Gatesy & Arctander 2000a, Kuznetsova et al. 2002) and Antilopini (Matthee & Davis 2001). They have been grouped in a variety of supposed clades; one permutation linked duikers with reduncines, hippotragines and alcelaphines (Gatesy & Arctander 2000b), another placed them with caprines, alcelaphines and hippotragines (Castresana 2001), and still other studies highlighted a close association with the Klipspringer (Hassanin & Douzery 1999, Matthee & Robinson 1999a, Hassanin et al. 2012). Thus, geneticists have invoked virtually every major bovid tribe as sister taxa for the Cephalophini!

202

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Family Bovidae

Transformation of the head in three related antelopes. Frontal and profile diagrams of Oribi Ourebia ourebi (top left), Grey Rhebok Pelea capreolus (top right) and Mountain Reedbuck Redunca fulvorufula (lower right) compared with reconstruction of hypothetical ancestor (lower left). Note depth of masseter is an indicator of fibre in diet.

Reconstruction of ancestral African antelope skull.

Duikers share forest environments with the dwarf antelopes of the genus Neotragus, but differ in many aspects of their anatomy and behaviour (see Kingdon 1982). Instead of relying on scarce vegetation, duikers have enlarged their diet to include some animal matter, but rely to varying degrees on fallen fruits, seeds and other plant debris from the canopy. While their social systems show some adaptive variation, they tend to live spaced out in territories and they have not elaborated horns beyond short spikes. The radiation of duikers appears to originate in a very diminutive ancestor with the larger species being among the most recently evolved (Kingdon 1982, Jansen van Vuuren & Robinson 2001). This implies that duikers might represent a secondary radiation deriving from a diminutive Neotragus-like ancestor and the arguments for this origin were detailed in Kingdon (1982) and are summarized on p. 220. As for dating the divergence of duikers, the date of 6.3–5.6 mya (Jansen van Vuuren & Robinson 2001) seems rather late and extrapolated dates of 12.3–8.9 mya (Hassanin et al. 2012) might be possible. The Klipspringer has no close genetic relatives (Gatesy et al. 1997, Matthee & Robinson 1999a, Rebholz & Harley 1999), and can be characterized as the single highly specialized survivor of a primary stem lineage. Some molecular resemblances with duikers have been observed and could suggest a phylogenetic connection, but the degree of specialization in the Klipspringer is so great that any very early relationships with later groups, whether duikers or caprines, are now obscured. What is clear is that this species

has paralleled some of the adaptations of goats and chamois and its early capture of the rocky mountain niche in Africa has probably discouraged the expansion of more recently evolved wild caprines into any habitats other than those in the Sahara and the highest reaches of Ethiopia. Whether these similarities are wholly convergent, or not, is debatable but some peculiarities of caprines would seem more likely to have found their roots in something like an early oreotragine or aepycerotine than in any other basal Antilopinae. Molecular analyses have suggested that the caprines might share ancestry with several superficially non-caprine antelopes and their common ancestor must have been very close to being a stem lineage. It is in this context that Aepycerotini is treated here as a true basal lineage that may be as close to Neotragini as it is to Oreotragini. The Impala is the single surviving species of the Aepycerotini. At least one synthesis of recent studies (Hernández Fernández & Vrba 2005; and see Hassanin et al. 2012) has suggested that this apparently basal group has given rise to three major secondary radiations, the Caprini (over 30 spp.), the Hippotragini (7 spp.) and the Alcelaphini (6 spp.). Following Gentry (1978) and Vrba (1984), many authors have treated Aepyceros as a primitive alcelaphine, a relationship that has been shown to be only remotely true because it may be as closely related to the hippotragines and caprines. The more advanced members of the Caprini are late, mainly Asiatic and unquestionably belong to a secondary radiation. The Hippotragini are mostly arid-adapted grazers, living at low densities in arid or impoverished zones. The Alcelaphini, by contrast, are high density, ecotone/catenary grazers in open but less arid habitats. All three groups can be regarded as secondary radiations deriving from a single stem lineage. However, the living survivor of that stem lineage, the Impala, bears more resemblance to one of its derivatives, Beatragus, an alcelaphine, than it does to any of the horse antelopes or caprines. The radiation of Antilopinae is summarized on p. 201 as a composite tree in which the dates of splits or branchings (as currently understood) are derived from disparate sources. The prolific radiation of antelopes has attracted several theoretically minded scientists, and the complex diversity of species has been broken down into various simpler categories (e.g. Schlosser 1904, Jarman 1974, Gagnon & Chew 2000). Vrba (1980) sought to explain their evolution and diversity by an ‘effect hypothesis’ in which less speciose ‘generalists’ were contrasted with speciose ‘specialists’. Every such hypothesis has been hostage to the slender informationbase that was available at the time of its inception, but recent field and laboratory studies have significantly improved our knowledge and potential for understanding one of the major radiations of large mammals in the world. The current spectrum of Antilopinae embraces a diversity of folivorous, graminivorous (and frugivorous) niches and the ability of so many species to inhabit the same landscapes is based upon very precise foraging and digestive techniques. The details of many of these herbivorous strategies are described in the profiles that follow, but the likely development of antilopine niches can be summarized as follows. The initial traits that distinguished early Antilopinae (in contrast to Bovinae) were small size, stable home-ranges and improved temperature and water regulation that allowed them to inhabit 203

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Table 8.  Diversification by habitat of Bovinae and Antilopinae. Asian origin

Large. Grazing, humid/unstable habitats

Asian origin. Africa entry 15 mya

Broad spectrum foliage gleaning

Bovini

Tragelaphini

African origin (17 mya?). Solitary gleaning (fruit, etc.), closed habitats

Cephalophini

Small, solitary, initially drier habitats

Dwarf bovids

Early Africa entry (22 mya)

African origin

Grazing in moist valleys

Reduncini

Mainly African

Gleaning in open arid habitats

Antilopini

African origin

Enlargement pioneer. Browse closed-to-open

Aepycerotini

Large. High-density grazing, less arid habitats

Alcelaphini

African origin

Large. Low-density grazing, impoverished zones

Generalized diets, impoverished habitats

drier habitats than their competitors. Their ruminant digestion ensured that they could maximize the extraction of nutrients and were therefore metabolically superior to the hyraxes, herbivorous macroscelids, rodents, chevrotains and larger non-bovid herbivores that preceded them. That dietary advantage is undoubtedly one reason for the success of antelopes in Africa. Among Antilopinae, members of the tribe Antilopini have become ‘gleaners’ for high-quality foliage in arid or semi-arid habitats. Body sizes range from small to moderately large and the more advanced species are relatively social. Among the most conservative of Antilopinae are the grysboks and Steenbok (Raphicerini), while most arid-adapted of all the small conservative tribes are the Madoquini. The Reduncini have moved furthest from this pattern and have become specialists in exploiting tropical valley grasslands, often at very high densities. Body sizes range from medium to large. Neotragini (Neotragus and Nesotragus) are minuscule and solitary gleaners of ground-level foliage in or on the edge of true forest. The Cephalophini are also gleaners, but mainly of fallen fruit and have developed many interesting sub-strategies, some of which are directly dependent on the wasteful feeding habits of primates, hornbills and bats. They are mostly solitary and territorial and range from 3.5–80 kg.

Hippotragini

Caprini

The Alcelaphini have become large-bodied ecotone and catenary grazers, mostly in open habitats and tend to live at relatively high densities. The Hippotragini are large-bodied antelopes that have adapted to arid habitats or impoverished zones and live, for the most part, at low densities. Caprini are predominantly a late Asian radiation that evolved in direct competition with Cervidae. Many are adapted to extreme seasonal variations in diet and have colonized or recolonized mountain habitats in between glaciations. In Africa, a unique monospecific genus (Ammotragus) inhabits desert hills, mountains and plateaux and the slopes of desert valleys in the Sahara, and a second genus, the ibexes Capra, have colonized mountains down the western shores of the Red Sea as far as C Ethiopia. Antilopinae span a wide range of body sizes, weighing as little as 1.5 kg (Royal Antelope Neotragus pygmaeus) to 300 kg (several spp.). Many of the larger Antilopinae have interesting coat patterns that show increased contrast along the margins of limbs or those body regions that are typically counter-shaded in small cryptic species. Common origins for all antilopines are evident in similarities of patterning among members of different tribes. Antelope skulls range from tiny compact structures with short spike horns, large preobital fossae to enclose large facial glands above

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Occlusal views of upper right toothrows. (Upper left) Oribi Ourebia ourebi (grazer). (Upper right) Mountain Reedbuck Redunca fulvorufula (grazer). (Centre left) Cape Grysbok Raphicerus melanotis (generalized diet). (Centre right) Aders’s Duiker Cephalophus adersi (partial frugivore). (Lower left) Suni Nesotragus moschatus (‘concentrate selector’). (Lower right) Bates’s Pygmy Antelope Neotragus batesi (strict folivore). Sketches of ‘signal geometry’ in three antilopine species; from left Beisa Oryx Oryx beisa, Sable Antelope Hippotragus niger and Waterbuck Kobus ellipsiprymnus. Note eye emphasis or eye suppression.

a short, finely pointed muzzle to heavy, elongated skulls surmounted by heavily reinforced horn cores. In spite of their diversity, horns share a basic structure among all Antilopinae: differential rates in the growth of horn sheaths and in the thickness of keratin deposits are the main determinant of species-specific horn shape and these have been explored in some detail in Kingdon (1982). The teeth of Antilopinae have been widely misinterpreted, and in one of the first efforts to classify bovids, Schlosser (1904) broke down the wide variety of dental types into ‘boodonts’ (with a long shallowly rooted toothrow with molar lobes and basal pillars) and ‘aegodonts’ (with hypsodont teeth, deeper teeth and roots in a shortened toothrow). This classification became entrenched in the literature on bovids for nearly 80 years and was misleading because the teeth of different lineages responded to tougher diets and increased body size in very similar ways. Thus, larger-bodied, more hypsodont lineages, such as Hippotragini and Reduncini, became unnaturally clumped on the basis of similarities that are primarily due to larger size and more grass in the diet. A shift from softer diets in smaller antelopes to harder diets in larger

Oribi Ourebia ourebi myology.

ones takes place within Antilopini, Reduncini and Caprini, while the largest antelopes in Reduncini, Hippotragini and Alcelaphini have all become hypsodont independently (Kingdon 1982). The transformation of a typical ‘early’ antilopine toothrow can be illustrated by comparing the rows of Raphicerus melanotis with those of Ourebia ourebi (as an intermediate type) and Redunca fulvorufula (as a specialized grazer). The masticatory action of a browser slices rather than mills soft leaves and the toothrow can be compared with the action of pinking shears or a long saw-edged blade: this functional morphology is most evident in the Neotragini. To turn a long narrow toothrow over to a macerating action, milling edges across the face of each molar can be increased by simply folding enamel surfaces while the spaces in between the layers of crenellated enamel are bulked out with dentine filler. The combination of extra enamel folds and pillars, more soft dentine and greater depth and rooting both improves the milling action of mastication and lengthens the life of the tooth. The front end of the toothrow declines in importance among grazers because a side-to-side or lateral mode of mastication, while more efficient for milling hard grasses, needs to be concentrated and needs maximum buttressing for the mechanical advantage to be optimal, hence the decline of premolar teeth in grazers. All antelopes share longish, slender lower limbs and large-eyed, horned heads above longish necks but proportions differ greatly depending on size, habitat, range and other factors. The smaller, more conservative types have hunched postures and are able to make short, very fast dashes into cover but have no stamina. The larger, grazing species tend to have straight backs, longer legs, necks and faces and many are able to run far and fast, especially those living in open plains where there are numerous predators. The social behaviour of antelopes has been widely studied, and has been extensively summarized in several works, in particular Estes (1991a). The more conservative species are territorial and residential, with male and female territories generally shared. Many of the larger species are highly mobile and land tenure and behaviour are varied (Jarman 1974). Antelopes occupy all the major habitats of Africa from forests to deserts and from sea level up into the high mountains. Jonathan Kingdon 205

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Tribe NEOTRAGINI Dwarf Antelopes Neotragini Sclater & Thomas, 1894. The Book of Antelopes 1: 2.

Suni Nesotragus moschatus.

The tribe Neotragini has long been the acknowledged ‘waste-paper basket’ receptacle for a diverse collection of small-sized antelopes that resembled each other in their short straight horns, compact skulls, preorbital glands and slender legs. However, several recent studies have shown that the Neotragini, traditionally including Dorcatragus, Madoqua, Neotragus/Nesotragus, Ourebia, Oreotragus and Raphicerus, is paraphyletic (Georgiadis et al. 1990, Gentry 1992, Gatesy et al. 1997, Hassanin & Douzery 1999, Matthee & Robinson 1999a, Rebholz & Harley 1999). Since the differences between these antelope genera have long found acknowledgement, tribal status within Neotragini has herein contracted to only three former members of the Neotragini. All the rest have been allocated to other tribes. With the advent of molecular science and the prospect of objective measures of genetic affinity, several former neotragines have been

shown to be more distantly related to one another than they are to other, less conservative groups. Furthermore, their resemblances can be interpreted as retentions indicative, in several instances, of their basal position in the radiation of African antelopes. The dwarf antelope genera Neotragus and Nesotragus constitute one such basal group, but more than one genetic study has associated the little Suni Nesotragus moschatus with another basal species, the much larger, rather gazelle-like Impala Aepyceros melampus (Georgiadis et al. 1990, Hassanin & Douzery 1999, Matthee & Robinson 1999a, Hassanin et al. 2012; but see Matthee & Davis 2001).This has served to emphasize the genetic isolation of Neotragus and has accelerated abandonment of the former ‘waste-paper basket’ category. Neotragini is here considered to embrace two species in the genus Neotragus, Bates’s Pygmy Antelope N. batesi and the Royal Antelope N. pygmaeus, and the monospecific genus Nesotragus for the Suni.

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left:

Suni Nesotragus moschatus adult male myology. Suni Nesotragus moschatus skeleton.

above:

Neotragine species resemble one another in retaining their attachment to a narrow range of forest types and retaining, too, a conservative dentition and feeding habits. It would seem likely that those primitive antelopes that succeeded in abandoning such dietary and ecological attachments became transformed and effectively founded entirely distinct lineages. The earliest precursors of such lineages were probably somewhat Suni-like or grysbok-like and the retention of a way of life typical of the earliest antelopes continues to give the behaviour and ecology of Nesotragus and Raphicerus a special interest. On a personal note, the decision, in 1964, to embark on a regional inventory of mammals and to entitle it An Atlas of Evolution in Africa (Kingdon 1971–1984) was my realization of the significant role that the then‘Neotragine’ antelopes must have played in bovid evolution. Contemporary molecular studies have borne out the validity of that proposition. Other aspects of the contemporary biology of this tribe are discussed under Neotragus and Nesotragus and in the species profiles.

Suni Nesotragus moschatus juvenile head. Details of sensory hairs around muzzle, eyes and ears.

Jonathan Kingdon

Genus Neotragus Dwarf Antelopes Neotragus C. H. Smith, 1827. In: Griffith et al., Anim. Kingd. 5: 349.

Most recent classifications include the allopatric Royal Antelope Neotragus pygmaeus, Bates’s Pygmy Antelope N. batesi and the Suni Nesotragus moschatus in Neotragus (e.g. Grubb 2005). However, the species have been split into three genera in the past: Neotragus sensu stricto for the Royal Antelope, Hylarnus for Bates’s Pygmy Antelope and Nesotragus for the Suni. In this work, only Neotragus and Nesotragus are retained (see genus Nesotragus profile for further discussion). Neotragus in this sense only occurs in the rainforests of western and central Africa (including N. pygmaeus and N. batesi) – the only other small rainforest antelopes apart from duikers.The Suni, here retained in the genus Nesotragus, occupies a wider range of more mesic habitats in eastern Africa from Kenya south to KwaZulu–Natal. Members of Neotragus are dwarf antelopes (shoulder height 20– 33 cm, mass 1.4–3 kg), with relatively long hindlegs, and pelage

plain coloured or with traces of agouti-speckling or streaked. The rhinarium is naked. Preorbital glands open by a pore; inguinal and pedal glands are both present. There are no lateral hooves. Females have two pairs of nipples. Horns are present in the ? only, directed backward more or less in the plane of the forehead. They are short, thin and smooth in both Royal Antelope and Bates’s Pygmy Antelope (cf. the longer and stouter horns, with up to 20 well-marked annuli, in Suni). In the skull, a premaxillo-maxillary vacuity is no more than a kink in the Royal Antelope, but vestigially present in Bates’s Pygmy Antelope. Apart from the Suni, this character is unknown in other extant bovids except the Impala Aepyceros melampus (Ansell 1972). Its possible evolutionary significance was discussed in Kingdon (1982) and elsewhere in this volume. A small ethmoid fissure is present in 207

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Cartesian co-ordinates applied to the nasal maxilla and premaxilla of: Suni Nesotragus moschatus (left), Bates’s Pygmy Antelope Neotragus batesi (centre), Royal Antelope Neotragus pygmaeus (right).

the Royal Antelope, but absent in Bates’s Pygmy Antelope. In the former, the nasals are posteriorly narrow and pointed, whereas they are broadened and blunt in the latter species. There are large but shallow preorbital fossae accommodating the preorbital glands (larger in ??). The incisiform teeth are small, the first incisors broad and others narrow. Upper milk canines are absent in Bates’s Pygmy Antelope (and in the related Suni), but often present in the Royal Antelope. The cheekteeth are low-crowned, much like those of Raphicerus, and the premolars well developed.

Neotragus differs from Raphicerus in the shape of the muzzle, larger preorbital fossa, less upright horns, longer tail and presence of inguinal glands. Gene analysis suggests Neotragus and Nesotragus are distantly related to other traditional neotragine antelopes, and are probably better considered the only members of the tribe Neotragini (Hassanin & Douzery 1999, Matthee & Davis 2001), a course that is followed in this work. Peter Grubb

Neotragus batesi  Bates’s Pygmy Antelope (Dwarf Antelope, Bates’s Dwarf Antelope) Fr. Antilope de Bates; Ger. Batesbockchen Neotragus batesi de Winton, 1903. Proc. Zool. Soc. Lond. 1: 192. ‘Efulen, Bulu Country, Kamarun [Cameroon], 1500 ft [457 m] above the sea’.

Taxonomy  Polytypic species with two recognized subspecies. Originally designated as a monospecific genus, Hylarnus (De Winton 1903), it has also been allocated to the subgenus Nesotragus (Haltenorth 1963), but Nesotragus is here retained only for the Suni N. moschatus. Synonyms: harrisoni. Chromosome number: not known. Description  Very small antelope, with short muzzle, large eyes and long thin legs. Pelage sleek and fine mahogany-brown, darker on back, crown and forehead. White spots at the base of ears; chin and throat white.Young may have a cream spot above eye. Preorbital glands large, especially in ??, round and not invaginated. Chest, belly, inner parts of upper forelegs and hindlegs, front fetlocks white. Lateral hooves rudimentary. Pedal glands are present. Tail short, inconspicuous brown, lighter below. Females are 6% larger (total length) and 21% heavier than ??. Narrow folivorous teeth. Horns present in ? only, small, smooth, weakly ringed at base, parallel to forehead line, tips slightly converging. In contrast to the Royal Antelope N. pygmaeus and the Suni, the ethmoid fissure is absent in Bates’s Pygmy Antelope. There is a vestigial hiatus between the maxillary and intermaxillary (the premaxillo-maxillary vacuity, as in the Suni), and there is no anteorbital vacuity. Bates’s Pygmy Antelope Neotragus batesi.

Upper-right toothrow of Bates’s Pygmy Antelope Neotragus batesi.

Geographic Variation N. b. batesi: SE Nigeria, east of Niger R. to Cross R., and S Cameroon, Equatorial Guinea, Gabon, SW Central African Republic and Congo. N. b. harrisoni: NE DR Congo and SW Uganda. More contrasted pelage, with more white on the legs (see Thomas 1906).

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Bates’s Pygmy Antelope Neotragus batesi skulls, adult male above, subadult below. Neotragus batesi

Similar Species Neotragus pygmaeus. Only occur in forests west of Dahomey Gap. Smaller in size, more reddish pelage, rufous collar thinner than in Bates’s Pygmy Antelope; lateral hooves absent. Nesotragus moschatus. Only occur in forests and thickets east and south of the Eastern Rift Valley. Larger in size, freckled dark brown pelage; horns of ?? longer, finely annulated and slightly uprising; lateral hooves absent. Philantomba monticola. Broadly sympatric. Larger in size and of more stocky build, grey or grey-brown. Distribution  Endemic to Africa. Patchily distributed in moist lowland forest in three disjunct regions: SE Nigeria, east of the Niger R. to the Cross R.; S and SE Cameroon (south of the Sanaga R.) to SW Central African Republic (west of the Sangha R.), Gabon, and NW and SW Congo; and NE DR Congo, north and east of the Congo–Lualaba, extending marginally into SW Uganda (East 1999). The gap in distribution may be due to ancient dry corridors during past glacial periods. Today, swamp forests from N Congo and DR Congo may constitute a barrier of unfavourable habitat. Habitat  Bates’s Pygmy Antelopes inhabit moist lowland forest. They prefer dense, low undergrowth along rivers, tree falls within mature forests, areas regenerating after logging or cultivation, road sides, village-gardens and plantations where they exploit patches of high-quality forage growing where light reaches soil (Dragesco et al. 1979, Feer 1979, Hart 1985). According to Mbuti hunters of NE DR Congo, they avoid damp, swampy habitats (Carpaneto & Germi 1989). In NE Gabon and S Cameroon, they apparently are excluded by drought during dry seasons (expressed as rainfall minima in the driest season, 25–50 mm per month) (Feer 1979). Abundance  Locally very abundant within optimal habitat such as in cocoa or cocoa plantations mixed with secondary forest in NE Gabon (35–75/km2; Feer 1979); uncommon in the Lopé N. P. in

Gabon (White 1994, P. Henschel pers. comm.). Densities over more extensive areas of forest are 1.5–2.2/km2 (Hart, J. A. et al. 1996, Fa & Purvis 1997). In forests of S Central African Republic, encounters of Bates’s Pygmy Antelopes during net hunts represent only 0.2% of total artiodactyls (Noss 1999). In the Ituri Forest, NE DR Congo, it represented 3.6% of the captures by nets (Carpaneto & Germi 1989). Here, estimated densities in mature forest are similar to those in regrowth (Wilkie & Finn 1990). Despite higher hunting pressure, encounter rates within 14 km from a village in Dja Reserve in Cameroon were higher than at distances further away (Muchaal & Ngandjui 1999). East (1999) estimated a total population size of about 220,000 individuals. Adaptations  Typically slow-moving while foraging, with a highstepping and hesitant gait in dense vegetation. Ratio of free height under chest to shoulder height is higher than for duikers (tribe Cephalophini).When alarmed may freeze a long time before jumping for cover. Flees quickly at short distances, occasionally with barking. Active both at night and day, with rests around mid-day and in the early and middle evening. In cultivated zones, animals rest in the densest areas by day and relatively more open areas by night. At night they move 51% less in open habitats (generally rich in food) than in other habitats. Lactating // and young prefer dense protective habitats (Feer 1979). Foraging and Food  Folivorous; classed as browsers by Gagnon & Chew (2000) in a comprehensive review of dietary preferences in African Bovidae. Bates’s Pygmy Antelopes delicately pluck leaves, buds and shoots, choosing small young items, but also consuming very large leaves (e.g. Dioscorea, Calocasia). They browse selectively on more than 200 species in cultivated areas in Gabon (Feer 1979), with recorded preferences for species in the genera Brillantaisia, Momordica, Phaulopsis, Cyathula and cultivars, notably sweet potatoes, peanuts and peppers. Near villages, they adapt their home-ranges 209

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to the seasonal rotation of clearings and growth of small plants; on average, half of their home-range is composed of dense herbaceous strata (Feer 1979). Their diet in deep forest is poorly known. They consume four species of forest fruit and various leaves according to Mbuti Swa from the Ituri Forest (Carpaneto & Germi 1989). Cerling et al. (2003) found that three different teeth taken from a single individual from the Ituri Forest had the most 13C-depleted bioapatite values from the more than 1000 extant mammals they had analysed, indicating that the species effectively has no capacity to survive on coarse grass or leaf diets. Social and Reproductive Behaviour  Most often solitary (61% of observations, Feer 1979); two or more adults are rarely seen within short distance (15 cm), large pedal gland, a striking superciliary line, the absence of a strongly marked break on the haunches between the dark croup, light colored flanks and lower haunches. Craniometrically, P. walteri can be differentiated from the two other species in the genus by a clearly smaller nasal constriction and cranial height (Colyn et al. 2010). Colin Groves

Philantomba maxwelli  Maxwell’s Duiker Fr. Céphalophe de Maxwell; Ger. Maxwellducker Philantomba maxwellii (C. H. Smith, 1827). In: Griffith et al., Anim. Kingd. 4: 267. Sierra Leone.

Lateral view of skull of Maxwell’s Duiker Philantomba maxwelli.

Maxwell’s Duiker Philantomba maxwelli.

The species is named for Sir Charles William Maxwell, who was Governor of Senegal in 1809 and then of Sierra Leone (origin of the type) in 1811. Taxonomy  Ansell (1972) and Ralls (1973) recognized three subspecies: the nominate form from Senegal and Gambia to Sierra Leone; P. m. lowei (including the form danei) from the islands in the Rokelle R. in Sierra Leone and with unknown limits; and P. m liberiensis, from Liberia eastwards. Grubb & Groves (2001) recognize two subspecies: the nominate form, P. m. maxwelli, is distributed through the species range except for the Yatward and Sherbro Is., where it is replaced by the subspecies P. m. danei¸ which is smaller (skull length under 136 mm). Synonyms: danei, frederici, liberiensis, lowei, philantomba, whitfieldi. Chromosome number: 2n  = 60; Maxwell’s Duiker has only acrocentric X chromosomes (Hard 1969, Robinson et al. 1996b).

Description  Small-sized antelope varying in colour from light, sandy-brown to greyish-brown or brownish-black. The narrow and angular face tapers off to a small pointed muzzle. The upper jaw clearly surpasses the lower jaw and the mouth droops, which gives the Maxwell’s Duiker a grim facial expression. There is no definite crest between the horns. The curved preorbital glands are heavily demarcated, and studded with as many as 25 pores. There is frequently a lighter brow-like line above the large black eyes. The oval ears are frequently fringed with white, and together with the tail, which is white below, are constantly flicked back and forth, making conspicuous fieldmarks. The belly and inside of legs are paler than the rest of the body. The slightly elongated rump and relatively narrow neck give a lighter build to this animal than to other duikers. The legs are very slim and relatively long, and the buttocks are angular. The tail is fully haired and untufted (Grubb & Groves 2001). There are no inguinal glands, and the well-developed pedal glands lie in a subcircular sac at the end of a narrow canal (Pocock 1910). Horns are short (seldom more than 6 cm), heavily ridged, thick at the base, and bent slightly inwards and backwards. Horns are frequently lacking in //, and when present are usually much shorter and less heavy at the base (Wilson 2001).

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Geographic Variation P. m. maxwelli: distribution as for species, but not present on Yatward and Sherbro Is.There is some evidence of geographical variation in anatomical features, but patterns of variation are complex (Grubb & Groves 2001). The frequency of nuchal hair reversal increases from west to east. Most measurements of skull size increase from east to west between Togo and Liberia, but there are discrepancies from this pattern in Ghana, and the pattern does not hold for the extreme eastern and western parts of the range. The length of horns in ?? decreases from east to west, but again, Ghana does not entirely fit the pattern, and horns are largest in the extreme west. The proportion of // with horns decreases westwards from 100% in Nigeria and Togo to only 5 out of 80 in Liberia, but there are few data for the west of the range. Geographical variation in coat colour is masked by a high degree of individual variation (Grubb & Groves 2001). P. m. danei:Yatward and Sherbro Is. Small size: skull length 128–136 mm (cf. >136 mm in P. m. maxwelli); nasals 40–44 mm (cf. >45 mm in P. m. maxwelli); preorbital 64–65 mm (cf. >66 mm in P. m. maxwelli); and toothrow 36–41 mm (cf. >41 mm in P. m. maxwelli). Similar Species Philantomba monticola. The two species are separated geographically, with Maxwell’s Duiker occurring to the west of the Cross R., and the Blue Duiker to the east. Blue Duikers are significantly smaller; according to Grubb & Groves (2001), the largest skull of a Blue Duiker they recorded was 139 mm, whereas this is closer to the minimum for Maxwell’s Duiker; also differs in colour (with strongly marked break on the haunches between the dark croup and the light colour of the flanks and lower haunches; cheeks are darker and superciliary streaks less prominent); length of tail (usually less than 100 mm); a shallower, smaller pedal gland; and cranial characteristics (skull narrower across the zygomata; orbital borders more protuberant; rostrum more

Philantomba maxwelli

constricted; narrower palate; free ends of nasals narrower on average) (Grubb & Groves 2001). Distribution  Endemic to Africa. Widespread within the Upper Guinean forest zone, in lowland forests and extending into forest– savanna mosaics, from W Gambia and SW Senegal east to the Cross R. in Nigeria (East 1999, Wilson 2001). Distribution is restricted by the drier savanna–sahel zone in the north. Since they do occur in gallery forests within savanna woodlands in parts of West Africa (e.g. N Côte d’Ivoire and SW Burkina Faso), it is reasonable to assume that they may occur in SW Mali, though there are no confirmed records to date (Wilson 2001).Their current distribution is probably not much changed from their historical distribution, although much of their original habitat has been modified or lost. Habitat  Found in mature and secondary forests, gallery forests and forest patches, coastal scrub and farmland (Aeschlimann 1963, Davies 1987, Newing 1994, 2001, Grubb et al. 1998).They have a preference for dense undergrowth, but occasionally penetrate into fields close to forest patches. They adapt well to farmland and in Sierra Leone they occur on montane grassland over 1400 m on the Loma Mts. The animals do not require permanent water within their territories, even if nursing, provided enough fresh fruits are available (Nett 1999). Abundance  Abundant throughout the forest zone in both mature forests and secondary vegetation, but decreasing where hunting pressure is high. Davies (1991) estimated population densities of 15–30/km² in mature logged forest in Gola N. P., Sierra Leone, and Wilson (2001) thought this was the most common antelope in the country. This is also the most common duiker in Côte d’Ivoire, where Newing (1994, 2001) recorded a density of 79/km² in a mixed, hunted farmland near Taï N. P. and 63/km² in lightly logged primary forest with minimal hunting, within the Park. Elsewhere in Côte d’Ivoire, Nett (1999) recorded 16–20/km² in heavily hunted and logged semi-deciduous forest patches in the eastern part of the country, and Hoppe-Dominik (1989) found 2.7/km² in the savanna– forest mosaics of Marahoué N. P. In the National Park of Upper Niger, Guinea, Maxwell’s Duikers had the highest density (3.7/km²) of a total of ten species of ungulates present (Brugière et al. 2005). East (1999) suggested a total population size of 2,137,000 individuals – likely a conservative estimate. Adaptations  Maxwell’s Duiker is the smallest and most opportunistic of seven sympatric duiker species in the Upper Guinean forest block. It inhabits even heavily degraded forest patches and hides in the dense cover of the exotic Akyempong weed species Chromolaena odorata (Newing 1994, 2001, Nett 1999). Wilson (2001) suggests that numbers may even have increased as a result of forest clearance for farmland and the spread of Akyempong. The success of Maxwell’s Duikers in secondary vegetation types is doubtless made possible by a very broad diet including a large variety of small fruits and leaves, and an ability to adapt to a more leafy diet at times of fruit scarcity (Newing 1994). In contrast, the sympatric species are significantly larger than Maxwell’s Duiker (Bay Duiker Cephalophus dorsalis, Black Duiker Cephalophus niger, Yellow-Backed Duiker Cephalophus silvicultor and Jentink’s Duiker Cephalophus jentinki) and/or more restricted to mature forest (Zebra Duiker Cephalophus zebra, 225

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Jentink’s Duiker and Ogilby’s Duiker Cephalophus ogilbyi) or to secondary forest (Yellow-backed Duiker) (Newing 2001). Foraging and Food  Maxwell’s Duikers are primarily frugivorous, but also eat leaves and mushrooms and may depend on leaves at times of fruit shortage. The two most comprehensive studies of feeding ecology to date are those of Hofmann & Roth (2003) and Wilson (2001). Hofmann & Roth (2003) examined 139 stomachs from the bushmeat market of Toumodi in Côte d’Ivoire.These authors recorded 78 different types of fruits eaten, with Maxwell’s Duikers showing a particular preference for Nauclea latifolia, Ficus capensis, Canthium vulgare, Blighia sapida, Griffonia simplicifolia, Alchornea cordifolia, Phoenix reclinata and Spondias mombin. Wilson (2001) examined 250 stomachs from bushmeat markets in Ghana, and listed 33 species of fruit, including Ficus spp., Solanum spp., Blighia spp. and Uvaria spp.; all fruits eaten were from cultivated lands, secondary forest or savanna. Both Wilson (2001) and Hofmann & Roth (2003) recorded an average of three fruits per stomach. In the eastern semi-deciduous forests of Côte d’Ivoire, hunters named the fruits of Ricinodendron heudelotii, Strombosia glaucescens, Trichilia monadelpha (= T. heudelotii), Ficus exasperata, Celtis adolfi-fridericii and Ceiba pentandra as their primary food sources (D. Nett pers. obs.). Aeschlimann (1963) examined two stomachs of Maxwell’s Duikers in the evergreen forest zone and found seeds of Turraeanthus africanus and Pycnanthus angolensis; fruits of Musanga cecropioides; leaves of Griffonia simplicifolia, Ficus barteri, Baphia nitida and Fagara parvifolia, and several varieties of mushrooms. Newing (1994, 2001) listed 26 species of fruits found in eight stomach samples and a further two species eaten during direct observations of free-living animals in mature forest. Hofmann & Roth (2003) and Wilson (2001) both recorded animal matter in the form of ants from stomachs. In suitable habitats Maxwell’s Duikers are said to follow groups of colobus monkeys and guenons, in order to profit from fallen fruits.The guenons include the Lesser Spot-nosed Guenon Cercopithecus (cephus) petaurista, Mona Monkeys Cercopithecus mona and Diana Monkeys Cercopithecus diana. In heavily degraded forests Maxwell’s Duikers are often found in close association with squirrels, which may take over the primates’ ecological role when the latter are no longer abundant (Nett 1999). Maxwell’s Duikers have a bimodal diurnal activity pattern with peaks in the early morning and late afternoon. Social and Reproductive Behaviour  Maxwell’s Duikers live in monogamous pairs or occasionally polygamous groups (e.g. Taï N. P.; Newing 1994) with or without young, with a common territory. Newing (1994) recorded home-range sizes of about 5 ha in mature forest and 3.6 ha in secondary vegetation, while Nett (1999) recorded 18.4 ha in relict forest.Territory size seems to increase with the age of the animal (e.g. 12.5 ha for one subadult compared with an average of 20 ha each for three adult animals). Territory size also appears to decrease in accordance with increasing available resources (especially fruits) and shows no seasonal changes. Overlap between territories of neighbouring groups is small (0–23%) and temporary. In a shared territory the borders of the female’s territory sometimes lie outside the male’s, suggesting that both sexes mark and defend against conspecifics (Nett 1999). Newing (1994) observed physical fights between neighbouring ?? and chases of ?? or male–female couples by neighbouring ??. Ralls (1975) observed Maxwell’s Duikers in captivity and documented agonistic behaviour in both

sexes, even though fighting was much more aggressive in ??. The author described Maxwell’s Duikers as extremely pugnacious with the ability to chase even other antelopes such as the larger Bay Duiker. Territorial boundaries are marked by middens and scent-marking. Mutual scent-marking with the preorbital glands strengthens the bonds of a monogamous pair; it is also carried out by two individuals of the same sex prior to heavy fighting. All marking is carried out by both sexes. Courtship is initiated by the ?, who pursues the / and attempts to lick the base of her tail while the / may be lifted off her hindfeet. Ralls (1973) reports that the tail of an oestrous / is often wet and frayed from the male’s intensive chewing and biting. Circling noseto-tail, flehmen (lip curling) and laufschlag (leg tapping) precede mounting attempts; the / only stands still once she is ready to mate, and copulation is very brief. Maternal care is reduced to a minimum in Maxwell’s Duikers even though the mother grooms the young by licking its fur. The young does not follow its mother but is laid up in a protected spot until about three months old, when it begins eating significant amounts of fruits and leaves (Wilson 2001, H. Newing pers. obs.). Two kinds of nasal vocalization accompany fighting, sexual and anti-predator displays: an often repeated short snorting and the louder intense alarm-call. The latter seems to have disappeared in some areas where hunting pressure is heavy, since hunters try to attract the animals by imitating these calls, and even playbacks no longer elicit a response. This might be an adaptation of Maxwell’s Duikers to human pressure (Nett 1999). Reproduction and Population Structure  Births occur throughout the year. In Côte d’Ivoire and Nigeria there is a suggestion of two birth peaks – one in the main dry seasons, from Dec to Mar, and one from Jul to Sep (Aeschlimann 1963, Happold 1987, Newing 1994), butWilson (2001) could not find any evidence of a birth peak in Ghana. Of 131 uteri examined by Wilson (2001), every one had the foetus implanted in the right uterine horn, although ovulation took place from either ovary. A single young is born that resembles the parents in colour, and weighs between 600 and 950 g (Aeschlimann 1963, Wilson 2001). Basic information on length of gestation period and age at sexual maturity remains contradictory. Numerous authors have cited a gestation period of 120 days (e.g. Aeschlimann 1963, Ralls 1973); however, Kadjo (2000) gave gestation as 188 days and Wilson (2001) gives a mean gestation length of 205 days (n = 4; range 198–213). Hofmann et al. (1998) recorded first oestrus in // of a heavily hunted wild population at six months. Wilson (2001) recorded four captive // conceiving at between 8 and 12 months, and earliest successful mating by a ? at 10 months. However, Kadjo (2000) recorded first oestrus only at 18 months. Females become receptive 3–5 days after giving birth. Hofmann et al. (1998) found a male to female sex ratio of 1 : 1.04 in the markets of Toumodi, Côte d’Ivoire (n = 1721) and 1 : 1.11 in Kumasi, Ghana (n = 858). The ratio of adult to young was 1 : 2.86 and 1 : 1.75, respectively. The maximum birth rate was 1.8 young/ year/reproductive /; 81–93% of the adult // were pregnant. These high figures may have been due to hunting pressure. Maxwell’s Duikers can live for more than ten years in captivity (Aeschlimann 1963, Ralls 1973).

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Predators, Parasites and Diseases  Common predators are African Rock Pythons Python sebae, Crowned Eagles Stephanoaetus coronatus, Leopards Panthera pardus and African Golden Cats Profelis aurata. Helminths recorded include nematodes (genera Ostertagia, Bunostromum, Setaria, Skrjabinodera, Trichuris) and cestodes (Avitinella) (Round 1968, Kamara 1975). In a study of the ticks associated with wild mammals in Ghana, Ntiamoa-Baidu et al. (2005) recorded the following species from duikers in Ghana: Haemaphysalis parmata, H. leachi, Ixodes aulacodi, I. cumulatimpunctatus, I. moreli, I. muniensis, Rhipicephalus senegalensis, R. simpsoni and R. ziemanni. Maxwell’s Duiker has not been identified as an important vector for diseases of domestic animals and humans although it is doubtless susceptible to all major ungulate diseases. Conservation  IUCN Category: Least Concern. CITES: Not listed. Maxwell’s Duikers are widespread and seem to support hunting pressure better than other duikers, due to a broad tolerance of habitat disturbance and diet (Newing 2001). Nevertheless, they are among the most hunted ungulates in much of their range, and around 30% of the hunted ungulates in secondary forest zones in Côte d’Ivoire are Maxwell’s Duikers, which means an average of nearly 3% of the hunted mammal populations in this region (Nett 2002). Some 30% of the delivered biomass in the bushmeat restaurants in the region of Taï N. P. is made up of Maxwell’s Duikers (Caspary et al. 2001). Intense poaching in Comoé N. P. in Côte d’Ivoire led to numbers of Maxwell’s Duikers declining by more than 90% within 20 years, resulting in a low density estimate of 0.04/km² (Fischer & Linsenmair 2001a). Brugière et al. (2005) also recorded significant decreases in this species between 1997 and 2002 (whereas other ungulate species increased or remained stable), and attributed this to changes in hunting patterns in the area. Whereas hunters used to come from the whole of the Haut Niger region and hunted in large groups for several weeks in the Mafou forest prior to its gazettement as a core area of the Park in 1997, solitary or small groups of hunters now originated

mainly from nearby villages and enter the forest for short periods (less than a day), hunted opportunistically (almost exclusively by night), and hence focused on the most abundant species. Maxwell’s Duikers are still common in several protected areas in their range, particularly Ziama and Diécké Forest Reserves, National Park of Upper Niger and Mt Nimba Strict N. R. (Guinea), Gola N. P. and Tiwai Island Game Sanctuary (Sierra Leone), Sapo N. P. (Liberia), Taï, Azagny, Mont Sangbe and Marahoue National Parks (Côte d’Ivoire) and Kakum, Bia and Mole National Parks (Ghana). Measurements Philantomba maxwelli TL (??): 860 (820–940) mm, n = 184 TL (//): 890 (830–950) mm, n = 91 T (??): 140 (120–160) mm, n = 184 T (//): 150 (120–160) mm, n = 91 HF c.u. (??): 200 (180–240) mm, n = 184 HF c.u. (//): 210 (180–230) mm, n = 91 E (??): 62 (58–69) mm, n = 184 E (//): 60 (58–69) mm, n = 91 Sh. ht (??): 350 (320–420) mm, n = 184 Sh. ht (//): 360 (330–420) mm, n = 91 WT (??): 7.5 (6.5–11.2) kg, n = 184 WT (//): 8.0 (6.5–12.0) kg, n = 91 Ghana (Wilson 2001) Includes only non-pregnant //. Average body weight for 131 pregnant // was 9.1 kg (range 7.5–12.6) Maximum recorded horn length is 6.6 cm for a pair of horns from Sierra Leone (Rowland Ward) Key References  Aeschlimann 1963; East 1999; Nett 1999, 2002; Newing 1994, 2001; Ralls 1973, 1975; Wilson 2001. Dorothe Nett & Helen Newing

Philantomba spp. frontal views of Maxwell’s Duiker P. maxwelli (left); Blue Duiker P. monticola congicus (centre); and Blue Duiker P. m. monticola (right). Several current subspecies may well prove to be full species.

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Philantomba monticola  Blue Duiker Fr. Céphalophe bleu; Ger. Blauducker Philantomba monticola (Thunberg, 1789). Resa uti Europa Africa, Asia …, 2: 66. South Africa, ‘Lange Kloof’; since identified as borders of Western and Eastern Cape, Uniondale and Humansdorp Dist., Langkloof, 33° 48´ S, 23°  to 24° 30´ E; see Grubb (1999: 21).

Blue Duiker Philantomba monticola adult female.

Taxonomy  Ansell (1972) recognized 16 subspecies, noting that the range limits and validity of many require further investigation. Grubb & Groves (2001), in their review of the classification of the duikers, recognized 12 subspecies, divided in two major groups, a greylegged and a red-legged group (see Geographic Variation). Synonyms: aequatorialis, aequinoctialis, anchietae, bakeri, bicolor, caerula, caffer, congicus, defreisi, fuscicolor, hecki, ludlami, lugens, melanorheus, musculoides, minuta, nyasae, pembae, perpusilla, pygmea, ruddi, schultzei, schusteri, simpsoni, sundevalli. Chromosome number is 2n = 60, the same as P. maxwelli (Hard 1969, Robinson et al. 1996b). Description  The smallest duiker in Africa, which varies greatly in colour and has differentiated into a large number of regional and insular forms. Blue Duikers have large eyes, somewhat swollen nostrils, small ears, lined with off-white hair, and a very wide, flexible mouth. The forehead profile is relatively flat. The pelage varies greatly in tonality and colour ranging from near black in some subspecies to slate-grey, bluish-grey and various shades of brown. The length and density of hair-cover varies between populations. The lower legs are generally lighter in tone than the back and the black

hooves are narrow and pointed. There are no inguinal glands, and the pedal gland (which secretes a whitish liquid) has a simple orifice, unlike that of Maxwell’s Duiker P. maxwelli which has a subcircular sac at the end of a narrow canal between the hooves (Pocock 1910). The preorbital gland exudes secretions through a series of pores that are arranged along an oblique arc across the swollen gland (the pores in most other duikers have an almost straight alignment). The secretion, while fresh, has a pale bluish-grey colour, and the chemical compounds are discussed in detail by Burger & Pretorius (1987).The tail, which is vigorously whisked up and down (not side to side), has very conspicuous white hair lining on its underside and a nearly black dorsal surface. Compared with Maxwell’s Duiker, the skull is notably smaller, shorter (typically 139 mm), and with longer tail, as much as 160 mm in length, whereas in Blue Duiker few specimens have tails longer than 100 mm. Also differs in colour, lacking the strongly marked break on the haunches between the dark croup and the light colour of the flanks and lower haunches; the cheeks are lighter and superciliary streaks more prominent (Grubb & Groves 2001). Distribution  Currently, many populations occur nearly con­ tiguously across all of central Africa, but the two major clusters of subspecies, as described above, represent a significant and potentially long-standing geographic separation. Past climatic fluctuations are likely to have fragmented both northern and southern types, but two principal barriers appear to have maintained the primary north–south divide. These barriers are the Congo R. in the west and the semiarid corridor that, in East Africa, separates well-established, southeastern, and mainly coastal, biota from a more recently expansive, predominantly western and equatorial biota (Kingdon 1971, 1982, 1990). Until recently, the Blue Duiker ranged throughout much of central, eastern and southern Africa. Localized populations occurred wherever suitable forest and thicket existed and this is still one of the few species of duiker that continues to survive over most of its historically known range (East 1999). In the west, the species occurs eastwards from the Cross R.; Happold (1987) mapped the Niger R. as the western 229

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authors (e.g. Bowland 1990) make mention of Blue Duiker in the Magaliesberg Mts in Limpopo Province, but Rautenbach (1982) never recorded them anywhere in the former Transvaal. There are, as yet, no confirmed records from Swaziland (Monadjem 1998) and none from southern Mozambique (Smithers & Lobão Tello 1976), suggesting a break in distribution from N KwaZulu–Natal to E Zimbabwe/C Mozambique. Habitat  Thrives in a wide range of forested and wooded habitats, including mature and secondary forests, gallery forests, dry forest patches, coastal scrub farmland and regenerating forest from sea level up to 3000 m. Bowland (1990) found the species in Podocarpus mist forests on the KwaZulu–Natal coast. In eastern Africa and eastern DR Congo, the species has a preference for areas with dense undergrowth, but animals that were living at exceptionally high densities in Gabon were thought to prefer areas with relatively little undergrowth (Dubost 1980). Blue Duikers can persist in small patches of modified or degraded forest and thicket, even on the edge of urban centres. Availability of free-standing water is not an essential habitat requirement (Dubost 1980, Mockrin 2010). Philantomba monticola

limit, but the Blue Duiker seems to be absent or extinct in the intervening area, although Maxwell’s Duiker occurs (see Wilson 2001 for discussion of western limits). This species then ranges east through the forest and ecotone belts of S and C Cameroon, Gabon, Equatorial Guinea (including the island of Bioko), Congo, Cabinda (Angola), most of DR Congo, across S Central African Republic and locally north, with populations in gallery forests and dry relict forest patches in the north of the country. Recorded at 8° 17´ N on the Koumbala R., in Manovo-Gounda-St Floris N. P. (Malbrant & Maclatchy 1949), through extreme SW Sudan (Azande lowlands) and SE Sudan (Imatong and Dongotona Mts). Its range in East Africa comprises a number of disjunct populations in forest patches of S and W Uganda, W Kenya and E and S Tanzania and possibly in W Rwanda and Burundi, as well. The species is absent from the dry country east of L. Tanganyika, but occurs again in the coastal forests and scrub of Mozambique, Kenya and in Tanzanian forests. Andanje et al. (2011a) have confirmed the presence of the species in low numbers north of the Tana R. in the Boni-Dodori forests in N Kenya, a range extension of some 200 km. The Blue Duiker occurs on the islands of Pemba, Zanzibar and Mafia (Moreau & Pakenham 1941, Pakenham 1984, Kock & Stanley 2009) and there have been somewhat implausible suggestions that one or more of these isolates could have been the result of human introduction. It is also found in forested areas of the Tanzanian Southern Highlands (including Ufipa) and the Eastern Arc Mts (Usambara, Uluguru and Udzungwa). The species’ southern range includes localized populations in N and C Angola, where the current distribution is not well known (but is mapped south to at least 15° S by Crawford-Cabral & Veríssimo 2005), through remnant forests and thickets in Zambia, Malawi, E Zimbabwe and parts of C Mozambique on both sides of the Zambezi (East 1999, Wilson 2001). In South Africa, this species is primarily confined to the evergreen forest and thicket along the coast and adjacent hinterlands from N KwaZulu– Natal to the eastern Western Cape Province; there are no records of the Blue Duiker occurring west of George (Wilson 2001). Some

Abundance  Usually the most abundant duiker in communities where it occurs, and co-exists with up to seven other small ungulate species (including up to five other duikers) in some central African forests. Dubost (1980) recorded densities of 70/km2 in a small area (74 ha) at Ipassa, Makokou, in NW Gabon, though it is unlikely that such high densities can be extrapolated over larger areas. In the Ituri Forest, DR Congo, non-hunted populations surveyed on 5 km2 study areas range from 21/km2 in mixed evergreen forest, to 10/km2 in large stands of mono-dominant Gilbertiodendron dewevrei forest, representing about half the total small ungulate population in a community of eight species and 17–18% of small ungulate biomass (Hart 2000). Most population surveys report densities from 5 to 35/km2 across the species’ range, with the exception of some very small populations in isolated habitats in southern Africa (Payne 1992, Bowland & Perrin 1994, Hart 2001, Newing 2001, Wilson 2001, Lannoy et al. 2003). East (1999) estimated total population size at more than 7 million animals. Adaptations  The ecological and demographic success of Blue Duikers is made possible by several factors, not least among which is their extraordinary fecundity, which, in turn, seems to depend upon their ability to maintain a high nutritional plane across a wide range of habitats. Blue Duikers’ consistently good metabolic and reproductive condition is assisted by a rumen anatomy (e.g. densely papillated rumen and fungiform papillae) that is typical for antelopes with concentrated nutrient diets (Faurie & Perrin 1995), but the animals are also sustained by an exceptionally broad diet. In addition to small fruits, seeds, gum, fungi and fallen flowers, they eat standing and fallen foliage, including, remarkably, dry fallen foliage especially when fruits are in short supply (Kingdon 1982, Hart 1985, Newing 2001, Wilson 2001). Possible explanations for such dietary adaptability are more complex than would appear at first sight. For example, their small body size allows them to occupy smaller patches of habitat and exploit dispersed food sources, but the combination of small size and a more leafy diet is counter-intuitive, even contradictory, in the

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Lateral and palatal views of skull of Blue Duiker Philantomba monticola.

sense that most ruminant bovids become better able to cope with difficult foliage diets the larger they are: yet this small duiker is better able to sustain itself on dead leaves than many of the larger duikers. Philantomba is a basal group that derives from the earliest beginnings of the duiker radiation (Jansen vanVuuren & Robinson 2001).Therefore, the Blue Duiker’s ability to shift to a leafy diet may be as much a reflection of its phylogenetic history as a proximate or late specific adaptation. Duikers are thought to derive from small, already folivorous ancestors with the larger species having made ever-more decisive secondary shifts into frugivory and omnivory (Kingdon 1982). In the course of evolution the folivorous habits of early, small duikers would have pre-adapted them to buffer temporary shortages of the more nutritious elements in their diet. This residual flexibility, deriving from their folivorous ancestry, might have combined with their small size to give them advantages over less flexible competitors. It is possible, therefore, that the ecological success of the Blue Duiker is of very long standing. Furthermore, a long evolutionary history, relative to other duikers, could have special implications for understanding progressive adaptation by some Blue Duiker populations to the localized conditions of their region or phytogeographic zone. This offers scientists unique opportunities to study refinements of physiological adaptation along several parameters or gradients (such as altitude, latitude, relative humidity or vegetation type) as well as the role of genetic drift or selection in isolates. Here is an economically important forest mammal that spans an enormous geographic range, has adapted to strong seasonal changes on the forest floor while regional populations have further refined their adaptation to quite localized conditions. While this duiker remains reasonably numerous across most of its far-flung range the opportunity should be seized to try to understand the foundations for its extraordinarily wide range and sustained ecological success. Wilson (2001) has pleaded for an intensification of research on digestive physiology and reproduction in duikers (and on this species in particular), before it is too late. The most eye-catching feature of this species’ behaviour is the continuous flagging, up and down, of the tail, the fringe of which is composed of slightly crinkled white hairs that reflect light exceptionally well. In appropriate lighting conditions this flashing resembles a discrete heliographic signalling device. Quite remarkably for a species that characteristically spends much time alone, this flashing (which in the

Blue Duiker Philantomba monticola.

dim lighting of the forest understorey may be the only visible feature of an otherwise sombrely coloured animal) would appear to be oriented towards other animals in its dispersed community, including individuals that share a territory. Of special interest is the development of horns in the // of some populations and their absence in others. Extrapolating from the Klipspringer Oreotragus oreotragus, it is possible that recently, or at some time in the past, there was a correlation between high population densities and the development of female horns. Given the very substantial differences in density that have already been recorded, this subject offers opportunities for interesting and fundamental research. Foraging and Food  Blue Duikers feed selectively from the forest floor on a wide diversity of ripe and unripe fallen fruits and seeds. They also eat freshly fallen foliage, flowers, pieces of bark, fungi, resin (in particular exudates from Albizia spp.) and some animal matter (Wilson 2001). Of 18 stomachs examined byWilson (2001) in Chirinda Forest, Zimbabwe, the average stomach contained 75% fruits, 15% browse, 5% flowers and 5% fungi. In Gabon, foliage averaged 20% of rumen contents collected throughout the year, while over 67% of contents consisted of small fruits and seeds measuring 0.5–2 cm, the balance being made up of flowers, animal matter and fungi (Dubost 1984). Gautier-Hion et al. (1980) recorded that stomach contents contained 79% fruit, 20% leaves and very small quantities of animal matter, flowers and gum. On the other hand, in KwaZulu–Natal, Bowland (1990), also from an analysis of stomach contents (n = 12), found that about 70% was made up of dicotyledonous leaves and 23% seeds and fruits, probably because fruit was an unreliable resource whereas freshly fallen leaves were available year-round and abundant. Similarly, Hanekom & Wilson (1991) reported that dicotyledonous leaves constituted 57% of stomach contents from Tsitsikamma N. P., South Africa. None the less, Bowland (1990) recorded a strong correlation between fruit and freshly fallen leaves in the diet, suggesting a much greater preference for the former. Animal matter is not uncommonly recorded in stomachs (Grimm 1970, Gautier-Hion et al. 1980, Dubost 1984, Wilson 2001), and Dubost (1984) witnessed Blue Duiker hunting ants by licking them off the ground; ingestion of 231

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Diptera was less common, although duikers were seen snapping at them in flight. In Chirinda Forest, the fruits of the various Ficus species were the most favoured fruits, with duikers eating large numbers of fruits (one stomach contained over 65 F. chirindensis fruits); other fruits eaten included Croton sylvaticus, Dovyalis macrocalyx, Rauvolfia caffra and Diospyros abyssinica (Wilson 2001). In Uganda, the seeds and fruit of the genera Maesopsis, Ricinodendron, Cordia, Musanga and Pycnanthus and the leaves of the genera Mammea and Mildbraediodendron have been recorded as preferred foods (Kingdon 1982). Most of these genera occur in the Ituri Forest and have been recorded in duiker diets (Hart 1985). Other important foods recorded in the Ituri Forest include fruits and seeds from a variety of species in the families Caesalpiniaceae, Sapotaceae, Sapindaceae, Sterculiaceae, Rubiaceae, Ulmaceae and Euphorbiaceae. The seeds of Pancovia harmsiana, Landolphia spp. and Cola spp. are frequent in the diet. Blue Duikers prefer a number of fruits only at an unripe stage, notably Klainedoxa gabonensis, Irvingia grandifolia and Blighia welwitschii, all large trees that regularly abort large portions of developing fruit crops at an early stage. When ripe, these fruits are too large for Blue Duikers to handle, though they are eaten (and in the case of K. gabonensis and I. grandifolia, the seeds dispersed) by the larger duiker species (Hart 1985). The species also joins many other primary consumers by concentrating on seeds of Julbernardia seretii, Cynometra alexandri and Gilbertiodendron dewevrei during periods of mast fruiting, and body fat levels rise during these periods (J. A. Hart pers. obs.). Blue Duikers, like the larger duikers, though to a lesser extent, forage on several species of fruits with large armoured seeds (Ricinodendron heudelotii, Chrysophyllum pruniformes, Celtis adolfi-friderici, Canarium schweinfurthii) that they regurgitate during rumination and disperse (Hart 1985, Feer 1995). In feeding trials with captive animals, Blue Duikers were consistently selective and had reduced capacity to digest structural carbohydrates compared with several of the larger duiker species (Hart 1985, Faurie & Perrin 1993, Newing 2001, Plowman 2002). Preferences of forest fruits and seeds determined by paired food choices in trials were not correlated with any single nutritional or chemical constituent (Molloy & Hart 2002). Blue Duikers preferred fruits over foliage, but when confronted with less preferred fruit choices, they increased foliage intake to a greater extent than larger species (White-bellied Duiker Cephalophus leucogaster and Bay Duiker C. dorsalis). Fruits and seeds with high levels of tannins and indigestible fibre were preferred if they also contained readily digested simple carbohydrates and protein. Blue Duikers produce large quantities of saliva, suggesting a capacity to bind and limit the effect of some digestion inhibitors and toxins. High dietary diversity may prevent rumen fermentation from being overwhelmed by any specific plant toxin in the foods. Blue Duikers forage slowly and methodically over small areas, picking over the forest floor for individual food items; they browse standing vegetation only infrequently. In common with other duikers, they locate and follow groups of colobus monkeys and guenons, as they pass over their home-range. The duikers feed from fruits, leaves and other plant parts that fall beneath the primates. Activity peaks in the early morning and late afternoon with relative inactivity in the middle of the day and at night (Bowland & Perrin 1995, Wilson 2001). Blue Duikers are variably active at night, possibly on a seasonal basis (Bowman & Plowman 2006).

Social and Reproductive Behaviour  Pairs or small groups live permanently on small home-ranges (Dubost 1983, Bowland & Perrin 1995). Territories may be grouped in loose aggregations and animals appear to form a dispersed social unit possibly composed of related individuals. Dubost (1980) found that territories on a 74 ha study area in Gabon with a very high density of Blue Duikers were exclusive and averaged 2.5–4 ha. He noted that discontinuities in the terrain, such as large fallen trunks, small streams (wet or dry) and the pathways of larger animals served to demarcate boundaries. At the heart of each territory was a regularly inhabited zone or core area from which brief excursions were made, mainly to feed on temporary food sources. Core home-ranges of radio-collared Blue Duikers in the Ituri Forest ranged from 2.6 to 11.9 ha and were similar in size for both ?? (mean = 6.2 ha, n = 10) and // (mean = 6.0 ha, n = 9), which is not dissimilar from a hunted population in Congo (Mockrin 2010). Males and // in adjacent home-ranges spent over one-third of their time together. Subadults of both sexes dispersed away from the maternal home-range. Dispersal distances were 1 km in a monitored population in mono-dominant Gilbertiodendron dewevrei forest (Mockrin 2010). Neighbouring radio-collared individuals sometimes aggregated into temporary groups of four to five animals, in particular when pursued by hunters. Of 11 ?? and 12 // monitored by Dubost (1980), only one, a subadult /, changed her mate, indicating that couples generally tend to be stable and maintain permanent pair-bonds. He found that young // left the parental home-range at age 1–1.5 years old whereas ?? tended to stay on until they were nearly two years old and fully mature, at which point they appeared to depart voluntarily. In spite of apparent tolerance within families, Dubost found that resident ?? were intolerant of other fully adult ?? on their territories and captured adults of either sex were extremely aggressive to any adults placed in the same pen. Dubost found that the ? of a mated pair greatly reduced the time spent with his mate immediately after parturition and he even observed one or two unknown subadult ?? keeping company with postpartum //. Blue Duikers of both sexes mark tree trunks, branches and other objects in their territories with their preorbital glands. Dung may be somewhat aggregated in a home-range, but they do not create dung middens. Adult ?? and // and young of both sexes greet, groom and mark each other by touching their facial glands upon contact (Dubost 1980, Bowland 1990, Wilson 2001). Blue Duikers make a highly distinctive ‘sneeze-whistle’, which is commonly uttered when the animal has been disturbed and is running away. Populations of hunted duikers whistle more than those that are less hunted, the latter often moving away without signalling (Croes et al. 2007). The whistle is mimicked by hunters, and actually serves to attract animals, suggesting that this species-specific loud call is a generalized auditory advertisement (Kingdon 1982). A less distinctive and relatively quiet snort signifies a lower level of excitement. Individuals apparently seeking contact with another animal utter a soft groan, especially ?? in pursuit of // (Dubost 1983). Animals also stamp their forefeet, which may release secretions from pedal glands between the hooves, and ?? grate their horns on plant stems. Courtship is initiated by the ? who follows the / closely and practises laufschlag (leg tapping) when the / permits it. Females that

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Philantomba monticola

Blue Duiker Philantomba monticola.

are not in oestrus tend to evade the ? and keep their tail down. If pursuit of an unreceptive / is sustained she may even utter a ‘sneezewhistle’ (Dubost 1983). Animals bleat in distress and are said to make cat-like yowls.After birth the young does not follow its mother but keeps hidden until it is about three months old. The temporary absenteeism of ?? after the birth of their mate’s lamb has been interpreted as a mechanism to reduce risks of predation on the young (Kingdon 1982). Reproduction and Population Structure  The gestation period has been variously estimated at 4–7 months, with estimates based in part on figures for the similar congeneric Maxwell’s Duiker. Aeschlimann (1963) gives a gestation period of four months or about 120 days. Mentis (1972) reports a figure of 167 days, a figure similar to the 188 days recorded by Kadjo (2000). Dittrich (1972) reports 205 days, the same figure cited by von Böhner et al. (1984) and Wilson (2001). The latter author notes that he determined gestation based on 19 known instances where copulation was observed and the day of birth recorded. An analysis of foetal growth rates of Blue Duiker in the Ituri Forest supports a gestation period of 120–150 days, with a lower likelihood of a gestation period over 200 days (J.A. Hart pers. obs.). Shorter gestation periods would also be consistent with the relatively high fecundity observed in a number of populations. Animal keepers at Epulu Okapi Station who maintained a colony of Blue Duikers over a number years also report (unpublished) that gestation is about four months. New observations, possibly supported by hormone studies, will be required to clarify gestation period for this species. Von Böhner et al. (1984) reported that, in captivity, ?? are sexually mature at nine months, and // at 6–17 months; Hofmann et al. (1998) recorded first oestrus in // of a heavily hunted wild population at six months.Wilson (2001) notes that female Blue Duikers usually become sexually mature before 13 months, and that the earliest a / became sexually mature was eight months. He notes that ?? become sexually mature much later than //, from 11–14 months. Births occur throughout the year (Brand 1963, von Ketelhodt 1973, 1977a, Dubost & Feer 1992, Wilson 2001) with lowered birth rates around the dry season. A single young is born. Fullterm foetuses in the Ituri had mean weight of 726 g (range 590– 950, n = 13) or 13.2% of mean maternal body weight. There is a postpartum oestrus of 3–5 days following birth (OIA 1991). Wilson

(2001) reports an inter-birth interval of 202–248 days (n = 32 captive births), while von Ketelhodt (1977a) reports a mean interbirth interval of 266 days in captive animals. In a multi-year project inspecting animals killed by local hunters in the Ituri Forest, 78.6% of 196 adult // presumed to be multiparous were pregnant and 28% were both pregnant and lactating, the highest percentage of the six species of duikers occurring at the site (other species ranged from 0 to 25% for adult // both pregnant and lactating). Composition of Blue Duiker milk is reported on by Taylor et al. (1990) and von Ketelhodt (1976b).Young are reported to be weaned in five months. Captive animals have lived for 16 years (Weigl 2005). Using mark-and-release in an area of undisturbed rainforest (near Makokou, NE Gabon), Dubost (1980) determined that 62% of the population were reproductively mature animals (of even sex ratio) with 38% immature, i.e. less than one year old. Over one-third of young died in their first year of life, whereas adult annual mortality rates ranged from 7.3 to 10.4% (and see Hart 2000). In the Ituri Forest, sex ratios in a heavily hunted population were even but were skewed toward ?? (1.28 : 1 and 1.22 : 1) in two unhunted populations. Pre-reproductive individuals represented 33% of an unhunted population in mixed forest, but less than 18% of an unhunted population in mono-dominant Gilbertiodendron dewevrei forest. In the latter forest type, duikers also had lower mortality and higher rates of emigration (Hart 2000). In unhunted populations, female Blue Duikers reach maturity at about the same time that their third molar (M3) erupts. Females in hunted populations mature earlier, with 20–45% of subadults with the second molar only erupted with full udder development or pregnancy. Less than 5% of // with the second molar erupted were sexually mature in the unhunted populations (Hart 2000). Predators, Parasites and Diseases  Common predators in­ clude the African Rock Python Python sebae, the Crowned Eagle Stephanoaetus coronatus and the Leopard Panthera pardus (Wilson 2001). Wilson (2001) reported Crowned Eagles being a major predator in NE Zambia, and several other studies of the diet of Crowned Eagles have recorded Blue Duikers as being important prey items (Jarvis et al. 1980, Boshoff et al. 1994; and see Vernon 2001); predation by these raptors is also reported in Uganda and Tanzania (J. Kingdon pers. obs.). Blue Duikers represented less than 5% of eagle prey in one survey in the Kibale Forest, in an area with a very high primate biomass, and where primates were the dominant eagle prey (Mitani et al. 2001). Leopards and African Golden Cats Profelis aurata are primary predators in the Ituri Forest and Blue Duikers were the single most abundant prey species in felid diets. However, they were not killed disproportionately in relation to their abundance, and were not the most selected prey (Hart, J. A. et al. 1996). Annual predation rates of 2.3 kills per km2 and 1.1 kills per km2 were recorded over a four-year study of radio-collared animals monitored in mixed and mono-dominant forest, respectively (Hart 2000). Annual per capita predation rates of Blue Duikers were 14–34% those of the larger duikers, which are selected prey of Leopards. In Tsitsikamma N. P., Hanekom & Wilson (1991) reported Blue Duiker remains found in 27% of Leopard scats and 7% of Caracal Caracal caracal scats. In a health evaluation of 95 individuals of five species of duikers (including 37 Blue Duikers) in the Ituri Forest, all the animals examined were healthy and in physically good condition. Faecal parasite ova 233

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Family Bovidae

were detected in about one-third of animals examined, comparable with frequencies in most other duiker species. Serologic titres for bluetongue, epizootic heamorrhagic disease, infectious bovine rhinotracheitis and leptospirosis were recorded, indicating widespread exposure to these diseases (Karesh et al. 1995). Round (1968) provided a checklist of helminth parasites recorded from Blue Duiker at the time, including Haemonchus lawrencei, Oesophagostomum eurycephalon, Setaria caelum, S. dipetalamotoides, S. labiotapapillosa, Acuaria dartevelli, Moniezia expansa and Stilesia hepatica. Jooste (1984) reported additional parasites from three animals taken in E Zimbabwe, including Cooperia chabardi, Trichostrongylus axei, which Boomker et al. (1986) also recorded from a Blue Duiker from Tsitsikamma N. P. Boomker et al. (1991b) reported on helminth parasites of three Blue Duikers from KwaZulu–Natal, including: Taenia hydatigena, Gonglynema sp. and several Trichostrongylus spp. The only record of an arthropod parasite in Blue Duiker is that of a Rhipicephalus sp. recorded by Karesh et al. (1995). Conservation  IUCN Category: Least Concern. CITES: Appendix II. Arguably the most important wild ungulate economically and ecologically in Africa, the Blue Duiker is one of the most frequent species reported in bushmeat surveys across the continent (Wilson 2001) and withstands hunting pressure better than most of the larger duikers (Hart 2000,Waltert et al. 2006,Van Vliet et al. 2007). Unlike many of the other forest duikers, Blue Duikers tolerate and even thrive in a range of human-modified habitats, even in the vicinity of settlement, and often persist well in small habitat patches. However, some South African populations may be vulnerable to isolation and local extinction due to habitat fragmentation (Lawes et al. 2000), while the long-term decline since the mid-1970s of at least one Blue Duiker population in the coastal Cape forests is attributed to decreasing habitat quality associated with raising ambient temperatures brought on by climate change (Seydack et al. 1998). In East Africa, Blue Duiker are now relatively rare in the Eastern Arc Mts, being apparently absent from Nguru, Ukaguru, Rubeho and Pare. In the Udzungwa Mts, it has a very localized occurrence, but it is generally absent from lowland forests, such as Matundu. In some places this may be due to poaching, but competition with very abundant Sunis might be another factor in localities where trapping pressure is still light (F. Rovero pers. comm.). None the less, Blue Duikers occur in large and stable numbers in many protected areas across the bulk of the species’ range (East 1999). Blue Duikers adapt well to captivity (Bowman & Plowman 2006) and have been considered as potential candidates for domestication and for use as a ruminant model in research (OIA 1991). However, singleton births and high aggression levels between adult ?? limit their potential for husbandry.

Like other duikers, the Blue Duiker is largely dependent on the rain of plant parts falling from the canopy. This derives, in large part, from the activities of arboreal mammals such as primates and fruitbats, and of birds. How the elimination, or depletion, of arboreal animals might affect duikers is unknown. The difficulties of inferring effects of hunting on duikers are compounded because heavy hunting hits both arboreal and terrestrial fauna and the details of interactions between arboreal and terrestrial communities are seldom taken into account. Measurements Philantomba monticola TL (??): 715 (627–731) mm, n = 10 TL (//): 700 (645–727) mm, n = 10 T (??): 75 (64–83) mm, n = 10 T (//): 78 (61–81) mm, n = 10 HF c.u. (??): 170 (156–190) mm, n = 10 HF c.u. (//): 165 (150–185) mm, n = 10 E (??): 55 (52–60) mm, n = 10 E (//): 56 (53–61) mm, n = 10 Sh. ht (??): 340 (330–360) mm, n = 10 Sh. ht (//): 350 (320–370) mm, n = 10 WT (??): 4.5 (4.0–5.3) kg, n = 10 WT (//): 5.4 (3.8–6.1) kg, n = 10 KwaZulu–Natal, South Africa (Wilson 2001) TL (??): 675 (630–700) mm, n = 10 TL (//): 680 (650–710) mm, n = 8 T (??): 88 (79–91) mm, n = 10 T (//): 88 (83–95) mm, n = 8 HF c.u. (??): 175 (165–180) mm, n = 10 HF c.u. (//): 171 (170–185) mm, n = 8 E (??): 56 (50–59) mm, n = 10 E (//): 55 (50–60) mm, n = 8 Sh. ht (??): 340 (320–350) mm, n = 10 Sh. ht (//): 340 (330–350) mm, n = 8 WT (??): 4.8 (3.9–5.4) kg, n = 10 WT (//): 5.3 (4.0–6.5) kg, n = 8 Chirinda Forest, Zimbabwe (Wilson 2001) Maximum recorded horn length is 7.3 cm for a pair of horns from Irumu in DR Congo (Rowland Ward) Key References  Bowland 1990; Dubost 1980, 1983; Hart 1985, 2000; Kingdon 1982; Mockrin 2010; Wilson 2001. John A. Hart & Jonathan Kingdon

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Sylvicapra grimmia

Genus Sylvicapra Common Duiker Sylvicapra Ogilby, 1837. Proc. Zool. Soc. Lond. 1836: 138 [1837].

These are the ‘bush duikers’, which appear to have reversed many of the deep-forest adaptations of the Cephalophini as a whole, such as the wedge-shaped form and the swept-back horns. They are slenderly built, with long legs (particularly the distal segments) and have a longer neck than other duikers. With Philantomba, this genus shares the elongated ischium, the evenly haired (untufted) tail, the somewhat curved horns, the somewhat tubular orbits, and the oblique occipital plane and paraoccipital processes. On the other hand, the hairs of the pelage are agouti-banded, and the hairs on the forehead are restricted to a coronal tuft, as in Cephalophus. The horns, which are largely restricted to ??, have more of an upright direction than in other duikers, and the coronal tuft is dominated by forwardly directed hairs. The dorsal profile of the

skull is straight, rather than convex as in most other duikers, and the nasal opening is enlarged. According to Jansen van Vuuren & Robinson (2001), this genus is not, as has been previously supposed, the most phylogenetically distinct of the duikers (that position belongs to Philantomba). Further, recent molecular evidence reveals that Cephalophus is paraphyletic with respect to Sylvicapra (Hassanin et al. 2012). Normally, a single species is recognized in this genus, but two or more could prove to be valid; for example, Grubb & Groves (2001) suggested that the taxon coronata, from far West Africa, may be a distinct species. Colin Groves

Sylvicapra grimmia  Common Duiker (Grey Duiker, Bush Duiker, Grimm’s Duiker) Fr. Céphalophe de Grimm (Céphalophe couronne, Céphalophe du Cap); Ger. Kronenducker. Sylvicapra grimmia (Linnaeus, 1758). Syst. Nat., 10th edn, 1: 70. ‘Habitat in Africa’; based on a specimen seen by Grimm in the fort at Cape Town (Thomas 1911: 153) so now known to be South Africa, Western Cape Prov., Cape Town.

that he thought corresponded well with general biogeographic patterns, while allowing for high-altitude isolates of lowland parent populations as well as supposed hybrid swarms from ‘overlap zones’. Meester et al. (1986) listed six subspecies for the southern African subregion. Grubb & Groves (2001) recognized 14 subspecies (including one undescribed form from Mt Kilimanjaro). Wilson (2001), who examined thousands of skulls and skins in many museums and dozens of live animals throughout the species’ range, concurred with this treatment, which is followed here. As the distribution of S. grimmia is continuous in sub-Saharan Africa and some forms show individual variation, there are obviously substantial intergradations between different subspecies or regional populations and it is currently impossible to accurately delineate boundaries between the subspecies listed under Geographic Variation. Synonyms: Grubb (2005) lists 38 synonyms. Chromosome number: 2n = 60; the X chromosome is a metacentric (Robinson et al. 1996b).

Common Duiker Sylvicapra grimmia.

Taxonomy  Haltenorth (1963) recognized 19 subspecies of Sylvicapra grimmia and these, with some changes, were accepted by Ansell (1972). However, Ansell indicated that the validity and distributional limits of many were doubtful. Kingdon (1982, 1997) broke down more than 30 named forms into eight regional groupings

Description  The Common Duiker is a medium-sized animal with a body colouration that varies from light grey in Botswana through many ranges of grey, yellow-ochre to almost red on the Nyika Plateau in Malawi to specimens in West Africa that are grey on the hindquarters to rufous coloured on the shoulders. In Malawi, Mozambique, E Zambia and even parts of Zimbabwe a distinct white ring occurs around the eyes (hence the name orbicularis for the subspecies occurring in that region). The colour of the ears varies, with some animals having a great deal of white inside the ears and others no white at all. The ears are comparatively long and pointed. A long tuft of hair occurs on the head and in many cases it is very long and pointed and in other animals short and ‘square-topped’. Colour of the frontal tuft varies from black to grey and even rufous. A dark-coloured frontal blaze is found in almost every animal, but 235

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occasionally no dark band occurs at all (especially in animals from East Africa). This frontal blaze varies in colour from light brown to pitch-black and occurs from the rhinarium to the front of the eyes and occasionally even beyond to the frontal crest. Underparts are snow-white in some populations to greyish-brown in others. Many Zimbabwean and Zambian specimens have snow-white bellies. Body hairs are fairly short in most animals, but several specimens examined in Botswana had hair that were, in places, in excess of 6 cm long; in the central Kalahari of Botswana, both long- and short-haired duikers can be found in the same population (Wilson 2001). Some isolated montane populations have long hair, such as those on Mt Kilimanjaro and Mt Elgon, likely an adaptation to the environment. The legs are generally the same colour as the body and the fetlocks brownishblack to pitch-black, the black colour often extending up the front of the legs to the body. The tail underneath is white and reddish-brown to black above, but again this varies with animals even in a small area. Albinos, characterized by white fur and pink eyes, are also recorded (Wilson 2001). Preorbital glands are present, and consist externally of a row of pores exuding from a bare streak of skin. The length of this streak varies, being much shorter in specimens from Ghana than those from Botswana (Wilson 2001). The secretion consists of a thick black melanincontaining component and a thin, yellowish clear liquid.The secretion is heterogeneous, comprising water, mucus, a thin yellowish oil and large concentrations of heavy waxy material in varying proportions. Burger et al. (1990) isolated 33 constituents of this preorbital gland secretion. Pedal glands are also present in all four feet, those on the forefeet being deeper and larger, and contain a creamy-white secretion. Inguinal glands are also present, as much as 65 mm long, and contain large quantities of a white to creamy secretion with a powerful odour (Wilson 2001). The size of the hooves depends a great deal on the environment and soil on which the animal lives.Those on Kalahari sand have very long hooves while those on the Nyika Plateau are short and rounded. Lateral (false) hooves are present in all animals. Females are generally larger than ??. Males have a pair of stout pointed horns; occasionally // have been recorded with horns (Miller 1912, Shortridge 1934, Ansell 1960b, Wilson & Clarke 1962). Shortridge (1934) considered that 10% of all Common Duiker // in Namibia have horns. Horned // appear to be more frequent in some localities than others (Wilson 2001).Wilson (2001) found that in E Zambia, in a sample of 980 female skulls examined, 0.6% had horns. In Zimbabwe, of 720 female skulls, 3.8% had horns (Sebungwe area) and in Botswana in a sample of only 55 female skulls, 12.7% had horns. In all cases horns appearing in // were short, thin and often stunted. The skull has a straight profile, enlarged nasal opening, and somewhat tubular orbits. The median palatal notch is very broad, and V-shaped (Grubb & Groves 2001). The dentition of the Common Duiker is diphyodont and similar to most other bovids. Riney & Child (1960) established ageing criteria for the Common Duiker based on a series of skulls with known collection dates supplemented by a few captive ‘known-age’ animals ranging from a neonate to one individual aged 21.5 years. This was revised by Wilson et al. (1984) after working with a very large collection of more than 90 knownage skulls (and see Wilson 2001). As with some other ungulates the Common Duiker has a modified lower canine tooth that has become the last incisiform.The first premolar has also been lost in the process

Lateral view of skull of Common Duiker Sylvicapra grimmia.

of evolution. At birth the Common Duiker has either partially or fully erupted lower milk canines and incisors with erupting upper and lower premolars; interestingly, P3 was usually the first premolar to be fully erupted and not P2 as one would expect. At the age of 26 months both mandibular and maxillary permanent molariform teeth are fully erupted and in wear. There is very little variation in the age of eruption and replacement of all molariform teeth making it a very useful feature for age determination (see Wilson 2001: Table 16). However, there is considerable difficulty in distinguishing deciduous incisiform teeth from permanent ones because of the great variability in eruption. The Common Duiker is one of the few ungulates in which the canines and incisors are replaced last and long after the eruption of the third molar. The final eruption and replacement of the canines and incisors usually takes place at between 30 and 38 months. Dental abnormalities, including, for example, having one or two upper canines present or the lower second premolar absent, are discussed by Riney & Child (1960) and Wilson (2001). Geographic Variation  As already mentioned above, the distribution of the Common Duiker is continuous in sub-Saharan Africa, and therefore it is currently impossible to delineate exactly where one subspecies starts and another takes its place. However, several of the subspecies do have clear distinguishing characteristics and some of these features are discussed below. There is considerable colour and size variation in this species even within very limited areas. S. g. grimmia: in the extreme south of the continent this subspecies is a great deal greyer than any other subspecies (it is known as the Grey Duiker in the Cape). It is not in any way tawny-coloured, and there is no white on the belly, but there are traces of white on the throat and on the upper half of the inner side of the legs. Roberts (1951) described medium development of speckling and an incomplete face-blaze, which does not extend to the frontal tuft on the head. S. g. caffra: occurs north of S. g. grimmia and common in KwaZulu– Natal and Limpopo Provinces of South Africa, S Mozambique (Coguno) and E Zimbabwe. It is speckled, fawn-brown to greyyellow in colour with the mid-dorsal zone being much darker. The face-blaze is nearly black and well marked up to the eyes. Some white occurs on the belly, but the underside is mainly light buff. The tail is nearly all black on the upper surface and white below (Grubb & Groves 2001). Intergrades between S. g. grimmia and S. g. caffra occur in the ‘Caffraria’ (Algoa Bay area). S. g. steinhardti: occurs throughout Namibia; south as far as Port Nolloth in the Northern Cape of South Africa; north to Namburi and Cahama in Angola, and east to Kazungula and Chobe district in Botswana. This subspecies is very pale sandy-coloured or even fawn. Very little speckling. In this subspecies the face-blaze fails

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to reach even the level of the eyes. The undersides are off-white, of the forest belt at 1980 m and the two subspecies are isolated chest and throat buff. from one another by montane forest. It is much greyer than hindei S. g. splendidula: the subspecies occurring in Angola, from Mupa with heavier speckling, more marked face-stripe, black pasterns district north to Kunungu, DR Congo, and across the Congo R. to and drabber tone on underparts. It has long, thick fur (Grubb & Odzala (Congo). It also occurs in the south-east of DR Congo to Groves 2001). the southern end of L.Tanganyika, north-east to Uvira and in Zambia S. g. lobeliarum: Afroalpine zone on Mt Elgon. Has a pinkish-grey coat west to the Luangwa Valley and north as far as Mpika. It is also with heavy speckling and a wide face-stripe, which is chocolatefound in W Zimbabwe (Grubb & Groves 2001). This subspecies is brown. The ears are short and pointed and the pasterns black. In the largest of all the common duikers. It has a light reddish-ochre most of its characters this race seems to be the most distinctive of colour and a short face-stripe. Specimens from Hwange N. P. in the afroalpine forms. Zimbabwe are intergrades between S. g. steinhardti and S. g. S. grimmia subsp. nov.: this is the duiker described by King (1975) splendidula. The Latin name (splendidula), meaning ‘splendid animal’, from Mt Kilimanjaro. This distinct form has a grey-brown colour, is appropriate, given its bright, contrasting colour, unparalleled in thick black face-blaze, reduced black on tail, and off-white any other form of Sylvicapra. underparts not extending down the inner sides of the limbs. It differs from altivallis in having heavier speckling, less black on tail, foreleg stripe usually reaching the knee, ‘squared off’ ears and a shorter tail (Grubb & Groves 2001). S. g. madoqua: Ethiopian Highlands, both east and west of the Rift Valley, as high as 2990 m in the Sahatu Mts and as low as 610 m at Hawash Station. It is generally grey-buff to brownish-ochre with heavy speckling and the mid-dorsal zone darkened. The face-stripe is complete in the highland form, less so in those from the lowlands. According to Grubb & Groves (2001) this is a very variable race, difficult to define although the various component samples do differ as a whole from the Kenya races and from campbelliae. S. g. campbelliae: savanna country from Burundi and Karagwe in the south-east to north of Bahl-el-Ghazal; also occurs in the northeast of DR Congo, southernmost Central African Republic, S Chad, N Cameroon and Nigeria, west to Ghana, Burkina Faso (Fada-N-Gourma) and Sierra Leone. This is a dull ochre to greyCommon Duiker Sylvicapra grimmia orbicularis crouching. buff duiker, heavily speckled and dark on the mid-dorsal zone. It has a dark face-blaze, which extends to the crest, the tail black S. g. orbicularis: occurs north of the Zambezi R. in Mozambique, above and brown pasterns. The foreleg stripe is well marked. The Malawi and in Zambia east of the Luangwa R. It is also present belly is buff-white to white and no throat patch. throughout Tanzania (perhaps absent from the extreme far north- S. g. pallidior: distribution of this subspecies is not accurately known, west), along the Kenya coast and inland to Voi and the Juba R. in but it does occur in the Sahel Zone from Mani (furthest west Somalia. This beautiful subspecies has a distinct white ring around locality) as far east as Gallabet on the borders of the Ethiopian the eyes in specimens from E Zambia and much of Malawi. The Highlands; north as far as Jebel Marra Darfur; south as far as general body colour is light fawn with a narrow face-blaze, the tail Fort Archambault. S. g. pallidor is much paler than the previous is black and so are the pasterns. Females are considerably larger race, it is pale buff and weakly speckled but still has a mid-dorsal than ??. The controversial form walkeri is a melanistic example darkening. The face-blaze extends to the crest. of S. grimmia (Grubb 1988) and can be assigned to S. g. orbicularis. S. g. coronata: this subspecies has only been recorded from a small area S. g hindei: Kenya highlands, east of the Great Rift Valley, as far east as on the borders of Guinea, Guinea-Bissau, Senegal and Gambia. It Machakos and Sultan Hamud and south as far as Moshi in Tanzania. is a beautiful bright orange-yellow colour with no speckling and a This subspecies has much speckling along the mid-dorsal region mid-dorsal darker shade, which is almost reddish.The face-blaze is on a generally ochre-coloured body. The face-blaze is bold, broad red and not extending to the crest. The foreleg stripe is indistinct and dark and extends up to the frontal tuft of hair on the head. and not reaching the knee. It has yellowish white underparts and The belly is white, throat buff with no white line. This is a small no throat streak. This is the most distinctive of all the subspecies subspecies, but has a broad skull. and could possibly be raised to specific rank depending on its S. g. nyansae: in Kenya, west of the Rift Valley west to Busoga district relationship with the neighbouring campbelliae (Grubb & Groves in Uganda and to the eastern levee of the Nile in S Sudan north to 2001). Reseires in Ethiopia on the Blue Nile. This subspecies is brighter than S. g. hindei, but with less mid-dorsal darkening and much whiter Similar Species underparts. The face-blaze is deep brown rather than blackish. Ourebia ourebi. Occurs on open grassland and could be confused with S. g. altivallis: Afroalpine zone of the Aberdares and also probably the Common Duiker in places where both species occur. Much on Mt Kenya. On Mt Kenya, S. g. hindei reaches up to the edge taller, with longer horns, and a much longer neck. Oribi normally 237

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occur in small family groups, whereas the Common Duiker is a solitary animal.

E and SE Gabon, where little is known of the species’ distribution (East 1999, Wilson 2001).

The Common Duiker is the only duiker species found in open savanna woodlands and, therefore, unlikely to be confused with any of the other duikers (although, to a lesser extent, the Red-flanked Duiker Cephalophus rufilatus occurs in some open country in West Africa in the Guinea Savanna zone).

Habitat  Although the Common Duiker is typically a savanna woodland species, it is often found in relatively open country and even extends into the alpine zone in some mountainous areas such as on Mt Kenya (to 4900 m;Young & Evans 1993) and Mt Kilimanjaro (to 5600 m; see King 1975, Grimshaw et al. 1995), the highland plateau to 3300 m in the mountains of Arssi and Bale in Ethiopia (Yalden et al. 1996), and at altitudes above 2200 m in the Drakensberg Mts of South Africa (Vincent 1962, Rowe-Rowe 1994). Although the species is generally associated with rather open country and woodland it nevertheless still requires at least some small patches of bush in which to hide. This is well demonstrated on the open montane grasslands on the Nyika Plateau in N Malawi and NE Zambia. In this area the species is often seen on the open grasslands but once disturbed it rushes into the edges of the montane forests to hide. Wilson (2001) identified six vegetational types as habitats for the Common Duiker: Brachystegia–Isoberlinia woodland – one of the preferred habitat types; Mopane Colophospermum mopane woodland – while the Common Duiker is not plentiful in this habitat type it nevertheless can still be found in many places where Mopane woodland occurs; Montane Grasslands – the species is common in this habitat and especially on the border of such grasslands where Uapaca and stunted Brachystegia woodland is found; Kalahari/Acacia veld – the species is common in these dry areas where it can go without drinking water for months at a time; Guinea Savanna – as with the southern African savanna woodlands, the species is plentiful in the northern Guinea savannas; and Terminalia woodlands, on Kalahari sand – a much-favoured habitat. In reality, the Common Duiker can be found in almost every habitat type in sub-Saharan Africa, with the exception of dense evergreen rainforests, and in more open areas as long as sufficient cover exists in which it can hide.

Distribution  The Common Duiker is one of the most widely distributed antelopes on the African continent, and in spite of dense human populations in many areas its historical distribution has remained largely unchanged, managing to survive where many other species of large mammals have been exterminated. In West Africa, the Common Duiker is widespread and common in savanna woodlands from C and S Senegal, Gambia and Guinea-Bissau east through N Guinea, SW Mali, N Côte d’Ivoire, S and C Burkina Faso, N Ghana, Togo, N and C Benin, N and C Nigeria, SW Niger and N Cameroon. They are naturally absent from the Guinean rainforests, being entirely absent from Liberia; however, there is one old record from NE Sierra Leone (Stanley & Hodgson 1929; and see Grubb et al. 1998). From Cameroon, their distribution extends through S Chad, Central African Republic and S Sudan to north-east Africa. Here, while still widespread and common in many areas in W and C Ethiopia, they are now rare and hardly ever seen within their former range in S Somalia and SC Eritrea; they are recorded only from the Forêt du Day in Djibouti, but there have been no confirmed records in several decades (Künzel et al. 2000, Heckel & Rayaleh 2008). In East Africa, the Common Duiker occurs throughout Kenya (except semi-arid and arid northern rangelands from which it is naturally absent), Tanzania, Uganda, Rwanda and Burundi, west into N, E and S DR Congo.They then occur southwards widely in all countries, with the exception of the arid coastal deserts of Namibia (East 1999,Wilson 2001). In central Africa, the Common Duiker occurs in the savannas of C and SE Congo, and in neighbouring

Sylvicapra grimmia

Abundance  The Common Duiker can be found in very large numbers in many places in Africa. In several places where game elimination as a means of controlling tsetse flies has been carried out, tens of thousands of individuals have been shot and yet today the species is still very common in those places. For example, 83,784 duikers were shot in Zimbabwe between 1924 and 1945 (White 1954), and yet the species is still very common in these areas today. Similar operations were undertaken in the Chipata district of E Zambia (see Wilson & Clarke 1962 and Wilson & Roth 1967 for details), yet they still occur widely in this part of the country. Wilson (2001) summarizes some recorded densities of Common Duikers from various localities in Africa in different vegetation types using line transects, including: 3.2/km2 in the Guinea savanna of Mole N. P. in N Ghana; 2.9/km2 in Brachystegia–Isoberlinia woodland in Lukuzuzi G. R. in E Zambia; 2.0/km2 in Acacia/Combretum/Grewia savanna in the south-west lowveld of Zimbabwe; 2.4/km2 in the Matopo Hills of Zimbabwe; and 9.0/km2 in the Baikiaea woodland of Hwange N. P. East (1999) indicates that aerial surveys generally produce estimates of population densities in the range of 0.01–0.15/km2, such as in W N. P. (Burkina Faso) (Belemsobgo & Chardonnet 1996), and Pendjari (Benin) (Chardonnet 1995). However, East (1999) notes that aerial surveys underestimate the numbers of this small, secretive

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species by a large but unknown factor. Ground surveys in areas where the species is common often produce density estimates of the order 0.3–1.7/km2. As the Common Duiker is a species that is capable of altering its periods of activity depending on disturbances caused by local conditions it becomes extremely difficult to assess the size of populations accurately.Therefore, ground surveys are often inaccurate if undertaken at a time when duikers are not active. The very best times of the day for surveys are very early in the morning soon after sunrise and late in the afternoon just before sunset and early evening. East (1999) estimates the total population for sub-Saharan Africa as 1,660,000, indicating that this is a conservative figure. The present author believes the total population of the Common Duiker is more likely to be in the order of 10 million animals. Adaptations  This duiker is a most adaptable species and capable of occupying a wide range of habitat types. Apart from the Rwenzori Red Duiker Cephalophus rubidus on the Rwenzori Mts, the Common Duiker appears to be the only species of African antelope able to adapt to habitats high up on the mountains, often to the snowline. It also occurs in the hot and seasonally dry valleys of the Zambezi and Limpopo valleys and is very common in the waterless and very dry Kalahari Desert in Botswana and Namibia. An experiment carried out on a captive duiker in Zambia indicated that one animal went without water for 40 days (Wilson 1966a). Dietary flexibility, the ability to survive without surface water and fecundity all contribute to this species’ success, but the basis of its adaptive flexibility is undoubtedly physiological, an aspect of the Common Duiker’s biology that awaits a comparative investigation. Studies of the preorbital gland secretion of male and female Common Duikers have shown that two compounds, both thiazole derivatives, are present in higher concentrations in ?? than in //. This is the only consistent difference between the secretions of the sexes, and coupled with the fact that only ?? mark territories, could be seen as evidence that these two compounds play a role as sex recognition cues and in territorial behaviour (Burger et al. 1988, 1990). There is a great increase in the size of the preorbital gland of the ? while the / is in oestrus and he will mark trees, shrubs and even stones far more vigorously than normal. The secretion, which is squeezed out by rubbing the gland against some hard object, crystallizes and forms a dark stain on the object marked. In E Zambia, and on several occasions in Zimbabwe, the author has observed a ? rubbing his preorbital glands on the hindquarters of a / in his enclosure, leaving the secretion on her. Sikes (1958) also observed a ? marking a / in Nigeria. During a detailed study of food and feeding habits in Matobo N. P. (W Zimbabwe), Wilson (2001) watched a duiker feeding under foraging monkeys: every now and then the duiker would look up into the trees to see where the monkeys were, seemingly waiting for them to drop more flowers to the ground. Such behaviour reveals Sylvicapra as a typical duiker and can be presumed to have derived from forest-dwelling ancestors. Foraging and Food  The Common Duiker is a browser and frugivore, rarely taking grass, and has been recorded feeding on a very large number of plants. In one study, in E Zambia, the leaves of 44 different plants were identified from the rumens of 191 duikers. In addition, from the same sample 33 different fruits and seeds were recorded as well as 15 species of flowers (Wilson 1966a).

A detailed study of the food and feeding habits of the Common Duiker was carried out in Matobo N. P. (see Wilson 2001).Tame male and female duikers were used for the study in granite hills where the vegetation consisted of Colophospermum mopane in addition to Brachystegia spp. and Julbernardia globilfora, Parinari curatellifolia, Terminalia sericea and Peltophorum africanum were also common. Contrary to what was expected, green leaves of large trees and shrubs were not the most important foods eaten by the duikers studied. Most of the leaves of the larger trees are well out of reach unless the trees occurred in a shrub form or as seedlings. Fallen leaves of some of the large trees were often gleaned from the ground even though they were partly dry, as in the case of Kirkiana acuminata, which were eaten in large amounts during the dry season. Instead, leaves, stems, flowers and fruits of many small herbs were eaten in very large quantities, for example, Commelina welwitschii, C. benghalensis, Ipomoea omaneyi and Sida cordifolia. In this study 65% of the duikers’ total feeding time was spent feeding on these plants. The duikers often climbed among the boulders at the base of rocky outcrops in the study area and both animals fed on herbs growing in the litter between the boulders. The duikers often pulled entire small plants out of the soft soil among the boulders and ate the entire plant, including the roots. After herbs, fruits and seeds were most important in the diet. The fruits, seeds and flowers of many species of trees and shrubs were eaten, when available, in very large quantities. The fallen dry pods and also green pods of Acacia robusta and A. nilotica were eaten in fair numbers. The following fruits were eaten whenever they were available and formed a major part of the duikers’ diet: Ximenia caffra, Dovaylis caffra, Flueggea virosa, Flacourtia indica, Ziziphus mucronata, Grewia flavescens, Solanum sp. and Pseudolachnostylis maprouneafolia. Both green and dry pods (in bunches) of Dichrostachys cinerea were much sought after as were the ripe bunches of Lannea discolor. Various species of Ficus fruits were also greatly relished and in some instances eaten in very large quantities. Flueggea virosa was an important tree as the tiny white fruits were eaten, when available, in vast amounts. On numerous occasions the ? was seen to stand on his hindlegs in order to get to fruits and leaves of shrubs that were just out of reach. This was particularly noticeable when feeding on the ripe fruits of Ximenia caffra, Flacourtica indica, Grewia spp. and Flueggea virosa. The flowers of most species were eaten, some in large numbers, while in other species only a few were eaten. In nearly all cases the flowers had fallen from the larger trees and these were gleaned from the ground. The duikers were like vacuum cleaners as they moved about sucking up the flowers. Flowers eaten included: Bauhinia petersiana, Cassia abbreviata, Brachystegia spp., Hibiscus sp., Loranthus sp., Dichrostrachys cinerea and Acacia spp. In the case of the small legumes, Crotaloria podocarpa, C. sphaerocarpa, C. virgulata and Dolichos daltonii flowers were often plucked from the plants and eaten in bunches together with any green pods and leaves that were present on the plant. In one part of the male duiker’s home-range there were four Cassia abbreviata trees. Vervet Monkeys Chlorocebus pygerythrus were on two occasions seen feeding on the recently flushed yellow flowers and on both occasions the male duiker picked up some of the fallen flowers from the ground and ate them. In the Matobo Hills study, although the leaves of trees and shrubs were not the most important dietary item, the following were eaten: Grewia flavescens, recently flushed Parinari curatellifolia, Dichrostachys cinerea, Acacia nilotica, A. robusta, Combretum hereroense, Dombeya 239

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rotundifolia, Combretum apiculatum, Ximena caffra, Pavetta schumanniana, Terminalia brachystema, Flueggea virosa, Lannea edulis, Flacourtica indica, Ziziphus mucronata, Pseudolachnostylis maprouneifolia, Pterocarpus angolensis and newly flushed Protea angolensis. Albizia amara, Burkea africana, Euclea divinorum, Lannea discolor, Azana garekeana, Faurea saligna, Peltophorum africanum and Rhus tunuinervis were occasionally eaten but only in very small quantities. Dichrostachys cinerea, Flueggea virosa, Combretum hereroense, Flacourtia indica and Combretum apiculatum were some of the more important trees from which leaves were picked and eaten. The leaves and fruits of the small tree Securingea virosa were often eaten in very large quantities. Taking all three years of the study into account, only 25% of the total feeding time was spent feeding on the leaves of large trees. As soon as the wet season ended in late Mar the trees and other plants started drying out. This normally started with the small herbs and climbers and ended with the leaves of the large trees falling to the ground. By late Jul there were very few large trees with leaves still on them and the country looked extremely dry. It was at that time of the year that fruits were still falling and these became extremely important in the duiker’s diet. By Aug temperatures were already increasing and fires had swept through the countryside including the study area. With the increase in temperature after the winter, many trees and shrubs started flushing new leaves. By Sep and Oct, long before the rains fell in the middle of Nov, the vegetation in the entire study area, for each of the three years of the study, was attractively green and provided good supplies of food for browsers. With the commencement of the wet season in late Nov and early Dec, most of the small herbs and climbers flushed, and for the following five months there was an abundance of vegetation again. The onset of the dry season led to the drying out of the vegetation and both the male and female duikers had to cover a larger area in order to find food. A lot of time was spent slowly walking about sniffing at dry plants and looking for food. By Aug both duikers were ranging over an area almost twice the size of the area occupied during the wet period and both duikers appeared to have lost weight. Soon after the first rains had fallen the duikers fed on a small quantity of freshly sprouted grass yet, even though the grass was luxuriant, it did not represent even 2% of their diet. Interestingly, the rains brought an abundance of harvester termites. On seven occasions (four times with the / and three times with the ?) the duikers were seen feeding on termites emerging from holes in the ground. On one occasion the ? was seen to lick up at least 50 harvester termites Trinervitermes rhodesiensis as they emerged. On the other six occasions not more than between 10 and 20 termites were eaten. With the rains, fungi also became evident. Three species were eaten by the duikers, but only one or two plants at a time. After eating the top umbrella part of the fungus the duikers would occasionally also eat the entire stem. Once the / dug a plant out of the ground and tried to dislodge the remaining small parts of the stem that remained in the ground. For over eight minutes she dug with her front hooves into the soft ground, removing the remains of the fungus and picking up and eating all the small pieces. During the study, it was noticed that resin that exuded from the trunk of Acacia robusta and Acacia karroo trees were eaten by both the male and female duikers. They pulled soft balls of resin from the trees and these were chewed thoroughly before being swallowed. On one occasion the ? removed six balls of resin from one tree and

spent another 10 minutes at the tree licking the bark and removing small pieces of bark. Common Duikers will also feed on cultivated crops, including beans, sweet potatoes, tomatoes and groundnuts. Analysing faeces in Grants Valley, Eastern Cape, South Africa, Kigozi (2003) found that chicory provided more than one-third (35.6%) of the winter diet and a substantial proportion (14.4%) of the spring diet of the Common Duiker. The species is well known for its tendency to eat meat, and there are records of Common Duikers feeding on Helmeted Guineafowl Numida mitrata chicks, ducklings, Red-billed Queleas Quelea quelea, a Laughing Dove Stigmatopelia senegalensis, mopane caterpillars Conimbrasia belina, lizards and insects (Smithers 1971, Wilson 2001). Hofmann (1973) once saw a duiker take a striped mouse Rhabdomys sp. The Common Duiker is independent of water and even when water is available the animal rarely drinks (Skinner & Chimimba 2005). Social and Reproductive Behaviour  Common Duikers are generally solitary animals. However, when a / is in oestrus a ? and / will come together. They will often stay together for several days at a time and once mating has taken place they will separate again. Pairs will nevertheless still remain in the same general area and will come together from time to time, especially if a tree producing fallen fruit is in their home-range and in that case they will often be seen together.When two duikers are seen together it is usually an adult ? with an adult /, but a / with her young is another common pairing. When three duikers are seen together it is usually an adult pair and a young animal that has not yet been weaned. One of the most important factors influencing distribution, territorial behaviour and the home-range in the Common Duiker is food supply. The home-range is determined by the availability of the preferred foods at different times of the year. Therefore, the daily activity and home-range of the duiker is determined by its need to obtain sufficient food and this basic function is complemented by its drive to reproduce. As Allen-Rowlandson (1986) has pointed out, territorial behaviour that limits a species to any given area is likely to affect its performance. A number of authors have briefly mentioned that the Common Duiker does not often move very far and is restricted to a small area (Dixon 1964, Wilson 1966a, Dunbar & Dunbar 1979). In the Ethiopian Highlands, Dunbar & Dunbar (1979) found that known duikers occupied home-ranges that overlapped only to a limited extent. Allen-Rowlandson (1986) found that some adult ?? shared part of their home-ranges with //. He found that the mean size of the home-range of Common Duikers in KwaZulu– Natal was about 21 ha, which varied between 21.1 and 27.4 ha. In a study in E Zambia, Wilson (2001) found that one adult ? had a home-range of 13.2 ha and another a home-range of 15.7 ha. These two ?? had distinct home-ranges that did not overlap, while adult // had home-ranges of about 13 ha, which overlapped with each other and also with those of ??. Adult ?? that are sexually mature will defend their territory and drive out any competing adult ??. However, subadult ?? are tolerated and not driven away. The Common Duiker has a number of behavioural patterns that characterize courtship, including: the ? chasing the / when in oestrus; flehmen; constant flicking of the tongue; low-pitched bleats; laufschlag (leg tapping); false mounting attempts; biting of the female’s hindquarters and tail; and increased marking of objects with secretion from preorbital glands and final mounting of /. Although

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Sylvicapra grimmia

Common Duiker Sylvicapra grimmia.

these various forms of behaviour are connected with courtship, they do not necessarily follow each other sequentially and some adult ?? do not exhibit all the behavioural patterns. The chasing of the / in oestrus is often very pronounced and ?? in captivity can become most aggressive and will not tolerate other sexually mature ?? in the same enclosure. Very often a ? would chase a / in oestrus to near exhaustion and in captivity the / would often lay down in some secluded spot in order to get away from the ?. If that happened he invariably persisted in attempting to get her to her feet by pawing her with his front feet, or even biting her. If the / rose to her feet the chase would continue.This chasing behaviour was often observed in the wild early in the morning and again late in the afternoon and at sunset. In the male Common Duiker, flehmen is not as noticeable as it is in other ruminant species, but after close genital olfaction or after testing freshly voided urine from a / in oestrus, the ? would raise his head and a slight lip-curl could be seen. After genital olfaction, reproductive activity in the ? is increased considerably and this is also noticed if freshly voided urine is tested. It is therefore clear that olfaction is essential in the stimulation of reproductive activity in the ?. Frequent urination by the / is a common characteristic while she is in oestrus, which suggests maximum output of urine and, therefore, endrocine contents.

During courtship the ? follows the / persistently but only seriously begins to court her when she proves to be in oestrus. The flicking of the male’s tongue and low-pitched bleats are closely linked and can be clearly observed in ?? following a / in oestrus. The male’s tongue will also occasionally come in direct contact with the female’s hindquarters, her tail and even her vulva. Low-pitched bleating is a vocal communication between the adult ? and / during courtship. The / on the other hand also bleats from time to time but not nearly as much as the ? and usually only when running away. Another form of mating behaviour consists of the ? striking out with a stiff, straight forelimb (a behaviour known as laufschlag). He nudges the / either between the hindlegs or on her hindquarters. If the / moves slowly away the ? will follow and flehmen and laufschlag continue. However, once the / stands the ? attempts to mount her, and false mounting attempts by the ? are extremely common. In this behaviour the ? shows no serious attempt to clasp the / with his forelegs as he would do if she is receptive. When he rises onto his hindlegs they are usually too far behind her to complete copulation. As soon as he mounts, the / takes a couple of steps forward and he drops onto all fours again. Often in these false mounting attempts the / would raise one or both of her hindlegs and kicks the ?. Either of these activities is sufficient to prevent mating, but does not prevent the ? from continuing to pursue the /. False mounting is carried on repeatedly and long before actual copulation takes place. In the false mounting position the ? will never rise to a fully upright stance on his hindlegs as he would when mounting a wholly receptive partner. It has been recorded in the wild and also with captive duikers that the ? will actually bite the female’s hindquarters and tail. The continual biting of the / in the same spot will often leave an area of about 4 cm diameter bare of hair and often raw and bleeding. A ? has also been recorded actually biting off the tip of a female’s tail, leaving a raw stump (Wilson 2001). Once the / is receptive and no longer runs from the ?, he rises onto his hindlegs, positioning them as close to those of the / as possible, and, clasping the / with his forelegs, mates with her. Intromission and ejaculation are performed very rapidly. Following ejaculation there is a short refractory period of about half an hour before mounting takes place again. The ? will often mate three or four times before moving off. He then loses interest in her but later in the day more mating bouts will take place continuing for as long as the / is receptive. Females retire to the cover of dense vegetation to give birth. Parturition seems to take place either early morning or late afternoon, or in the hours of darkness (Wilson 2001). The / cleans the lamb immediately following birth, and usually eats the afterbirth; young are precocious and stand up and attempt to suckle shortly after birth.The lamb is hidden near the birthplace, but the / remains in close association with her young for several days following birth, frequently visiting it (four to six times per day) and strengthening the mother–young bond. Lambs will suckle as much as eight times per day, usually early in the morning, late afternoon and during the night (Wilson 2001). Reproduction and Population Structure  The information that follows is summarized fromWilson (2001), unless otherwise indicated. Females become sexually mature when they are between 8.5 and 10.5 241

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months old; a known-age / born in captivity came into oestrus at eight months, and had her first lamb when 15 months old. Males become sexually mature about 3–4 months later. Ovulation takes place in the right horn of the uterus. Of 231 reproductive tracts dissected, Wilson (2001) recorded 110 cases where ovulation was from the left ovary and 121 from the right ovary. In all cases implantation was in the right uterine horn. The gestation period of 29 captive births, in which mating and subsequent births were witnessed, was between 189 and 216 days with an average of 200 days. Neonates have a mass of about 1.7 kg, but this can vary considerably depending on the size of the /. Newly born Common Duikers in Botswana have a mass almost twice those from Ghana, where adults are half the size of those in Botswana. Young gain as much as 600 g in weight in the first week of life (Wilson et al. 1984). Lactation lasts approximately 3–4 months and weaning is gradual, with young starting to nibble on leaves when only two weeks old. Common Duiker milk has a crude protein percentage of about 8.5%, ash 1.2%, moisture 79%, fat content 8.2% and lactose 3.9%. In view of the fact that the gestation period of the Common Duiker is so long for such a small animal and secondly that the / will often come into oestrus soon after giving birth (see below), there is no clear season of birth. However, Wilson (2001), when studying duikers in E Zambia, recorded young throughout the year, while in a similar study in the south-east lowveld of Zimbabwe where the rainfall is a lot less than E Zambia, young duikers were recorded from Nov to Mar, suggesting a birth peak. Females often come into oestrus within a week of giving birth and many // of breeding age are often pregnant and lactating at the same time. Very little information is available on the inter-birth interval of the Common Duiker and all the data that are available come from animals in captivity. Mentis (1972) mentions 267 days while von Ketelhodt (1977b) gave the average interval as 259 days. However, his records were for only one pair of duikers that produced 11 young over a period of seven years. From data collected by Wilson (2001) over an extended period of time from several localities in southern Africa, the mean interval between births was 237 days with a range of 199 to 271 days (n = 20). Jones (1993) gave a longevity record for S. g. grimmia in captivity of 11 years and 10 months, but Wilson (2001) noted that the oldest ‘known-age’ captive duiker reached an age of 26.5 years, while there were several other captive animals that attained ages of 15, 17, 19 and 21.5 years. Predators, Parasites and Diseases  The Common Duiker has a very large number of predators and there is hardly a mammalian carnivore or bird of prey that does not feed on this species. Two very detailed studies of the predators of the Common Duiker, undertaken in E Zambia and in the Matobo Hills in Zimbabwe, showed that at least 22 species of mammals and birds were recorded feeding on this duiker (Wilson 1966b, 2001). The adults are taken by, among others, Leopards Panthera pardus, African Wild Dogs Lycaon pictus, jackals Canis spp., Cheetahs Acinonyx jubatus and hyaenas, while young may be taken by baboons Papio spp., many of the larger birds of prey, including Martial Eagles Polemaetus bellicosus, Verreaux’s Eagles Aquila verreauxii and Wahlberg’s Eagles A. wahlbergi, as well as Giant Eagleowls Bubo lacteus and African Rock Pythons Python sebae. While the Secretary Bird Sagittarius serpentarius feeds on rodents, snakes and

Common Duiker Sylvicapra grimmia.

insects, there is a record of a young duiker being killed by one in Matobo N. P. (V. J. Wilson pers. obs.). Crotalariosis, intoxication due to the legume Crotalaria, is the only known cause of extreme overgrowth of horn affecting all four feet in domestic animals and some wild antelopes. In the Common Duiker, crotalariosis is the laminitic form of crotalism caused by an acute inflammation in the horn-forming tissue due apparently to some toxic process resulting from the ingestion of one or more species of Crotalaria. In the acute form there is great pain that later subsides. This induces abnormality of gait, which, together with the excessive growth, results in an elongated and deformed foot becoming noticeable after a period, often showing a succession of rings. Laminitis has been recorded in a number of Common Duiker in Matabeleland, Zimbabwe. Additional details of this disease can be found in Wilson (2001). A very large number of both external and internal parasites have been recorded from the Common Duiker, including roundworms (nematodes), flukes (trematodes), tapeworms (cestodes), and arthropod parasites such as flies, lice and ticks. Nematodes include species of the genera Cooperia, Haemonchus, Impalaia, Longistrongylus, Nematodirus, Ostertagia, Telodorsagia, Trichostrongylus, Trichuris and Setaria, as well as Oesophagostomum columbianum, Skrjabinodera kuelzii, Subulura distans and the lung-worm Dukerostrongylus kenyae (Round 1968, Dinnik & Boev 1982, Boomker et al. 1983, 1986, 1987, 1989a, Boomker & Reinecke 1989). Trematodes include Paramphistomum sp. (Boomker et al. 1983), Cotylophoson cotylophoson (Round 1968), Schistosoma bovis (Sobrer 1975) and Fasciola hepatica (Graber et al. 1980). Cestodes include Moniezia expansa, Cysticercus spp., Echinococcus spp., Avitinella centripunctata, Stilesia hepatica, Taenia spp. and Thysaniezia sp. (Round 1968, Graber et al. 1980, Boomker et al. 1987, 1989a). Arthropod parasites are also numerous, including: ixodid ticks, such as Amblyomma hebraeum, Boophilus spp., Haemaphysalis spp., Ixodes spp. and Rhipicephalus spp.; sucking lice, Linognathus breviceps and L. zumpti; biting lice, Damalina; and the louse fly Lipoptena paradoxa (Boomker et al. 1983, Horak et al. 1989).

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Sylvicapra grimmia

Conservation  IUCN Category: Least Concern. CITES: Not listed. The Common Duiker is in the very fortunate position that, at the present time, no special conservation measures are necessary to protect this species. Its wide distribution in sub-Saharan Africa, coupled with its adaptability and the fact that it is capable of eating almost anything, means that it survives in many areas where most other antelopes have been eliminated. It occurs in good numbers in many specially protected areas throughout sub-Saharan Africa. Measurements Sylvicapra grimmia TL (??): 920 (910–932) mm, n = 12 TL (//): 948 (932–954) mm, n = 14 T (??): 122 (100–149) mm, n = 12 T (//): 123 (101–146) mm, n = 14 HF c.u. (??): 262 (231–286) mm, n = 12 HF c.u. (//): 269 (234–289) mm, n = 14 E (??): 93 (90–107) mm, n = 12 E (//): 103 (92–125) mm, n = 14 Sh. ht (??): 433 (402–461) mm, n = 12 Sh. ht (//): 448 (386–478) mm, n = 14 WT (??): 11.2 (9.7–13.6) kg, n = 12 WT (//): 12.4 (10.3–14.7) kg, n = 14 NW Ghana (Wilson 2001)

HF c.u. (??): 330 (280–350) mm, n = 45 HF c.u. (//): 338 (240–360) mm, n = 48 E (??): 134 (113–151) mm, n = 45 E (//): 135 (116–153) mm, n = 48 Sh. ht (??): 570 (490–630) mm, n = 45 Sh. ht (//): 600 (490–680) mm, n = 48 WT (??): 19.3 (14.8–22.4) kg, n = 45 WT (//): 21.8 (16.1–26.3*) kg, n = 48 Kalahari Desert, Botswana (Wilson 2001) *The heaviest mass recorded by Wilson (2001); this / had a total length of 1280 mm and stood 680 mm at the shoulder Maximum recorded horn length is 18.1 cm for a pair of horns from South Africa (Rowland Ward) Key References  Allen-Rowlandson 1986; Dunbar & Dunbar 1979; Wilson 1966a, 2001; Wilson & Clarke 1962. Vivian J. Wilson

TL (??): 1000 (950–1060) mm, n = 126 TL (//): 1050 (980–1120) mm, n = 97 T (??): 120 (110–140) mm, n = 126 T (//): 130 (100–150) mm, n = 97 HF c.u. (??): 290 (270–300) mm, n = 126 HF c.u. (//): 290 (270–320) mm, n = 97 E (??): 115 (104–127) mm, n = 126 E (//): 115 (103–132) mm, n = 97 Sh. ht (??): 530 (500–560) mm, n = 126 Sh. ht (//): 530 (499–570) mm, n = 97 WT (??): 13.9 (12.9–16.6) kg, n = 126 WT (//): 15.4 (12.7–16.8) kg, n = 97 E Zambia (Wilson 2001) TL (??): 1080 (1020–1140) mm, n = 52 TL (//): 1110 (1060–1150) mm, n = 61 T (??): 130 (120–140) mm, n = 52 T (//): 130 (100–140) mm, n = 61 HF c.u. (??): 300 (290–310) mm, n = 52 HF c.u. (//): 310 (300–320) mm, n = 61 E (??): 125 (123–126) mm, n = 52 E (//): 126 (125–128) mm, n = 61 Sh. ht (??): 550 (540–560) mm, n = 52 Sh. ht (//): 560 (530–580) mm, n = 61 WT (??): 15.9 (14.5–17.3) kg, n = 52 WT (//): 18.2 (17.2–19.1) kg, n = 61 SW Zimbabwe (Wilson 2001) TL (??): 1100 (1010–1260) mm, n = 45 TL (//): 1165 (1050–1280) mm, n = 48 T (??): 132 (105–164) mm, n = 45 T (//): 153 (120–195) mm, n = 48

Common Duiker Sylvicapra grimmia.

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Family Bovidae

Genus Cephalophus Forest Duikers Cephalophus C.H. Smith, 1827. In: Griffith et al., Anim. Kingd. 5: 344.

Cephalophus is a genus that currently embraces all duikers except for the two or more species of Philantomba and the monospecific Sylvicapra. The separation of Philantomba based on morphological characters (e.g. Pocock 1910) has had strong support from molecular studies that indicate that these small duikers were the first of the extant species to diverge from the rest of the duikers (Jansen van Vuuren & Robinson 2001). The Common Duiker Sylvicapra grimmia occupies a much more ambiguous position, with different molecular trees positioning it wholly outside, marginally outside and wholly within the Cephalophus mainstream duikers (Heyden 1968, Robinson et al. 1996b, Jansen van Vuuren & Robinson 2001, Hassanin et al. 2012). Molecular studies have confirmed that duikers as a whole, as well as the genus Cephalophus, conform with the prediction (Kingdon 1982) that small duikers should be the more conservative while the larger duikers are among the most recently evolved (Robinson et al. 1996b, Jansen van Vuuren & Robinson 2001). According to the molecular studies of Jansen vanVuuren & Robinson (2001), the most consistently conservative Cephalophus species is the small Aders’s Duiker C. adersi, with another apparently early branch being the smallish and somewhat specialized Zebra Duiker C. zebra.The bulk of the radiation is made up of a large number of medium-sized ‘red duikers’ with a very densely complex genealogy and a later offshoot of ‘giant duikers’ and ‘fibre duikers’. The complexity of red duiker relationships may well be further complicated by long-term hybridization along the frontiers of expanding and contracting ranges. The most plausible match between the very tentative trees devised by molecular scientists and the biology and biogeography of living species invites the following sketch of possible evolutionary sequences. The relictual Aders’s Duiker, confined to Zanzibar and isolated forest fragments along the east African littoral, is the last ‘eastern peripheral’ relic of a once widespread, very early red duiker. The Zebra Duiker might be an equivalent ‘western peripheral’ species that has mitigated its marginal status by becoming more highly specialized. The rest of the ‘red duikers’ are of substantially larger size and can be divided into the following categories:

the White-bellied Duiker C. leucogaster, Red-flanked Duiker, Blackfronted Duiker C. nigrifrons and Rwenzori Red Duiker C. rubidus. 3. ‘High-forest frugivores’: species that only really flourish in highly diverse forests with multi-species fruiting trees and a broad spectrum of primates and birds assisting the wind in dropping high-quality debris from the canopy. Species in this category are highly competitive and suitably rich forests only support one species in this size class (15–25 kg). The situation is complicated by the likelihood that the older members of this class (Ogilby’s Duiker C. ogilbyi and allies) are being actively displaced by larger and more recently evolved forms (Peters’s Duiker C. callipygus, Weyns’ss Duiker C. weynsi and the Black Duiker C. niger). Molecular trees are somewhat contradictory in tracing the origins of the remaining Cephalophus species. All are large, heavily built animals and they probably derive from an early, large ‘high-forest frugivore’. They fall into two groups: the first comprises very large but slender, generalized duikers, including Abbott’s Duiker C. spadix and the Yellow-backed Duiker C. silvicultor, while the second consists of two mainly nocturnal ‘fibre duikers’ that are primarily nocturnal and seek out widely dispersed, heavy and often fibrous fallen fruit, namely the Bay Duiker C. dorsalis and Jentink’s Duiker C. jentinki. It is possible that nocturnal duikers may be longer-lived than other forms and this might well be an advantage for species that need to get to know a large home-range. Most duikers are subject to very high levels of predation, particularly by humans. They are easy to snare, net, call up to a lure, dazzle by flash-light and hunt with dogs. The rarer species are already close to extinction and many others are known to be in decline. As one of the most interesting and complex of bovid evolutionary radiations, they deserve much more attention than they have to date. The extinction of once common and widespread providers of high-quality meat will only impoverish further already marginalized forest people. Both research and conservation are urgent. Jonathan Kingdon

1. ‘Successful peripheral generalists’: relatively conservative species living in forests or galleries that are outliers geographically speaking and ecologically degraded (see map right). The relative poverty of fallen fruit and foliage ensures there are few other duikers with which to compete. This category includes the Natal Red Duiker C. natalensis and Harvey’s Duiker C. harveyi. The Red-flanked Duiker C. rufilatus could also come here, but fits better in the next category. 2. ‘Successful specialists’ (deriving from older lineages): species that have escaped competition by becoming specialized in forest types that are less well endowed with a rich variety of fruits.These forests range from mono-dominant patches such as the Gilbertiodendron ‘Limbali’ forests of central Africa to a variety of forest ‘margins’ such as actual forest edges (notably the broad swathe of forest mosaics and galleries along the northern boundaries of the main forest block), swamp forests at both low and high altitudes and the high, cold reaches of tall mountain massifs. Species in this category include

Distribution of the smaller red duikers. Note that all have peripheral or relict distributions or have specialized habits.

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Cephalophus zebra

Cephalophus zebra  Zebra Duiker (Banded Duiker) Fr. Cépalophe zébré (Cépalophe rayé); Ger. Zebraducker Cephalophus zebra Gray, 1838. Ann. Mag. Nat. Hist. 1: 27–30. Sierra Leone.

Zebra Duiker Cephalophus zebra.

Taxonomy  Monotypic. Ogilby’s (1837) account of Antilope doria is actually a reference to the Mhorr Gazelle Nanger dama mhorr (see Kuhn 1966). Following a request from Ansell (1980), the ICZN ruled that zebra is to be given priority over doria whenever the two specific names are considered synonyms (see also Grubb 2004 for a discussion). In a multiple analysis of mtDNA, Jansen van Vuuren & Robinson (2001) found the position of this species within the duiker molecular radiation particularly obscure. Parsimony analysis hinted that C. zebra and Aders’s Duiker C. adersi might be sister taxa, but this was not supported by maximum likelihood analysis, which was consistent with both species having basal positions between the ‘red duiker’ and ‘giant duiker’ complexes. Other, apparently anomalous linkages with Ogilby’s Duiker C. ogilbyi (which gives several indications of being an early, conservative relict species) could also imply that C. zebra derives from a relatively early stem duiker. Synonyms: doria, doriae, zebrata. In contrast to all other duiker species studied to date, the diploid number is 2n = 58, with a pair of submetacentric autosomes; the X chromosome is similar to that of other duikers karyotyped, but theY is one of the smallest acrocentric chromosomes (Bogart et al. 1977, Hsu & Benirschke 1977). Description  Small- to medium-sized duiker, with a panel of vertical, vivid black and cream stripes from behind the shoulder to the tail. There is a frontal rufous-coloured tuft, though it is never

very long, and it usually obscures the short horns (with the result that animals may appear hornless, especially //). Head, shoulders and lower legs russet red with hocks and leg joints nearly black; muzzle slate-grey and nose black. Hair on the neck and shoulders is distinctly shorter than on the body, while the underparts also have rather short hair. The number of dark brown or black vertical stripes varies between 12 and 16, sometimes with light shadow stripes in between, and the width of individual stripes varies greatly (Wilson 2001). Tail tufted, well haired and relatively long, black hairs intermixed above, but white below. Lateral hooves very small. Preorbital glands present. Tarsal glands prominent and tufted just below the heels of the back legs. Inguinal glands entirely different to those of any other duiker, with tiny round holes in the groin (Zeller & Kuhn 1991, Wilson 2001). Both sexes have small (about 4 cm) but robust, conical, smooth and very sharply pointed horns above a forehead and nasal area where the bone is massively thick and reinforced. The horn cores of the frontalia rise less above the profile line of the frontalia than in other species. The enlarged nasal cavity is particularly conspicuous in the skull. Preorbital depressions are relatively tiny. The second incisor and canine of the lower jaw are further reduced than, for example, in the Bay Duiker C. dorsalis or Black Duiker C. niger (Kuhn 1966). Geographic Variation  None recorded. 245

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Lateral, palatal and dorsal views of skull of Zebra Duiker Cephalophus zebra.

Similar Species  Several species, notably Maxwell’s Duiker Philantomba maxwelli, Bay Duiker, Black Duiker and Brooke’s Duiker (C. ogilbyi brookei) are sympatric with Zebra Duikers. However, the striped colour pattern of the Zebra Duiker renders this species quite unmistakeable. Distribution  Endemic to West Africa, ranging from E Sierra Leone (Moa R.) to Côte d’Ivoire (Niouniourou R.) within the primary forest zone. Most common in east-central Liberia. Its distribution and numbers have declined markedly because of forest destruction and excessive hunting for bushmeat. Its remaining strongholds are Sapo N. P. and other forests of SE Liberia, and Taï N. P. and adjacent forest reserves in Côte d’Ivoire (East 1999). Historical Distribution In Sierra Leone, the Zebra Duiker probably occurred quite widely in the southern and central moist lowland forests. It appears to be dependent on undisturbed primary forest and has retreated as forest has been cut down and converted to farmland (Teleki et al. 1990, East 1999). By the 1980s, it was very rare and localized and known to occur in only a few localities, such as Gola N. P. (Davies 1987), the eastern part of Gaura Chiefdom and, possibly, Western Area F. R. (Grubb et al. 1998). In Liberia, it appears to have been relatively common in some regions. In 1974–75, Jeffrey (1977) recorded that this was the third most frequent duiker species encountered in a survey of bushmeat in east-central Liberia. It was also the third most frequently encountered duiker species in the surveys reported by Kranz & Glumac (1983). It was thinly distributed along the Sehnkwen R., but common in the Buto Oil Palm Plantation, and reported by local observers to be abundant along the Sinoe R. (Peal & Kranz 1990). In Côte d’Ivoire, its distribution never extended further east than the Niouniourou R. (which lies between the Sassandra and Bandama Rivers), such that Zebra Duikers have historically always been restricted to the lowland forests in the south-west. Sightings of Zebra Duikers further to the east, for example in Comoé N. P. (see discussion in Fischer et al. 2002), are unsubstantiated.

Cephalophus zebra

Current Distribution Continued presence in Liberia was confirmed in 1997 in Sapo N. P. and from the forests bordering Côte d’Ivoire (East 1999). In Côte d’Ivoire the species is now confined to remaining areas of primary forest within its former range, namely Taï N. P. and the adjoining Haute Dodo–Rapide–Grah–Hana Forest Reserves, and the Cavally–Goin, Scio and Niegré Forest Reserves. Its main stronghold is Taï N. P., where it is seen regularly, and its abundance varies from common in the west to uncommon in the centre and rare in the east (Hoppe-Dominik et al. 1998). In Sierra Leone they persist in the Gola N. P. in the south-east (Lindsell et al. 2011, F. DowsettLemaire & R. J. Dowsett pers. comm.), but their presence in the west is unclear. The presence of Zebra Duikers in Guinea was not mentioned by Sournia et al. (1990) or Grubb et al. (1998), but East (1999) remarked that their presence was confirmed by a report from the Ziama and Diécké Forest Reserves, presumably a reference to Butzler (1994). Wilson (2001) thought it likely that they should occur. The species was not reported by Barrie & Kant (2006) in a rapid survey of Diécké in 2003. Habitat  Found in undisturbed primary forests and along their margins and in clearings, sometimes extending into secondary growth and swidden cultivation (Newing 2001). It favours intact lowland forest (notably the Sinoe and other river valleys in Liberia), but may also live in low montane and hill forests. Abundance  Assuming average population densities of 2.0/km² where it is known to be common and 0.2/km² elsewhere, East (1999) gives an estimated total population of about 28,000. However, Wilson (2001) regarded this as an overestimate, and doubted that there could be more than 15,000 animals across the range at most. Davies (in Grubb et al. 1998) estimated peak population density in little-hunted areas in Sierra Leone at 3–10/km2. Hoppe-Dominik et al. (2011) calculated a population density of 0.4/km2 in Taï N. P.

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Cephalophus zebra

using night counts and a total population of perhaps 2000 animals. The population trend is generally downwards because of bushmeat hunting and continuing destruction of West Africa’s few remaining primary forests.The only exceptions are a few localities where hunting pressures are low and/or there is effective protection against logging and poaching, such as the western section of Taï N. P. (East 1999, Hoppe-Dominik et al. 2011). Adaptations  Known to be a strong stimulus to the eye/brain sensory system, stripes may serve as a focus for social attraction (in which case the resemblance of these duikers to zebras may be more than superficial). An advantage of stripes in this species could be to inhibit goring of the soft abdomen during aggressive encounters. Thus, while other animals shield themselves from the worst effects of aggressive rivalry with thickened skin, bony plates and so forth, these duikers may have a subtler strategy for reducing excessive aggression (Kingdon 1997). Grubb et al. (1998) suggested that Zebra Duikers are both diurnal and nocturnal. Captive animals in Monrovia Zoo, Liberia, were active about 70% of the daytime (Newing 2001); animals in Taï N. P. were mainly active at night (Hoppe-Dominik et al. 2011). Foraging and Food  Diet comprises fruits and foliage. Details are unknown, but the Zebra Duiker does not appear to have specialized teeth or diet. The stomach contents of four duikers shot in Liberia indicated that at least 79% of plants eaten were fruits and seeds, and all four stomachs contained large numbers of the fruits of Diospyros sanza-minika; two stomachs contained seeds of Bussea occidentalis (Wilson 2001). The animal may use its head to break open the shells of larger fruits. In captivity, occasional consumption of meat (selfcaught mice) has been observed (Schweers 1981), and indeed one of the stomachs mentioned by Wilson (2001) contained the remains of an unidentified rodent. Social and Reproductive Behaviour  All of 35 field observations from Taï N. P. were of lone animals, suggesting a primarily solitary

Zebra Duiker Cephalophus zebra.

social system (G. Radl pers. comm.). Friendly relations between captive pairs involve mutual rubbing and licking, suggesting that breeding pairs are the normal social unit (as they are in many duikers). Because both sexes have horns and a thickened skull (which do not appear to correlate with a specialization in ecology or diet), it seems likely that bonded pairs share defence of their home-range against intruding duikers and their offspring against predators. Scarred heads suggest that the collisions that are normal in duiker confrontations are particularly vigorous and uninhibited in these exceptionally stocky and muscular antelopes (Kingdon 1997). Schweers (1984) conducted a detailed study of breeding behaviour and reproduction on captive Zebra Duikers in Frankfurt Zoo. Males will determine the female’s reproductive status by smelling her urine and performing flehmen. While a consorting pair mutually mark each other even outside of the female’s oestrous period, the frequency of this marking increases in the ? as the / approaches oestrus, with the ? applying secretions from his preorbital glands to the head, neck and back of the /. As the / approaches oestrus, the ? follows the / more closely, and licks her neck, hindlegs and anogenital region. The ? usually follows directly behind the /, sometimes making grunting sounds, his head stretched forward such that his neck is horizontal and the head is held at the height of the female’s tarsal gland. Mounting attempts occur when the / stops moving, and is preceded by the ? smelling and licking the female’s anogenital region; Schweers (1984) also recorded frequent high lifting of the foreleg past the female’s hindlegs (in contrast to between the legs as occurs with laufschlag in gazelles). Copulation is brief, lasting only a few seconds, but is repeated frequently. Birth in captivity is normally in the morning hours, and typically takes place while the mother stands, the neonates mainly appearing front first.The mother then eats the afterbirth.The young are precocious, but remain alone lying up until Day 20 (Schweers 1984). Expressions of sound are rare and, besides during mating, have only been observed in captivity during initiations of contact between mothers and their offspring, and during fighting (Schweers 1984). Reproduction and Population Structure  Detailed infor­ mation about reproduction is given by Schweers (1984) and Barnes et al. (2002) from observation of captive animals. After a postpartum oestrus, // are normally successfully mated as little as 10 days after birth (Schweers 1984). Gestation is between 221 and 229 days (although Pfefferkorn 2001 gives a range of 190–245 days) and the birth interval 241 days (Schweers 1984). A single young is born. The birth weight is around 1270–1550 g for ?? and 1300– 1750 g for // (Barnes et al. 2002). Neonates are born with the dark stripes visible, although they appear closer together, giving the young an overall darker appearance (Schweers 1984). The weight development during the first 60 days ranges from 47 to 74 g/day; at 40–45 days weights are around 4450 g for ?? and 3620–4620 g for // (Barnes et al. 2002). Young are able to nurse shortly after birth, and for the first few weeks will nurse about four times per day for several minutes (Schweers 1984); solid food is consumed from the fifth week. The young animal is weaned after about 95–111 days (Barnes et al. 2002). Horns begin to appear at 1–2 months of age, and at the age of 7–9 months juveniles have the adult colouration and have reached adult height (Schweers 1984). Males reach sexual maturity at about two years of age (Barnes et al. 2002). A wild-born animal survived to 13 years in captivity (Weigl 2005). 247

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Family Bovidae

Predators, Parasites and Diseases  The main predator in un­ disturbed habitat is undoubtedly the Leopard Panthera pardus. HoppeDominik (1984) showed that the Zebra Duiker and Black Duiker made up a large part of the spectrum of prey (up to 10%) in Taï N. P. Immature Zebra Duikers are present in the prey of the Crowned Eagle Stephanoaetus coronatus in Taï N. P., but in low frequency (S. Shultz pers. comm.). Further, the African Golden Cat Profelis aurata and the African Rock Python Python sebae are likely predators. There is no information available on parasites or diseases. Conservation  IUCN Category: Vulnerable A2cd, C1. CITES: Appendix II. Formerly widespread over much of Liberia and Sierra Leone, Zebra Duikers are declining fast as their habitat is being destroyed and commercial bushmeat hunting becomes more entrenched and comprehensive in its onslaughts. Wilson (2001) considered them the least adaptable to deforestation of all West African duiker species and therefore the least likely to survive hunting pressure and habitat degradation. Actual consumption of bushmeat was measured by Caspary et al. (1999) in a one-year study in the region of Taï N. P. For the year of 1988 they estimated that 73,000 subsistence hunters killed about 1500–3000 tonnes and a group of professional hunters killed 52–650 tonnes of bushmeat in the region (2700 km²). The share of weight of the Zebra Duiker was about 2.1% of meat at market stands in the east of the park (26 Zebra Duikers out of a total of 2171 animals killed) and 1.8% in the west (36 duikers out of a total of 5101 animals killed there). The share of the Zebra Duikers killed in the forests to the west of the R. Cavally in Liberia and smuggled into Côte d’Ivoire was around 5%. Of a total of 11,215 animals, 440 Zebra Duiker cadavers were counted. Data collected from hunters (representing approx. 3% of the total village population) in three villages in Sinoe County in Liberia over 10 months (2001–2002)

demonstrated that Zebra Duikers represented the fourth most commonly killed animal (R. Hoyt pers. comm.). Due to undiminished demand for game meat and unsustainable hunting, there is a high risk that species with restricted ranges such as the Zebra Duiker (which is dependent upon primary rainforests) will be extirpated. Existing protected areas therefore need to be extended, networked and their protection considerably improved. Important conservation areas for the protection of this species include the Gola complex of reserves in Sierra Leone, Sapo N. P. and Grebo National Forest (Liberia), and Taï N. P. and surrounding forest reserves in Côte d’Ivoire (East 1999, Wilson 2001). Measurements Cephalophus zebra HB (??): 846 (830–870) mm, n = 4 T (??): 125 (122–127) mm, n = 4 HF c.u. (??): 221 (215–227) mm, n = 4 E (??): 79 (77–80) mm, n = 4 WT (??): 13.5, 14.2 kg, n = 2 WT (/): 14.7 kg, n = 1 Liberia (Kuhn 1966) Wilson (2001) gives an average of 17 kg for five ?? (range 15–21 kg), and 18 kg for four // (range 15–23 kg). Shoulder height of // averages 460 mm, compared with 440 mm in ?? Maximum recorded horn length is 7.9 cm for a pair of horns from Liberia (Rowland Ward) Key References  East 1999; Kingdon 1997; Schweers 1984;Wilson 2001. Bernd Hoppe-Dominik

Cephalophus adersi Aders’s Duiker Fr. Céphalophe de Aders; Ger. Adersducker Cephalophus adersi (Thomas, 1918). Ann. Mag. Nat. Hist., ser. 9, 2: 151. Tanzania, ‘Zanzibar’.

The vernacular name is after William Mansfield-Aders, a zoologist for the government in Zanzibar in the first few decades of the 1900s. Taxonomy  Monotypic. The species has been considered con­ specific with both Natal Red Duiker C. natalensis and/or Peters’s Duiker C. callipygus (e.g. Heyden 1968, Dorst & Dandelot 1970, Haltenorth & Diller 1980), but is currently regarded as a separate species (Kingdon 1982, 1997, Grubb & Groves 2001, Wilson 2001, Grubb 2005). On the basis of a molecular study of duikers, Jansen van Vuuren & Robinson (2001) found that this species occupied a consistently basal position in the duiker radiation, confirming Kingdon’s (1982) prediction that this might be a relictual species from an early phase in the evolution of ‘red duikers’. Synonyms: none. Chromosome number: not known.

Aders’s Duiker Cephalophus adersi.

Description  A small to medium-sized duiker standing about 40– 45 cm at the shoulder when fully grown and weighing 8–12 kg (Kingdon 1982,Williams et al. 1995,Wilson 2001).The fur is soft and

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Lateral view of skull of Aders’s Duiker Cephalophus adersi. Occlusal view of the upper-right toothrow in Aders’s Duiker Cephalophus adersi.

above:

below:

Aders’s Duiker Cephalophus adersi.

silky, with a marked cow-lick or whorl of hair on the nape of the neck but little change in texture from the neck to the main body (Kingdon 1982, 1997). The muzzle is pointed, with a flat front to the nose. The overall colour is a reddish-ochre but grey on upper shoulder and back of neck and light fawn on the sides of the face, neck, shoulder and flanks. The bridge of the nose and crown are richly red with a prominent darker tuft at the horns. The underbelly is clear white. Aders’s Duiker can be distinguished from other red duikers by the white freckling on the lower limbs and a broad white band on the rump, which merges into the underparts (Kingdon 1982, Wilson 1987, Williams et al. 1995). Preorbital glands present; pedal glands long, narrow and well developed. Inguinal glands are absent. Aders’s Duiker have simple spiked horns that grow to a little over 5 cm in length (Kingdon 1982,Williams et al. 1995,Wilson 2001). In the skull, the preorbital fossae are deep with a sharply defined upper margin; there is no interfrontal groove and no marked frontal convexity. The nasals broaden distally (Grubb & Groves 2001). Geographic Variation  The species is monotypic, although Kingdon (1982) reported hunters’ assertions that animals from NE Zanzibar (Kiwengwa) were darker and heavier than animals from Jozani Forest (south-central), which were paler and lighter in weight (see Wilson 2001). Similar Species Cephalophus harveyi. Sympatric in coastal Kenya. Larger, redder and lacks white belly and buttock stripe. Distribution  Endemic to Africa. In Kenya, Aders’s Duiker has been described as once widespread in the forests, woodland and thickets north of Mombasa up to the Somali border (Kingdon 1982). However, as a result of rapidly shrinking suitable habitat, the population has succumbed to a severe decline in numbers. Until recently, the species was thought to have become confined to a very small population in the Arabuko-Sokoke Forest north-west of Kilifi (East 1999), which covers approximately 416 km². However, in 2004, an Aders’s Duiker was sighted in Dodori National F. R. north of the Tana River Delta on the N Kenya coast (Andanje & Wacher 2004), where earlier recorded by Gwynne & Smith (1974). Since then extensive camera-trapping work has found that Aders’s Duiker is the most frequently recorded antelope in the Boni-Dodori forests (Andanje et al. 2011a, b). One reason for their distribution and habits being poorly known is that Aders’s Duikers

inhabit coastal forests and thickets that often grow on very coarse coral rag, an inhospitable, waterless and, in places, impenetrable, habitat. Given such habitats and a presumably recent connection between the Zanzibar and coastal populations it is possible that the species may survive in other pockets of similar habitat elsewhere along the East African coast, especially in coastal habitats between the Pangani/ Rufiji and Galana/Tana rivers (Andanje et al. 2011b). Because they show no signs of being exceptionally specialized and because of their likely basal position in the duiker radiation, a primary constraint on their distribution may well be the existence of competition from other red duikers, suggesting that their overall distribution is intrinsically relictual (J. Kingdon pers. comm.). In Tanzania, Aders’s Duikers still occur on Unguja (Zanzibar) I., although the population has declined substantially and now faces an increasingly uncertain future. There is a possibility that Aders’s Duikers may have once occurred on Fundo I., off the coast of Pemba I. (see Williams 1998), and it is reported to have been introduced to Funzi I., Pemba (Kingdon 1997), but has since become extinct on both these islands. Archer & Mwinyi (1995) mention unconfirmed, but reliable, reports indicating a thriving population on Tumbatu I. – a small island lying off the north-west of Unguja I. While this was once certainly the case, the Aders’s Duiker population on Tumbatu has heavily declined and may now no longer exist. In February 2000, five Aders’s Duikers were translocated to Chumbe I. (a private marine reserve) lying off Unguja I. to complement a single previously translocated / (see Mwinyi & Wiesner 2003). A few individuals were also translocated to Mnemba I. off the north-east coast of Unguja and are breeding (K. Siex pers. comm.). Habitat  In Zanzibar, Aders’s Duikers inhabit tall, undisturbed thicket known locally as ‘msitu mkubwa’ (Archer 1994) in the waterless coral-rag on the eastern side of the island. Aders’s Duiker is very sensitive to habitat disturbance, but occasionally occurs in secondary thicket (Williams et al. 1995). In prime habitat south of Jozani Forest, the emergent tree layer is dominated by Mystroxylon aethipicum, Diospyros consolatae and sub-dominantly by Euclea schimperi. The shrub layer is dominated mainly by Polysheria multiflora and Canthium hibracteatum (Swai 1983). In Arabuko-Sokoke Forest, Aders’s Duikers are most often sighted in the undisturbed Cynometra vegetation, mostly on red soil, with fruiting vegetation (Kanga 1995, 2002). In Dodori National F. R., Aders’s Duiker has been recorded 249

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from dense coastal thicket with full canopy cover to ca. 4–5 m and a ground layer of leaf litter and tangled stems on sandy substrate (Andanje & Wacher 2004). Adaptations  Wilson (2001) commented that the preorbital gland of Aders’s Duiker was different to any other he had studied, being about one-quarter the size of the preorbital gland of even the smallest duikers. He found the gland to be near-vestigial and only a small part was active from which only small quantities of secretion could be squeezed. The gland of a ? had short rufous hair on top of the gland and above the opening, while below the slit there was no hair at all, and the slit of the gland was about 8 mm long and 2 mm wide. Wilson (2001) suggested that Aders’s Duikers might not mark their territories to the same extent as other duikers and might, therefore, be losing the use of their preorbital glands. On the other hand, J. Kingdon (pers. comm.) thinks it possible that C. adersi conserves an earlier or transitional stage in the evolution of the duiker type of preorbital gland. Aders’s Duikers survive in a very demanding and specifically coastal environment where they must periodically derive their water needs from the plant parts that they eat.To this extent they are coastal specialists and probably have appropriate physiological adaptations. Apart from this they do not exhibit very marked specialization in diet or foraging techniques and their distribution accords with the suggestion that their geographically and ecologically marginal existence may reflect a relictual status (J. Kingdon pers. comm.). These duikers are diurnal/crepuscular and very shy (Archer 1994, Kingdon 1997, Andanje et al. 2011a). Abundance  To date three surveys have been carried out on Unguja I. The first in 1982 (Swai 1983) estimated the Aders’s Duiker population to be in the region of 5000 individuals. A second, more detailed survey undertaken in 1995 (Williams et al. 1995) placed the population between 1200 and 2000. The populations were shown to be located in five main sub-populations with varying degrees of interconnectedness: Kiwengwa Forest in the north, localities in the central Jozani-Chwaka Bay area and Mtende in the south. A third survey carried out in 1999 (Kanga & Mwinyi 1999) placed the population at 614 ± 46 within the same study area as Williams et al. (1995). Although the surveys all adopted different methodologies, they show a very substantial decline in the population over a 17-year period. In 1995, Aders’s Duikers on Zanzibar were found to occur at an overall density of 4.5 ± 1.2/km2 in high thicket (comparing favourably with that of 4.3/km2 recorded by Swai [1983]), with higher population densities of up to 11.4 ± 5.2/km2 in limited areas of particularly undisturbed high thicket habitat (Williams et al. 1995). The impact of hunting was not controlled so the species must have once occurred naturally at higher densities than those measured. Until recently Aders’s Duiker in Kenya was thought to have become restricted to the Arabuko-Sokoke Forest where it was once common but is now considered very scarce. A very approximate figure of about 500 individuals was estimated in 1999 based on a small drive-count survey in the Arabuko-Sokoke Forest (Kanga 2002). In Arabuko, Aders’s Duikers were found to occur at a density of 2.8/km2 (Kanga 2002, 2003a). Overall, recent surveys have only sighted very low numbers (three in 1999, two in 2002 and two in 2003) (Kanga 2002, 2003b). Camera-trapping surveys have shown that Aders’s Duiker is the most frequently recorded antelope in the camera grids in the Boni-

Cephalophus adersi

Dodori forests and surrounds, being present at 51 of 52 camera sites, returning a standardized photo rate of 75 encounters/100 days across three grids dispersed over 1200 km2. In contrast, at Arabuko-Sokoke, a comparable sampling effort returned 0.09 encounters/100 days (Andanje et al. 2011a). Foraging and Food  Aders’s Duikers are browsers selecting for dicotyledonous leaves, seeds and fruit (Swai 1983). They show a particular dependence on the flowers and berries that grow prolifically from common trees, such as Diospyros consolataei, Cassine aethiopica and Euclea schimperi, and bushes such as Canthium spp. and Polyspheria (Kingdon 1997); Wilson (2001) recorded the fruits of D. consolataei, Ficus sur and Tetracella littoralis. In addition to these foods they will eat sprouts, buds and other fresh growth found at ground level (Kingdon 1997). Individuals may sometimes follow troops of Gentle Monkeys Cercopithecus mitis and Zanzibar Red Colobus Procolobus kirkii feeding on discarded fruits and dislodged edibles from the canopy above (Swai 1983, S. Imani pers. comm.). Feeding occurs from dawn to around 11:00h, followed by a period of rest and rumination, before becoming active again from about 15:00h when they continue to forage until nightfall (Kingdon 1997). The species has been reported as being independent of water (Kingdon 1982). Aders’s Duikers are sympatric with Harvey’s Duikers C. harveyi, Blue Duikers Philantomba monticola and Sunis Nesotragus moschatus on the mainland, and with Blue duikers and Sunis on Zanzibar. Little is known regarding their feeding and ecological separation, but camera trapping data suggests that Aders’s Duikers hold their own in relative numbers at key sites such as Boni-Dodori (75 encounters/100 days compared with 55 Suni encounters and two for Harvey’s Duiker), with broadly similar diurnal activities patterns in all three (T.Wacher pers. comm.). It can be predicted that diurnal feeding allows these duikers to benefit from monkey and bird activity in the canopy (Kingdon 1982).

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Aders’s Duiker Cephalophus adersi.

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Social and Reproductive Behaviour  Aders’s Duikers have been reported to live in pairs and to defend territories (Kingdon 1997). However, Aders’s Duikers have also been recorded singly (Williams et al. 1995); out of 3021 separate camera-trap records at Boni-Dodori, only 8.5% comprised two animals (mostly adult pairs, but also mother and young) with less than 1% involving trios (T. Wacher pers. comm.). Reproduction and Population Structure  Aders’s Duikers reportedly breed throughout the year (Rodgers & Swai 1988). During a six-month survey on Zanzibar, Archer & Mwinyi (1995) recorded 40 Aders’s Duikers shot by local hunters, 24 of which were pregnant. Pregnant // were recorded in all months, with three of eight and three of four pregnant // having large foetuses in Aug and Sep, respectively. Predators, Parasites and Diseases  Natural predators, apart from people, have been eliminated on Zanzibar, most notably the Zanzibar Leopard (sometimes afforded the trinominal Panthera pardus adersi), but on the mainland Leopards may still take individuals. At Boni-Dodori, predators such as Lion P. leo, Leopard, Caracal Caracal caracal, African Wild Dog Lycaon pictus and Spotted Hyaena Crocuta crocuta all co-occur with Aders’s Duiker (T. Wacher pers. comm.). A parasite taken from an animal in Zanzibar was described as Tricholipeurus pakenhami (Pakenham 1984). Conservation  IUCN Category: Critically Endangered A4cd. CITES: Not listed. Aders’s Duiker has undergone severe long-term population declines in both Kenya and Zanzibar (Tanzania) due to habitat loss and hunting pressure. In Zanzibar there has been a large amount of deforestation and forest degradation, particularly over the last 30 years, as the human population has increased. Much of the habitat loss has been driven by a demand for domestic fuelwood by a rapidly expanding urban population. This has led to extensive loss of habitat for Aders’s Duikers but also severe habitat fragmentation. Habitat destruction is probably the most significant threat to the species’ survival on Zanzibar, and one that is unlikely to be solvable in the short term. Jozani-Chwaka Bay N. P., which was gazetted in 2004, includes important but only small parts of suitable habitat for Aders’s Duikers as does the proposed Kiwengwa-Pongwe F. R. In Kenya, the Arabuko-Sokoke Forest is one of the last major remnants of lowland forests on the East African coast. Illegal woodcutting has led to the deterioration of the species’ habitat and contributed to the decline of the population. Kanga (2003b) reported that levels of wood-cutting increased between the surveys he carried out in 1998 and 2002. It is likely that habitat destruction will constitute the most difficult threat to address in terms of the species’ future security in Arabuko-Sokoke (Kanga 2003b). The discovery, then, of a significant population of Aders’s Duiker in the Boni-Dodori forest to the north improves the conservation prospect for the species, and highlights the need for improved management efforts in the Boni-Dodori forest (Andanje et al. 2011b). Aders’s Duikers are traditionally hunted in Zanzibar but an increase in hunting pressure likely followed the revolution of 1964 after which enforcement of the wildlife laws became largely nonexistent (Finnie 2004). It would appear that, partly as a result of

hunting, the mini-antelope populations of Zanzibar (especially Aders’s Duikers) have undergone long-term declines (for example, see Williams et al. 1995, Kanga & Mwinyi 1999). The Department of Forests and Non-Renewable Natural Resources began to address the hunting situation in Zanzibar in 1994, through community-based conservation initiatives. Although hunting has now come under an increasing level of control in Zanzibar (both at the village and governmental level) it remains a significant threat up to the present (Finnie 2002). In the Arabuko-Sokoke Forest, trapping is common. In 1991, more than 2600 households lived within 2 km of the forest and at least 33% carried out hunting and/or trapping (FitzGibbon et al. 1995). It is thought that the high level of snare-trapping in the Arabuko-Sokoke Forest represents a significant threat to the continued existence of Aders’s Duikers in this forest (Kanga 1995, 2002, Finnie 2004) as the species is a favoured target because of its sweet meat, which fetches a good price at market. Kanga (1995) stated that when hunters found an area where they suspected Aders’s Duikers occurred, they would saturate the pathways with a large number of traps. Hunters told Kanga that Aders’s Duikers were often caught in the 1970s, but that since the 1980s their hunting success with this species was low. However, Kanga’s (1995) observations can be contrasted with FitzGibbon et al.’s (1995) assertion, without documentation, that trapping for duiker species in 1991 was well within sustainable limits. The impact of hunting on the Aders’s Duiker population that may exist in the vicinity of the Boni-Dodori forests is unknown. Measurements Cephalophus adersi HB (unsexed): 714 (630–780) mm, n = 14 T (unsexed): 111 (60–138) mm, n = 14 E (unsexed): 78 (75–87) mm, n = 14 Sh. ht (unsexed): 404 (375–440) mm, n = 14 WT (??): 9.2 (8.7–10.2) kg, n = 9 WT (//): 9.0 (6.8–10.1) kg, n = 5 Zanzibar (Kanga & Mwinyi 1999) Wilson (2001) noted that Kanga & Mwinyi (1999) failed to indicate how their measurements were taken, and draws attention to the doubtful reliability of their measurements of the hindfoot (mean = 452 mm). Wilson (2001) gave the measurements of two adult ?? as TL: 823–830 mm; T: 100–122 mm; E: 73–81 mm; HF c.u.: 195– 214 mm; Sh. ht: 400–420 mm; and WT: 8.5–10.7 kg. Measurements for three // were TL: 790–846 mm; T: 120–131 mm; E: 68– 76 mm; HF c.u.: 210–225 mm; Sh. ht: 410–435 mm; and WT: 10.5–12.4 kg. Maximum recorded horn length for this species would appear to be that of a ? captured in Jozani Forest in 1999, measuring 5.3 cm and reported by Kanga & Mwinyi (1999). The longest pair recorded by Wilson (2001) was 4.8 cm, on an adult ? killed by hunters and measured by the side of a road near Jozani Forest, which exceeds the 3.8 cm pair recorded by Rowland Ward from the Arabuko-Sokoke Forest Key References  Kanga 2002; Kanga & Mwinyi 1999; Williams 1998; Williams et al. 1995; Wilson 2001. Andrew Williams

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Cephalophus rubidus

Cephalophus rubidus Rwenzori Red Duiker Fr. Céphalophe de Rwenzori; Ger. Rwenzoriducker Cephalophus rubidus Thomas, 1901. Proc. Zool. Soc. Lond. 1901 (2): 89. ‘Ruwenzori District’.

Rwenzori Red Duiker Cephalophus rubidus.

Cephalophus rubidus

of dark grey underlies uniform red tips of fur down mid-line of the back and neck, while underfur of the flanks is cream. Chin and belly white. Hindlegs almost black, dark brown markings on the joints Lateral view of skull of Rwenzori Red Duiker Cephalophus rubidus. of the forelegs. Tail bushy. Pedal and preorbital glands present. The horns, present in both sexes, reach 8–9 cm in length (Kingdon Taxonomy  This species has been treated as a subspecies of C. 1982). natalensis (Schwarz 1914) and as a subspecies of the Black-fronted Duiker C. nigrifrons (St Leger 1936, Ansell 1972, Groves & Grubb Geographic Variation  Currently believed to be restricted to the 1981). Because this species occurs on the same Rwenzori mountain upper reaches of the Rwenzori Mts, no significant variation is known range as another race of the Black-fronted Duiker (C. n. kivuensis), yet or likely. However, sightings of very russet duikers from the bamboo differs from it in features that suggest other affinities, Kingdon (1982, zone of the Bwindi forest in SW Uganda suggest a distinctive duiker 1997) regarded this as a distinct high-altitude red duiker, proposing in a relictual habitat once similar to that in which C. rubidus occurs on that it might have been pushed up into its present sub-alpine and upper the Rwenzori Mts. montane forest habitat by the later arrival of the Black-fronted Duiker. Molecular studies have also indicated that C. rubidus is a distinct taxon Similar Species with relatively old and ill-defined affinities with other red duikers, Cephalophus nigrifrons. Long-legged and long-hooved species from montane and lowland forest habitats. Kingdon (1982) suggested although admittedly the only material available for the analysis was that C. rubidus and C. n. kivuensis (which occurs in the lower altitudes a single tooth from a specimen in the Swedish Museum of Natural of the Rwenzoris Mts) might hybridize where their ranges overlap. History (Jansen van Vuuren & Robinson 2001), which may itself be misidentified.Wilson (2001) was reluctant to consider rubidus a distinct species, and Grubb & Groves (2001) and Grubb (2005) both considered Distribution  Endemic to Africa, where confined to the Rwenzori it a subspecies of C. nigrifrons. Synonyms: none. Chromosome number: Mts (Kingdon 1982, Grubb & Groves 2001). Although it presumably not known, but given their conserved karyotype it is likely to be occurs on the section of the Rwenzori Mts that lies within Virunga N. P. in DR Congo, the presence of this form in this country has not 2n = 60 (B. Jansen van Vuuren pers. comm.). been confirmed (East 1999, Wilson 2001). Description  A medium-sized duiker with a glossy rufous coat with long coarse hair on the neck changing to dense soft hair over Habitat  Mostly found in alpine and sub-alpine habitats between the hindquarters. Black or dark brown blaze on the forehead. Zone the snow/glacier line and bamboo zones, at altitudes of 1300– 253

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4200 m. They occur along the margins of Carex tussock bogs, in afroalpine moorland dominated by Helichrysum, Alchemilla, giant groundsels Dendrosenecio and giant lobelias Lobelia and in heath ‘woodlands’ dominated by tree heaths, Philippia and Erica. Abundance  Data are scarce, but daylight sightings and hunter’s middens reported before the mid-1960s suggest that it was once very abundant in its preferred habitats. East (1999) suggested that the population of the Rwenzori Red Duiker may number at least in the thousands. Adaptations  Apparently able to tolerate very low temperatures at night and frequent cold rain or snow, as well as periodic strong insolation during the day. The peculiarly soft, dense texture of the pelage on the body of this species would seem to be adapted to these peculiar conditions, although no attempt has been made to assess the insulating or water-proofing properties of the hairs. Activity periods are likely to be strongly influenced by the very frequent rain on the Rwenzoris. This species is of some significance for our understanding of mammalian adaptation to an extreme habitat, especially as such situations are rare in Africa. As the only large mammal to inhabit the high, cold and wet afroalpine vegetation zones of the Rwenzori Mts it offers unique opportunities for the study of physiology, dietary habits and behavioural repertoires in a ruminant that has several apparent equivalents in temperate Eurasia, North and South America (notably Goral Naemorhedus goral, Chamois Rupicapra rupicapra, Pudu Pudella mephistophiles and Musk Deer Moschus moschiferus). Molecular studies imply that this species is likely to derive from a population of duikers that preceded the evolution of the specialized Black-fronted Duiker (Jansen van Vuuren & Robinson 2001). While the legs and hooves of this species appear to be less specialized in their proportions than those of the Black-fronted Duiker, it is likely that less obvious physiological adaptations could be significant. Foraging and Food  No direct information is available, but the absence of fruit and abundance of afroalpine herbs (especially Alchemilla) and lichens in the sub-alpine zone suggest that these are possible staples. Rock-cress Arabis alpina, violets Viola spp. and ground orchids Disa spp. are also common, but other flowering plants are few

and small in size. Around bogs, the sedge Carex petitiana as well as the nettle Laportea alatipes might be eaten; a few high-altitude grasses, the heavy rain-exposed bark and roots of plants both large and small are all likely forage. Along the lower margins of its habitat the fruits of Rubus might be important and herbs such as goose-grass Galium, sorrels Rumex spp., wild parsley Peucedanum spp. and borage Cynoglossum spp. are very abundant and probable foods. Social and Reproductive Behaviour  Casual observations suggest that these duikers are generally solitary except when juveniles or subadults accompany their mothers or ?? are accompanying an oestrous /. Reproduction and Population Structure  Nothing is known about the timing of breeding or any other aspect of reproduction in this species. Predators, Parasites and Diseases  Known predators include African Golden Cats Profelis aurata and probably Leopards Panthera pardus. Large, migratory eagles sometimes fly over the Rwenzori Mts, but are not likely to be any more than a rare hazard. Conservation  IUCN Category: Endangered B1ab(iii); C2a(ii) (listed as C. nigrifrons rubidus). CITES: Not listed. A very vulnerable species owing to the ease with which it can be snared and the increase in illegal hunting within Rwenzori Mountains N. P., Uganda, the species’ only known protected area stronghold. This species has long been a highly desirable quarry for Bakonjo hunters from the inhabited foothills. As noted earlier, the species presumably occurs on the section of the Rwenzori Mts that lies within Virunga N. P., but the presence of this form within DR Congo requires confirmation. Measurements No reliable measurements are available. Grubb & Groves (2001) gave mean GLS as 172±4.2 mm (n = 3) for ?? and 180 mm for a single /. Key References  Kingdon 1982; St Leger 1936; Wilson 2001. Jonathan Kingdon

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Cephalophus leucogaster

Cephalophus leucogaster  White-bellied Duiker Fr. Céphalophe à ventre blanche (Céphalophe du Gabon); Ger. Weusbauch Ducker Cephalophus leucogaster Gray, 1873. Ann. Mag. Nat. Hist., ser. 4, 12: 43. ‘West Africa, Gaboon’ (Gabon).

White-bellied Duiker Cephalophus leucogaster.

Taxonomy  Type specimen named from an immature animal collected by Du Chaillu in Gabon. Rode (1943) mistakenly lumped leucogaster and ogilbyi as subspecies of the Bay Duiker C. dorsalis. Ansell (1972) included the form arrhenii as a subspecies of the Bay Duiker, but this form is actually attributable to C. leucogaster (Grubb & Groves 2001). Synonyms: arrhenii, seke. Chromosome number: not known, but likely 2n = 60. Description  Pale, sandy-coloured ‘red duiker’ with a distinctive narrow black dorsal patch, broadest in the middle back. A sleek animal with short dense hair, and gracile build, this species is one of the more delicately built of the forest duikers. Sides of face pale rufous; forehead contains a strong admixture of black hairs, darkening toward the nose producing a distinctive dark frontal blaze. Back of ear pinnae rufous admixed with dark hairs on the outer surface, and variably fringed with light hair on the upper rim and black hairs toward the base of the rim. Inner surface of ear with black patch on lower third of the ear. Both sexes have a coronal tuft of upright, bright rufous hairs between the horns. The dark dorsal patch begins as a narrow dark streak at the base of the neck, intergraded with dark rusty hairs over the shoulders, widening into a nearly pure black, distinctively marked, dorsal patch, widest in the middle of the back, where it sharply contrasts with the pale sandy-coloured sides, narrowing again and ending in a point just above the tail. Neck and sides pale rufous, sprinkled with darker hairs dorsally, and white hairs ventrally. Chest, abdomen and inside of upper legs pale cream, becoming nearly pure white on the lower abdomen. Limbs pale rufous admixed with darker hairs on outer surface on front legs and shoulder. Rear hocks with indistinct black patches. Lower limbs, from metacarpals distally, black. Tail pale rufous-sandy colour at base and along upper surface, with large brush of long black hairs, many with the distal third white-tipped. Preorbital glands, producing a greyish exudate, are present in both sexes, and run from under the eye toward the muzzle. Inguinal glands and pedal glands present in both sexes. Inguinal secretions

Lateral and dorsal views of skull of White-bellied Duiker Cephalophus leucogaster.

are rusty-coloured and are visible on the white ventral surface of some animals. There is little sexual dimorphism: ?? and // have similar pelage, but adult // average between 8% (Congo) and 11% (Ituri Forest, NE DR Congo) larger body mass than adult ?? (see Measurements). Pelage of newborns differs from adults in being overall pale grey, with less distinctive dorsal patch. This pelage is replaced by adult colouring before weaning. Cranium with broadened toothrow and palate, though the skull is not as robust or broad as in the Bay Duiker C. dorsalis. Horns present in both sexes, stout and strongly annulated at base, and variably over three-quarters of their length, turned inwardly at the tips in older ??. Geographic Variation C. leucogaster leucogaster: range of the species, except NE DR Congo. Colour darker and redder. Dorsal stripe narrow (e.g. between 21 and 84 mm on the Cameroon coast); slightly smaller in size. C. leucogaster arrhenii (includes seke): NE DR Congo. Colour paler, browner. Dorsal stripe broader (in Ituri 80–156 mm wide); size slightly larger. Similar Species Cephalophus ogilbyi. TheWhite-legged Duiker (C. o. crusalbum) is partially

sympatric with White-bellied Duiker in W Gabon. Distinguished by its overall more russet colour and white patches on its legs. C. dorsalis. Broadly sympatric in Gabon. Larger and overall darker in colour and with the dorsal band widest over the rump, not in the middle of the back. C. callipygus. Sympatric in western central Africa. More rufous; dorsal patch widest over the rump. C. weynsi. Sympatric east of the Ubangi R. Deeper rufous in colour and lacks the black dorsal patch. Distribution  Endemic to the central African lowland forest zone, and apparently restricted to forest below 1000 m. The White255

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bellied Duiker ranges in the west from S Cameroon (south of the Sanaga R.), through Gabon, Equatorial Guinea, N and SW Congo (where the species is reported in the forest savanna complex along the Atlantic coast), and extreme SW Central African Republic (East 1999, Wilson 2001). The provenance of museum specimens north of the Sanaga R. remains uncertain (P. Grubb, in Lamarque et al. 1990, Grubb et al. 2003). The species is found again with certainty in NE DR Congo in the Ituri Forest, Maniema and North Kivu. It is uncertain if the species ever occurred north of the Mboumu R. into the forest blocks of SE Central African Republic (for example, East [1999] and Wilson [2001] both map its possible occurrence from Bangassou Forest, from whence there are no confirmed records). There are no records from Cabinda (Angola) (Crawford-Cabral & Veríssimo 2005), but it may occur, perhaps ranging into the BasCongo province in DR Congo. The species’ occurrence in DR Congo’s central cuvette is also hypothetical (von Richter et al. 1990, Wilson 2001). Schouteden (1947) shows four records west of about 24° E along the Congo R., including two from locations on the left bank of the river, but it is uncertain if these specimens originated from these locations. A skin reportedly of this species has been reported from Salonga N. P., but no details are known, and the skin was apparently not examined; more recent surveys of animals taken by local hunters in the park failed to produce any evidence of the species (von Richter et al. 1990, Hart et al. 2008). There is no further evidence of the species’ occurrence south of the Congo R. The White-bellied Duiker is not known to range east of the lowland forest block bordering the Albertine Rift, although it was thought possible that it may exist in the Kayonza Forest, in W Uganda (Kingdon 1982). It may still occur in the forests west of the Semliki R. in DR Congo. The range of the White-bellied Duiker, with distinct western and eastern block populations, resembles that of another small forest ungulate, Bates’s Pygmy Antelope Neotragus batesi.

Habitat  The White-bellied Duiker is restricted to areas of extensive, intact closed tropical moist forest. It occurs in upland, terra firma, primary forest and older secondary forests. It is absent from areas of recent clearing and disturbance. The species does not occur in swamp forests and does not range into gallery forest or the forest–savanna ecotone north of the main forest block. Its characterization by Malbrant & Maclatchy (1949) as a common species of secondary forest and savanna ecotone and not of deep forest has not been supported by subsequent research. In the central Ituri Forest, radio-collared individuals frequented areas of open understorey, and tended to avoid dense stands of herbaceous thickets. In the Ituri Forest and the Nouabale–Ndoki area of N Congo this is one of the characteristic species in the monodominant mbau/limbali forests, a closed-canopy formation dominated by Gilbertiodendron dewevrei. Fruit and seed availability in monodominant forests is strongly seasonal with periods of very low availability juxtaposed by brief pulses of abundance during the supra-annual mast flowering and seed production by the dominant G. dewevrei (Hart 1985). In the Ituri Forest, where six species of frugivorous duikers, and the Water Chevrotain Hyemoschus aquaticus co-occur, population densities of all species except the White-bellied Duiker are lower in large monodominant stands than in the more productive, and seasonally less variable, mixed-canopy formations (Hart 1985, 2001). For Whitebellied Duikers, the monodominant forest appears to represent a competitive refuge from the more densely occupied mixed-forest areas, where combined densities of all small ungulates can reach 45–50/km2. In large monodominant stands, in contrast, total small ungulate densities are about half this value, and the White-bellied Duiker’s nearest competitor, the similarly sized, diurnal Weyns’ss Duiker Cephalophus weynsi, is absent. Abundance  A little known species over much of its range, the White-bellied Duiker is reported as uncommon or rare in most areas where it occurs. Dubost (1984) reported the species comprising less than 11% of all individuals reported in hunter catches in E Gabon (Makokou area). Elsewhere in Gabon, the species is most frequently reported in Lopé N. P., but is uncommon and poorly known elsewhere (Blom et al. 1990). White-bellied Duikers were represented by only 2 of 198 skulls collected in SW Central African Republic (Wilson 2001). Colyn et al. (1987) report the species as the least common among duikers in surveys of the Kisangani bushmeat markets. The species appears to be more common in the Ituri Forest where densities of unhunted radio-collared populations averaged about 4.5/km2 in mature mixed forest and about 6/km2 in monodominant Gilbertiodendron dewevrei forest, contributing about 15% and 35% of the total small ungulate biomass in mixed and monodominant forest, respectively (Hart 2000, 2001). East (1999), assuming average population densities of 2.0/km² where it is known to be common/abundant and 0.2/km² elsewhere, estimated the total population at 287,000 individuals. Adaptations  The White-bellied Duiker is a diurnal species that occupies large home-ranges. The species has long legs, a gracile build and a broadened, but lightly built dental arch and palate. These adaptations are associated with a trophic niche specifically adapted to widely dispersed high-quality food patches. This specialization provides the White-bellied Duiker with a

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relative advantage over more generalist duikers in monodominant forests where food resources can be uncommon and patchy, and are successfully exploited by the duiker that ‘gets there first’. In the Ituri Forest, radio-collared adults ranged widely over annual home-ranges that averaged 63 ha in mixed forest and 58 ha in mbau forest. These home-ranges were comparable in area with those of the larger nocturnal Bay Duiker, and nearly twice the area of the similarly sized diurnal Weyns’s Duiker. Radio-collared animals regularly patrolled widely dispersed, known fruit sources, and frequently accompanied troops of primates. While other duikers also feed beneath primates, White-bellied Duikers actively seek and follow moving troops to a much greater extent than other more sedentary duiker species. They feed not only on fallen fruit parts, but more remarkably are coprivorous, foraging selectively on primate faeces when these contain concentrations of nutrient-rich seeds, and in particular Landolphia spp. The relatively broad mouth and wide maw indicate that the species is capable of ingesting large-sized fruits. However, unlike the Bay Duiker, which has a still broader toothrow, the dentition of the Whitebellied Duiker is less robust, and lacks the developed cranial supports for expanded musculature to deal with coarse foods. It would seem that aggressive intra-specific conflict is sufficiently rare given the light build of the skull and lack of reinforced cranial boss. The wide ranging and wandering movements of the Whitebellied Duiker would appear to obviate the need for armoured defences typical of some of its immediate competitors, such as Weyns’s Duiker, which occupy smaller more intensively defended territories. Foraging and Food  The diet is dominated by fruits and seeds. Gautier-Hion et al. (1980) and Dubost (1984) recorded that fruit comprised almost three-quarters of stomach contents collected in Gabon. In captive feeding trials, White-bellied Duikers invariably selected fruits over foliage, even when these contained relatively higher levels of fibre (Molloy & Hart 2002). Hart (1985) found the White-bellied Duiker to be among the most dedicated frugivore of all the forest duikers in the Ituri Forest, with ripe fruits, unripe fruits and edible seeds comprising from 80% to 100% of rumen contents in all but the most extended periods of fruit and seed dearth. Seasonally dominant dietary items included seeds of Landolphia spp., and the unripe and still soft aborted fruits of Kalinedoxa gabonensis and Irvingia grandifolia. These relatively uncommon and ephemeral, but high-quality, foods were less frequently found in the diets of other duiker species. Like other duikers, White-bellied Duikers feed on caesalpinaceous mast when this is available. Fungi and fallen flowers picked up from the forest floor dominated diets during periods of low fruit availability. In common with other frugivorous duikers, the White-bellied Duiker disperses seeds of a number of species of forest trees (Feer 1995). These tree species have characteristically large, heavily armoured seeds, which duikers spit out undamaged when they ruminate. A number of these trees appear to be specialized for ungulate dispersal and the fruits are not regularly eaten by other frugivores. In the Ituri Forest, two of these species, Donnella pruniformes and another unidentified Sapotaceae, were characteristically eaten and seeds dispersed by White-bellied Duikers. Both tree species are uncommon, and produce large, soft, latex-rich fruits in small numbers over extended periods that are

optimally exploited by this species’ wide-ranging, patrol-foraging strategy. Social and Reproductive Behaviour  Male and female home-ranges overlap, and pairs are perhaps drawn together by the noise made by the primates that both sexes seek to accompany. Nevertheless, sexes often range separately. Radio-collared juveniles often travelled in company with their mothers; however, both sexes disperse from maternal home-ranges at the onset of sexual maturity. Reproduction and Population Structure  A wide seasonal spread of birth and pregnancy records in the Ituri Forest implies that some breeding goes on throughout the year. Seven of nine estimated birth months occurred during the late rains (Aug–Nov), when fruits are seasonally most abundant, a pattern also found in some other duikers in Gabon (Dubost & Feer 1992). In the Ituri Forest, about 60% of 20 adult // examined were pregnant. Gestation period has been estimated at six months (Dubost & Feer 1992). Females produce a single young. The weights of two full-term foetuses were 1300 and 1500 g. Predators, Parasites and Diseases  White-bellied Duikers are killed by Leopards Panthera pardus in the Ituri Forest (Hart, J. A. et al. 1996); however, the species is not among the most selected prey. Per capita annual predation of radio-collared White-bellied Duikers was lower than for nocturnal duikers, such as Bay Duikers and Yellowbacked Duikers C. silvicultor (Hart 2000). Serological surveys of duikers in the Ituri Forest revealed potentially significant exposure to bluetongue, epizootic haemorrhagic disease, infectious bovine rhinotracheitis and leptospirosis (Karesh et al. 1995). This study also recorded several ixodid tick species, including Haemaphysalis parmata, Ixodes cumulatimpunctatus and a Rhipicephalus sp. Conservation  IUCN Category: Least Concern. CITES: Not listed. White-bellied Duikers appear to be rare or declining in many areas of their range. The species is extremely vulnerable to overhunting and is one of the first species to drop out of the small ungulate fauna in areas subject to heavy snare pressure. Some populations may have become locally extinct in recent time with the advent of uncontrolled hunting and the bushmeat trade. In Ituri Forest, indigenous Mbuti hunters regularly catch White-bellied Duikers on net drives. Since the persistence of hunting is often determined by the frequency of capture of more abundant species such as the Blue Duiker Philantomba monticola, White-bellied Duikers could be hunted to local depletion or extirpation, even when already rare (Hart 2000). The species’ wide-ranging movements over large home-ranges expose it to higher risk by snare capture than more sedentary species. Important protected areas for this species include Lopé N. P. (Gabon), Monte Alén N. P. (Equatorial Guinea), Dzanga-Ndoki N. P. (Central African Republic), Odzala N. P. and Nouabalé-Ndoki N. P. (Congo) and Kahuzi-Biega N. P., Maiko N. P. and Okapi Faunal Reserve (DR Congo) (East 1999). Management of the bushmeat trade in Okapi Faunal Reserve, a stronghold for White-bellied Duikers, would improve overall prospects for the species’ conservation (Wilkie et al. 1998). 257

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Measurements Cephalophus leucogaster HB (?): 1050 mm, n = 1 T (?): 105 mm, n = 1 Sh. ht (?): 490 mm, n = 1 WT (??): 16.8 (14.7–18.1) kg, n = 15 WT (//): 18. 9 (16.8–21.0) kg, n = 23 Ituri Forest, DR Congo (J.A. Hart pers. obs.) WT (??): 15.5 (14–17) kg, n = 5

WT (//): 16.8 (15.5–17.5) kg, n = 8 N Congo (Wilson 2001) The record pair of horns measured 12.7 cm from Sette Cama, Gabon (Rowland Ward) Key References  Hart 1985, 2000, 2001; Wilson 2001. John A. Hart

Cephalophus natalensis Natal Red Duiker Fr. Céphalophe du Natal; Ger. Natal-Rotducker Cephalophus natalensis A. Smith, 1834. S. Afr. Quart. J. 2: 217. South Africa, KwaZulu–Natal, ‘Port Natal’ (Durban).

Lateral view of skull of Natal Red Duiker Cephalophus natalensis.

Natal Red Duiker Cephalophus natalensis.

Taxonomy  In a revision of the duikers, Grubb & Groves (2001) included Harvey’s Duiker C. harveyi as a single subspecies of the Natal Red Duiker C. natalensis (and see Jansen van Vuuren & Robinson 2001 and Hassanin et al. 2012). Ansell (1972) considered Harvey’s Duiker C. harveyi and Weyns’s Duiker C. weynsi as conspecific with C. natalensis and he offered a provisional list of 11 subspecies (including ‘walkeri’ from Malawi, which was actually a Sylvicapra; see Grubb 1988, 2005). Excluding these extralimital or mistaken forms, Ansell still retained four described subspecies of C. natalensis (as defined here). Following Swynnerton & Hayman (1951), Kingdon (1982, 1997) and East (1999), we provisionally treat C. natalensis and C. harveyi as separate species and recognize two subspecies within C. natalensis (following Meester et al. 1986). Synonyms: amoenus, bradshawi, lebombo, robertsi, vassei. Chromosome number: 2n = 60 (Robinson et al. 1996b). Description  A small duiker, often giving a first impression of being an all-over, uniform red. Although there is substantial regional and individual variation, the upperparts of the body are typically a deep chestnut-red, the lower part of the flanks and underparts pale chestnut, and there is a prominent crest of long, bushy, chestnut-coloured hair on the top of the head, which often conceals the horns. The sides of the muzzle, the sides and underparts of the neck, and the inner upper

surfaces of the limbs range from tawny or pale fawn to a rich red, only marginally lighter than the upperparts.The lower jaw behind the black or brown chin is always white and white or cream sometimes extends onto the upper throat. Ears short and rounded with a fringe of black hair on the outside margins; insides with three alternating black and white, rather ‘smudged’ markings.The back of the neck varies in colour with animals from the southern part of the range tending to uniform red while those in the north are more variable, sometimes having the entire nape light or dark grey. The pasterns and hind part of the hocks are also variable, sometimes red but more commonly darker in colour, with a tinge of dull violet or grey. The upperparts of the tail are the same colour as the back, darkening towards the whitish tip. Preorbital glands are present as a perceptible swelling below and in front of each eye. The gland exudes secretion along a streak of black skin that is studded with a row of small pores (Mainoya 1978); the secretion is a clear sticky fluid with a faint aromatic odour, the chemical constituents of which have been investigated by Burger et al. (1988). Interestingly, the preorbital gland secretions of the Natal Red Duiker and Common Duiker Sylvicapra grimmia contain two major constituents absent from the preorbital gland secretion of the Blue Duiker Philantomba monticola. Pedal glands are present, and discussed in detail by Mainoya (1978); there are no inguinal glands. Horns, present in both sexes (those of ?? twice the length of //), are short, straight, with coarse basal rings and longitudinal striations, but smooth towards the tips. Geographic Variation C. n. natalensis (including amoenus, lebombo): S and NE KwaZulu– Natal, E Mpumalanga and possibly S Mozambique. General description as above with strong erythristic tendency. C. n. robertsi (including bradshawi, vassei): Mozambique, north of the Limpopo R. northwards through S Malawi to SETanzania. Specimens

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from Mozambique tend to be more orange/rufous in colour on the upperparts of the body than their counterparts from KwaZulu– Natal. In addition, the head is darker and there are slight traces of a dark frontal blaze, not present in animals from KwaZulu–Natal (Wilson 2001). Grey napes are also more common (J. Kingdon pers. comm.). Comparing specimens from lowland S Tanzania (south of the Rufiji R.) with specimens of C. natalensis from many other parts of their range, Swynnerton & Hayman (1951) had no hesitation in allocating them to C. n. robertsi.

south as Pondoland (Bowland 1990) and even to George (Skead 1980); Rowe-Rowe (1994) shows the species occurring in small isolated populations along the coast south to about Oribi Gorge at about 31° S. Today they probably do not occur much further south than Durban.There are no confirmed records of this species from Zimbabwe (Smithers & Wilson 1979, Wilson 2001). East (1999) shows the species occurring in NE Zambia and N Malawi, but these records relate to Harvey’s Duiker, the Natal Red Duiker being known only from S Malawi (Ansell 1978, Ansell & Dowsett 1988). Red duikers from the area between the Ruvu and Rufiji valleys in E Tanzania approach C. harveyi in colouring and this area may represent a hybrid zone (Kingdon 1982). A similar hybrid frontier is implied by Wilson’s (2001) observations on the Nyika Plateau in N Malawi where animals he observed looked like intermediates between C. natalensis and C. harveyi. The relationship between these very closely related duikers awaits a scientifically rigorous resolution.

Similar Species Cephalophus harveyi. Larger red duiker ranging from N Malawi, NE Zambia, through suitable forested areas of C and NE Tanzania to SE Kenya and S Somalia. Legs dark grey to brownish-black; midfacial zone blackish; crest with more black in it. Bolder, more strongly contrasting facial and ear patterns. For intermediatelooking animals along the interface between C. natalensis and C. harveyi, see discussion in next section. Habitat  Throughout their range, Natal Red Duikers are associated with forests and dense thickets, occurring in riverine forest and in Distribution  Endemic to Africa. The Natal Red Duiker formerly thickly wooded ravines and dense coastal bush. In NE KwaZulu– occurred widely in coastal and riverine forests and thickets from SE Natal they occur up to about 200 m elevation (Rowe-Rowe 1994). Tanzania to NE KwaZulu–Natal. The species’ distribution within this They are independent of water; in KwaZulu–Natal, only one of range is patchy and discontinuous and the clearing of forests and the home-ranges of the duikers studied by Bowland (1990) had a thickets to make way for agricultural development has destroyed perennial water supply. much suitable habitat. Natal Red Duikers still occur locally in forest patches and coastal scrub in SE Tanzania, while in Mozambique they Abundance  The only reliable density estimates available for are still believed to occur widely in the eastern, coastal parts of the Natal Red Duikers are from KwaZulu–Natal, where Bowland (1990) country, as well as in a few scattered localities inland; they are recorded densities varying from 1 ind/0.5–1.0 ha in favourable reported to occur at high densities in Maputo G. R.Their distribution habitat (such as at St Lucia) to 1 ind/2.5–5.0 ha in less favourable range extends southwards through Mpumalanga and the Limpopo areas. East (1999) estimated the total population to stand at about Province of South Africa (in the Soutpansberg), Swaziland, to 42,000, but noted this could be an underestimate. Rowe-Rowe KwaZulu–Natal, where they occur mainly in the north-eastern parts (1994) gave the population size for KwaZulu–Natal as between 2000 (East 1999, Wilson 2001, Skinner & Chimimba 2005). The and 3000 animals; the largest population in this province occurs in southernmost limit of distribution has been given as extending as far Greater St Lucia Wetland Park (1000). A substantial part of the range of this species occurs in Mozambique, where an overall recovery in wildlife populations is likely to include this duiker.

Cephalophus natalensis

Adaptations  Natal Red Duikers are diurnal, with peaks in activity at dawn and dusk, and little evidence of nocturnal activity (Bowland 1990, Wilson 2001), a pattern that is confirmed by the frequency of their bones in the debris below nests of the diurnal Crowned Eagle Stephanoaetus coronatus in S Tanzania (J. Kingdon pers. comm.). They are shy and secretive, and if disturbed seldom stand for more than a few seconds before bounding away into thick bush, often emitting a hoarse alarm ‘whistle’. They are said to display a remarkable jumping ability; De Vos (1979) reported an instance of two adults clearing a 1.6 m high fence. Metabolic rates are high, and energy requirements are met by utilizing easily digestible and fermentable plant parts while less digestible parts (lignin, cellulose) are poorly digested due to the fast digesta passage rates (Faurie & Perrin 1995, Perrin et al. 2003). The rumen has been described as similar to that of the Blue Duiker, displaying many adaptations typical of concentrate selectors, including a densely, evenly papillated rumen for maximum nutrient absorption (Faurie & Perrin 1995). High rates of food intake, frequent rumination and rapid fermentation ensure efficient supply of energy and nutrients (Perrin et al. 2003). 259

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Foraging and Food  Natal Red Duikers are browsers and frugivores, feeding primarily on leaves (fresh and fallen), wild fruits and flowers, and shoots of low-growing shrubs. An analysis of the contents of 19 stomachs from KwaZulu–Natal found that dicotyledonous leaves had a relative occurrence of 66%, seeds and fruits 25% and flowers 1% (Faurie & Perrin 1993, Bowland & Perrin 1998). Although Gagnon & Chew (2000) reported a very small percentage (1%) of monocotyledons in the diet (and see Sponheimer et al. 2003b), Prins et al. (2006) recorded between 15% and 20% of monocots in the diet. Bowland (1990) directly observed Natal Red Duikers eating the freshly fallen leaves of Cussonia sphaerocephala, Strychnos spinosa, Celtis africana, Ficus stuhlmanii, Sapium integerrimum, Harpephyllum caffrum, Barringtonia racemosa, Apodytes dimidiata, Acacia robusta and Ziziphus mucronata. Heinichen (1972) recorded them feeding on Dichrostachys cinerea, Strychnos madagascariensis, Gardenia cornuta, Azima tetracantha, Asparagus falcatus and Justicia protracta. Other species recorded from the examination of stomach contents include Justicia sp., Commelina africana and Grewia sp. (Heinichen 1972). Elevated water and protein content of leaves influence the food choice of Natal Red Duikers, but there is no evidence that condensed tannin concentrations influence leaf choice, since some highly preferred species have tannin concentrations exceeding 18% while other rejected leaves have concentrations of less than 2% (Faurie & Perrin 1993, Bowland & Perrin 1998, Perrin et al. 2003). Wilson (2001), after Bowland (1990), provides a detailed list of dietary items in the diet of Natal Red Duikers recorded in southern Africa. Prins et al. (2006) investigated niche segregation among three small antelopes – Natal Red Duikers, Common Duikers and Sunis Nesotragus moschatus – in a coastal savanna woodland/forest mosaic in S Mozambique to determine whether they exhibited any obvious differences in diet to avoid competition. Some 80 dietary items were recorded being used by Natal Red Duikers. On average, only about 10% of the food items used by all three small bovids were specific to one species only. Diet overlap was considerable in the wet season, but the use of exclusive species was significantly larger in the dry season, and significantly larger for the Natal Red Duikers (Prins et al. 2006). Likewise, Blue and Natal Red Duikers are sympatric in many localities, with home-ranges of the two species often overlapping. Both generally have similar diets, and they exhibit no temporal separation since both are diurnal, with only subtle differences in spatial utilization (e.g. Natal Red Duikers sometimes forage beyond the forest margin) (Perrin et al. 2003). Social and Reproductive Behaviour  Natal Red Duikers are primarily solitary, although it is not uncommon to encounter a / with her offspring, or pairs or small groups of three to five (?? sometimes form temporary associations in the absence of a /). Home-ranges usually cover an area of 2–15 ha in size, and decrease in size with an increase in population density and availability of food resources. Communal dung heaps are used to demarcate the boundaries. The secretion from the preorbital glands is also dabbed onto stems and branches as a means of marking the home-range. However, there is no evidence of territoriality (as in Blue Duikers) and home-ranges may overlap by as much as 80–100%; there was also extensive overlapping of core areas of home-ranges (Bowland 1990, Bowland & Perrin 1995).

Vocalizations include an alarm whistle, and a loud ‘tchie-tchie’ sound, which is louder and more penetrating than that of the Suni (Heinichen 1972). Natal Red Duikers are known to ‘thump’ or ‘alarm stamp’ prior to fleeing. Natal Red Duikers have been recorded in association with Vervet Monkeys Chlorocebus pygerythrus; the monkeys were seen actively grooming a duiker, which itself solicited grooming on several occasions (Borland & Borland 1979). Reproduction and Population Structure  Breeding occurs throughout the year, and three-month-old lambs have been observed in Feb, Jul and Aug (A. E. Bowland pers. obs.); in captivity, young are also born throughout the year, with no discernible peak (Spence 2003). A single young is born; captive animals had an average mass at birth of 979 g (n = 2) and an inter-birth interval of 236 days (range 222–273, n = 5; Spence 1991). Gestation period is in the order of 7 months. Bowland, in Rowe-Rowe (1994), gives age at first conception at 18–24 months, but a captive / conceived when just under a year old (Spence 1991). Bowland (in Rowe-Rowe 1994) gives a potential life-span of 8–9 years;Weigl (2005) gives a longevity record in captivity of 15 years. Predators, Parasites and Diseases  Natal Red Duikers, in particular infants or juveniles, are likely to fall prey to a number of medium to large carnivore species (e.g. Roche 2003), as well as birds of prey (such as the Crowned Eagle; Wilson 2001) and African Rock Pythons Python sebae. Very little was known of the helminth parasites of Natal Red Duikers until the studies of Boomker and coworkers in KwaZulu–Natal (Boomker et al. 1984, 1991d). The latter authors summarized what was known at the time, listing 17 species of nematodes, two species of cestodes (Moniezia benedeni and Stilesia hepatica) and trematodes (paramphistomes). Ectoparasites in Natal Red Duikers have been investigated by various authors, primarily in KwaZulu–Natal; Horak et al. (1991a) examined 20 duikers from two reserves in NE KwaZulu–Natal and recorded the following ixodid ticks: Amblyomma marmoreum, Haemaphysalis leachi, H. parmata, Rhipicephalus appendiculatus, R. maculatus, R. muehlensi and R. evertsi. In addition, they recovered two species of lice (Damalina sp. and Linognathus sp.). Conservation  IUCN Category: Least Concern. CITES: Not listed. Natal Red Duikers have disappeared from large parts of their former range, largely as a result of the loss of suitable habitat in the face of expanding human settlement and agriculture, combined with the impacts of hunting. None the less, despite what is likely to be a gradual population decline across much of their range, they are not likely to be threatened as a species, and remain well represented by healthy populations in a number of protected areas, such as Selous G. R. (Tanzania), South Vipya F. R. (Malawi), Maputo G. R. (Mozambique) and Greater St Lucia Wetland Park, HluhluweiMfolozi and Ndumo G. R. (South Africa) (East 1999). Measurements Cephalophus natalensis TL (??): 864 (809–900) mm, n = 9 TL (//): 890 (855–950) mm, n = 8 T (??): 96 (70–115) mm, n = 8

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T (//): 103 (85–125) mm, n = 8 HF c.u. (??): 231 (220–245) mm, n = 9 HF c.u. (//): 227 (220–240) mm, n = 8 E (??): 77 (74–81) mm, n = 9 E (//): 77 (72–80) mm, n = 8 Sh. ht (??): 412 (380–480) mm, n = 9 Sh. ht (//): 418 (385–450) mm, n = 8 WT (??): 11.7 (9.8–12.6) kg, n = 9 WT (//): 11.9 (10.3–13.2) kg, n = 9 KwaZulu–Natal (A.E. Bowland pers. obs.)

E (??): 78 (74–82) mm, n = 5 E (//): 75 (73–77) mm, n = 7 Sh. ht (??): 419 (390–460) mm, n = 5 Sh. ht (//): 420 (388–460) mm, n = 7 WT (??): 11.9 (9.8–12.7) kg, n = 5 WT (//): 12.0 (9.9–13.6) kg, n = 7 Mozambique (Wilson 2001) Maximum recorded horn length is 10.4 cm for a pair of horns from Mufindi, Tanzania (Rowland Ward), but these measurements undoubtedly refer to Harvey’s Duiker C. harveyi.

TL (??): 869 (805–903) mm, n = 5 TL (//): 910 (850–920) mm, n = 7 T (??): 96 (80–106) mm, n = 5 T (//): 100 (84–114) mm, n = 7 HF c.u. (??): 230 (220–240) mm, n = 5 HF c.u. (//): 235 (219–240) mm, n = 7

Key References  Bowland 1990; Bowland & Perrin 1994, 1995, 1998; East 1999; Heinichen 1972; Skinner & Chimimba 2005; Wilson 2001. Michael Hoffmann & Anthony E. Bowland

Cephalophus harveyi Harvey’s Duiker Fr. Céphalophe de Harvey; Ger. Harveyducker Cephalophus harveyi (Thomas, 1893). Ann. Mag. Nat. Hist., ser. 6, 11: 48. Kahe Forest, Moshi District, Tanzania.

Natal Red Duiker, commenting that the red duikers he saw in Selous G. R. were very similar to these duikers. Synonyms: bottegoi, keniae. Chromosome number: not known, but likely to be 2n = 60 as in C. natalensis (B. Jansen van Vuuren pers. comm.). Description  A richly russet-red duiker with the red of the rump and back becoming lighter below. The dorsal portion of the head shows a very distinct black band that covers the bridge of the nose and becomes wider above the eyes. The centre of the crest is also black and this line is continued down the back of the neck where it gives way to a dark freckle. In southern parts of its range this melanic tendency extends to the dorsal portion of the neck, shoulders and, in the same area, the legs can be entirely dark sepia brown. Hundreds of camera-trapping photographs of Harvey’s Duikers from several sites in the Udzungwa Mts show that there is large variation in the extent of these dark portions of the pelage between individuals within the same area (Rovero et al. 2005, F. Rovero pers. obs.). By contrast, Harvey’s Duiker Cephalophus harveyi. animals from Mt Kenya tend to be more consistently red. As with the Natal Red Duiker, preorbital glands are embedded in a large swelling Thomas named the species for Sir Robert G. Harvey, a member of below and in front of each eye, the external expression of which the hunting party which took the type specimen. comprises a black streak of skin studded with a row of small pores (Mainoya 1978). Pedal glands are present, and discussed in detail by Taxonomy  Treated as a subspecies of Natal Red Duiker C. Mainoya (1978); there are no inguinal glands. natalensis by some authors (Heyden 1968, Ansell 1972, Groves & Grubb 1981, Grubb & Groves 2001, Wilson 2001, Grubb 2005; Geographic Variation  Populations in the northern part of their and see Jansen van Vuuren & Robinson 2001 and Hassanin et al. range (Kenya and N Tanzania) differ from those in SC Tanzania, but 2012), Kingdon (1982) considered that subsuming this very the boundaries or clinal areas between them remain to be elucidated. distinctive population in C. natalensis risked obscuring the complex It may eventually be possible to characterize these two populations evolutionary history of red duikers. None the less, likely hybrids as subspecies. Three names have been applied to this duiker: harveyi between C. harveyi and C. natalensis are known from some of the from Kilimanjaro, keniae from Mt Kenya and bottegoi from lowland lowland areas where their ranges appear to overlap (Kingdon coastal forest on the extreme northern edge of its range. These 1982). Wilson (2001) considers animals from N Malawi on the names should be treated as synonyms until their taxonomic status Nyika Plateau as intermediate between Harvey’s Duiker and the has been assessed. 261

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Lateral view of skull of Harvey’s Duiker Cephalophus harveyi.

Similar Species Cephalophus nigrifrons. Lowland forest block and montane isolates on Mt Kenya, Mt Elgon and the Aberdares. Long-legged, longhooved species with dark, finely freckled pelage. C. natalensis. A primarily southern African duiker, reaching its northerly limit in the Rufiji Valley, S Tanzania. Smaller; legs slightly grey or not at all; facial mid-line in front of eyes darker. C. weynsi. Mainly found in central Africa and Uganda extending as far as the Mau forest in Kenya. A larger species with longer, more arched muzzle and duller orange-red colouring. Kingdon (1982) suggested some red duikers from the Mau massif in Kenya might be hybrids between Weyns’s Duiker and Harvey’s Duiker. C. adersi. Restricted to Zanzibar and the East African coast (notably the Arabuko-Sokoke Forest and Boni-Dodori forests, but also, perhaps, other East African isolated coastal forest pockets). Smaller, with white underbelly, white freckling on the lower limbs, and a broad white band on the rump. Distribution  Endemic to Africa. Patchily distributed through various forest types in the moister eastern parts of C and SE Kenya and NE and C Tanzania and, marginally, in extreme S Somalia (including, formerly, the lower Shebelle and Juba Rivers where now probably extinct) and N Malawi (Wilson 2001). There is a single record of this species from the Zambian side of the Nyika Plateau (Ansell 1978; mapped by East [1999] as C. natalensis). Red duikers seen in the Harenna Forest on the Bale massif (Hillman 1988a, Yalden et al. 1996) and another population in Omo N. P. in SE Ethiopia (Schloeder & Jacobs 1996) probably represent this species (East 1999), although the Omo duikers could be Weyns’s Duiker C. weynsi, which occurs in the Imatong Mts of S Sudan, some 400 km to the south-west of Omo (East 1999, Wilson 2001; and see Grubb et al. 2003). Habitat  Found in both moist coastal forests and riverine gallery forests in the eastern parts of its range. It ascends into montane forests on Mt Kenya, the Aberdares and the Eastern Arc Mts in Kenya and Tanzania (see below). There it is found in both drier and moister forest types but predominantly in the vicinity of permanent water. In the Udzungwa Mts, Tanzania, it has been recorded in miombo woodland at 400 m; in dry, Commiphora-dominated bush at 1400 m; also up to about 2200 m in mixed upper montane forest and bamboo thickets (F. Rovero pers. obs.) and may well range to the highest forested slopes of Mt Luhomero to about 2400 m. It has also been reported as widespread in the highest Forest Reserves of the Uluguru Mts, which range to over 2400 m (Doggart et al. 2004). Abundance  Data scarce, but observations suggest that the species may be more abundant on the moister eastern or windward sides of

Cephalophus harveyi

massifs and scarcer on the drier western rain-shadow sides. Assuming an average population density of 2.0/km² in favourable areas and 0.2 in other parts of its range, East (1999) estimated a total population of 20,000 individuals, but notes that this could be an underestimate. A sharply downward trend in the overall population is evident in the rapid clearance and human settlement of its preferred riverine habitat. Extensive camera-trapping and line transect censuses conducted in the Udzungwa Mts indicated high variation in the density of Harvey’s Duikers even within relatively close and similar forest sites, presumably related to a combination of past and current hunting pressure and habitat features. Highest densities of about 13.3/km2 were recorded in Matundu Forest in ground-water semi-deciduous forest at 300–500 m within Udzungwa Mountains N. P. However, in areas of the same forest where hunting had occurred, the density decreased to 8.2/km2, and in the highly disturbed Uzungwa Scarp forest density dropped to about 2.1/km2 (Rovero & Marshall 2009). Similarly, in Mwanihana Forest, at an altitude of 300–1000 m, the density resulted in the range of 4.8–9.5 ind/km2 for moderately disturbed and undisturbed forests within the National Park (Rovero & Marshall 2009). These results confirm that East’s (1999) average density estimations for favourable areas might be underestimates and that more data are required before a sound, overall estimation can be attempted. Adaptations  Contemporary molecular studies suggest that the closest relatives of the harveyi/natalensis group are the swamp-forest adapted Black-fronted Duiker C. nigrifrons and woodland adapted Red-flanked Duiker C. rufilatus (Jansen van Vuuren & Robinson 2001). Since both the latter species have moved into habitats that are atypical of other duikers, the very generalized Harvey’s Duiker could be a passable model for their common ancestral type. Until very recently Harvey’s Duikers were abundant in the montane forests of East Africa and neighbouring coastal areas. This abundance and the absence of a clearly defined habitat preference confirms that this species fills the niche of medium-sized frugivorous antelope within

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Harvey’s Duiker Cephalophus harveyi.

this area. Over the greater part of its range this species co-exists with the much smaller Blue Duiker Philantomba monticola and, in some parts of its montane range, is sympatric with the diminutive Suni Nesotragus moschatus and the much larger Abbott’s Duiker, C. spadix. All of these species appear to occupy definably different niches. Where the range of Harvey’s Duikers abuts that of the slightly larger Weyns’s Duiker in Kenya, and that of the rather smaller Natal Red Duiker in Tanzania, hybridization appears to take place. In both instances these neighbours not only resemble Harvey’s Duikers in general morphology, but they clearly occupy the same generalized red duiker niche. Supposing the hybridizations with both neighbours are real there could be active competition between these three populations along the boundaries of their ranges. This suggests that there could be a long-term dynamic in which some species are expanding while others may be retreating. Pre-occupancy of a region confers some advantages on wellestablished populations but in any situation where there are advantages in being larger or smaller it can be predicted that one or other neighbour gains advantage and can expand its range. Overall faunistics suggest that Weyns’s Duiker has made a relatively recent eastward expansion. It is therefore possible that this expansion was

at the expense of Harvey’s Duiker. It is less easy to identify winners and losers along the C. harveyi/natalensis front, but overall trends support the idea that Harvey’s Duiker could be in an early stage of becoming an exclusively montane species, while the Natal Red Duiker is becoming entrenched as the dominant lowland red duiker, actively expanding its range northwards. If this is the situation with Harvey’s Duiker, it becomes important to maintain awareness of its distinctness as a population and pay attention to what is happening along its borders with the Natal Red Duiker and Weyns’s Duiker. Likewise, an already marginal status for Aders’s Duiker C. adersi (before it was threatened by settlement and hunting) could relate to ancient long-term competition with Harvey’s Duiker. Foraging and Food  Harvey’s Duikers are browsers, selecting almost exclusively a dicotyledonous diet with a high preference for leaves and fruit. Leaves from bushes comprised about 90% of rumen contents from three specimens in Kenya (Hoppe et al. 1981; and see Cerling et al. 2003). At times large quantities of yellow sodom apples (Solanum incanum) are eaten that are often swallowed intact (Hofmann 1973). Leaves, seeds and pods of Acacia spp., and leaves and fruits of 263

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the bushes Acokanthera frisiorum and Warburgia ugandensis are also eaten. Parinari excelsa is dominant in some montane areas and Ficus, Cordia and Lannea spp. are widespread; all are known foods. Within its range the following are known sources of fruit and/or leaves: Annona senegalensis, Anthocleista grandiflora, Tabernaemontana pachysiphon and species of the genera Bridelia, Canthium, Coccinia, Diospyros, Euclea, Grewia, Mammea, Parkia, Poliscias, Rhus, Sapium, Strychnos, Trema and Uapaca (Kingdon 1982, T. Jones & F. Rovero unpubl.). Digestion in the forestomachs has not yet been investigated. However, the strict preference for dicotyledonous leaves and fruits likely results in a high rate of volatile fatty acid production in the rumen. This is indicated by anatomical features such as a dense papillation of the mucosa to enhance absorption and inconspicuous thin rumen pillars (Hofmann 1973) resulting in a short rumen retention time, and a paucity of rumen protozoa (Hoppe et al. 1981). This contrasts with a surprisingly large capacity of the rumen, which is atypical for dicotyledon selectors and more typical of grass-eating antelopes. Harvey’s Duiker is primarily diurnal. In the Udzungwa Mts, activity pattern of this species, as indicated by camera-trapping times, resulted in 98% of more than 350 photographs being taken between 07:00h and 20:00h. Activity peaked between 07:00h and 10:00h and between 15:00h and 18:00h, and decreased significantly between 12:00h and 15:00h (F. Rovero pers. obs.). Social and Reproductive Behaviour  Little is known, but the species resembles other red duikers in being mostly solitary, although Wilson (2001) observed three adult male red duikers (here considered to be Harvey’s) together on the Nyika Plateau; in the same region, he recorded an adult Gentle Monkey Cercopithecus mitis grooming a female duiker. The solitary habit of this duiker has been confirmed from both direct observations and camera-trapping in the Udzungwa Mts, with only occasional (fewer than 10%) observations and photographs of pairs (T. Jones & F. Rovero unpubl.). Harvey’s Duikers have often been seen foraging below trees occupied by colobus monkeys and moving in association with the predominantly ground-foraging Sanje Mangabey Cercocebus sanjei; they are also groomed by this mangabey species (T. Jones & F. Rovero unpubl.). Other than facilitating access to food resources (i.e. fruits dropped by the monkeys), this behaviour must also increase anti-predator vigilance. As with some other red duikers, Harvey’s Duiker utters a shrill, fluty whistle when disturbed. If these animals live in a mosaic of territories (whether exclusive or overlapping), as is supposed, the most likely function of such calls is to inform conspecific neighbours as to the whistler’s movements as well as alert them to disturbance. It is not known how much information these whistles might contain about the identity of the caller or the nature of the disturbance. Reproduction and Population Structure  Wilson (2001) recorded newborns on the Nyika Plateau in Dec, Mar, Aug and Feb. There is no other published information. Predators, Parasites and Diseases  Known predators include Leopards Panthera pardus, notably in the Udzungwa Mts (Rovero et al. 2005). Predation by Crowned Eagles Stephanoaetus coronatus has been recorded in the Udzungwa Mts (F. Rovero & A. Bowkett unpubl.); other potential predators are Lions Panthera leo and Spotted Hyaenas Crocuta crocuta. Round (1968) documented cestodes from the genera

Moniezia and Cysticercus and a nematode (Setaria sp.) from Harvey’s Duiker. Hoppe et al. (1981) recorded a number of species of the protist Entodinium. Conservation  IUCN Category: Least Concern. CITES: Not listed. Harvey’s Duikers are declining due to hunting and destruction of habitat. East (1999) notes that this species is heavily hunted throughout its Tanzanian and Kenyan range, where animals are hunted with dogs in several areas; wire snares are probably the largest cause of mortality through most of its range. The species has lost most of its habitat in Somalia along the lower Shebelle and Juba Rivers. Clear evidence of decreased abundance related to habitat degradation and hunting resulted from comparison of line-transect and camera-trapping data across contrasting sites in the Udzungwa Mts (Rovero & Marshall 2004, F. Rovero pers. obs.). Important protected areas with known populations of Harvey’s Duikers include Aberdare N. P., Mt Kenya F. R., Arubuko-Sokoke, Boni, Dodori, Tana River and Shimba Hills National Reserves (Kenya), Bush Bush N. P. (Somalia), Bale Mountain N. P. (Ethiopia) and Mikumi, Udzungwa, Kilimanjaro, Lake Manyara and Arusha National Parks (Tanzania) (East 1999, Wilson 2001). Measurements Cephalophus harveyi TL (??): 873 (796–891) mm, n = 3 TL (//): 879 (849–910) mm, n = 4 T (??): 94 (88–96) mm, n = 3 T (//): 98 (90–108) mm, n = 4 HF c.u. (??): 230 (222–239) mm, n = 3 HF c.u. (//): 229 (219–240) mm, n = 4 E (??): 76 (73–80) mm, n = 3 E (//): 77 (74–79) mm, n = 4 Sh. Ht (??): 413 (390–460) mm, n = 3 Sh. Ht (//): 417 (399–449) mm, n = 4 WT (??): 11.3 (9.9–12.6) kg, n = 3 WT (//): 11.9 (9.7–12.9) kg, n = 4 Selous G. R., Tanzania (Wilson 2001) TL (??): 868 (802–913) mm, n = 6 TL (//): 886 (855–917) mm, n = 4 T (??): 96 (82–104) mm, n = 6 T (//): 101 (90–117) mm, n = 4 HF c.u. (??): 227 (220–239) mm, n = 6 HF c.u. (//): 229 (218–240) mm, n = 4 E (??): 78 (74–80) mm, n = 6 E (//): 79 (75–81) mm, n = 4 Sh. Ht (??): 416 (386–470) mm, n = 6 Sh. Ht (//): 419 (385–449) mm, n = 4 WT (??): 11.0 (9.7–13.2) kg, n = 3 WT (//): 11.8 (9.9–13.6) kg, n = 4 Arusha, Tanzania (Wilson 2001) Maximum recorded horn length is 12.7 cm for a pair of horns from Elgeyo Forest, Kenya (Rowland Ward) Key References  Kingdon 1982; Wilson 2001. Jonathan Kingdon & Francesco Rovero

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Cephalophus rufilatus Red-flanked Duiker Fr. Cephalophe à flancs roux; Ger. Rotflankenducker Cephalophus rufilatus Gray, 1846. Ann. Mag. Nat. Hist., ser. 1, 18: 166. ‘Sierra Leone, Village of Waterloo’.

Red-flanked Duiker Cephalophus rufilatus.

Taxonomy  In the nineteenth century, several taxonomists confused this species with Sylvicapra grimmia coronatus (Sclater & Thomas 1899). A densely haired neck and coarse coat ally this species with other coarse-haired red duikers. Jansen van Vuuren & Robinson (2001), using mtDNA, found a consistently close relationship with the Black-fronted Duiker Cephalophus nigrifrons, in spite of very different proportions, morphology and habitat (and see Hassanin et al. 2012). They found a slightly more distant relationship with the Natal Red Duiker C. natalensis, which has similar proportions and dimensions, and the larger Harvey’s Duiker C. harveyi. Two subspecies are usually recognized (St Leger 1936, Ansell 1972, Kingdon 1997), with the Chari R. thought to mark the boundary between them. Grubb & Groves (2001), who did not list subspecies, noted that two types of colouring are found at the extremities of its range: pale ochraceous with dorsal stripe sharply marked in the west, and darker, redder, with dorsal stripe more diffuse towards the eastern end of its range, and that skins from intervening areas show intermediate colouring. Size also varies rather sporadically. Synonyms: cuvieri, rubidior. Chromosome number: 2n = 60; the X chromosome is a metacentric (Hard 1969). Description  A prettily coloured, relatively short-legged duiker with bright orange-red on the face, neck and flanks, dark brown or grey limbs and similarly coloured dorsal ‘patch’. It belongs to the coarse-haired duiker group and its pelage, notably on the neck and throat, resembles that of the Black-fronted Duiker. The shiny black nose is broad and prominent. A dark line down the mid-line of the face is subject to much individual variation in breadth and colour. Likewise, the length and thickness of hair growing vertically from the crown varies individually, but is mostly black in colour. Underside of chin white, posterior surface of ear black, anterior surface lined with short white hairs except for black band close to lower margin. Underside light reddish. Tail with black tuft and its very frequent movements are always from side to side. Long, narrow pedal glands present on both fore- and hindfeet. Inguinal glands absent. Preorbital glands well developed in both sexes. Rostrum is long and narrow, and the frontals are evenly convex as far forward as the proximal nasals, as far back as bases of horns, which are raised up (Grubb & Groves 2001).

Dorsal and lateral views of skull of Red-flanked Duiker Cephalophus rufilatus.

Geographic Variation C. r. rufilatus: Senegal to Chari and Benue Valleys. Back and legs light grey. C. r. rubidior: Chari R. to Nile Valley. Back and legs dark grey. Similar Species Cephalophus nigrifrons. Niger R. to East Africa. A larger, long-legged, long-hooved swamp-forest and mountain dwelling red duiker. Philantomba maxwelli. Senegal to Cross R. A smaller, forest-dwelling duiker of overall dull brown colour. P. monticola. Cross R. to East and south-eastern Africa. A smaller, forestdwelling duiker of overall grey-brown colour. Distribution  Endemic to Africa, ranging from S Senegal, Gambia and Guinea-Bissau through to the Nile Valley in SW Sudan in forest–savanna mosaics north of the main forest block (East 1999, Wilson 2001). Formerly widespread in NW Uganda, as far east as the Albert Nile (Kingdon 1988, East 1999); a small relic population was discovered on the eastern side of the Nile in Bugungu G. R., immediately south of Murchison Falls N. P. (Allan 1996). The boundaries of its range are poorly defined and the species has been recorded in the southernmost regions of several ‘Saharan’ States (SW Mali, SW Niger and extreme S Chad). Not recorded from Liberia. Dowsett (1993) rejected all records from Gabon and Congo, though Grubb et al. (2003) remark that a supposed record of this species from the coastal lowlands of SE Gabon deserves further investigation. Habitat  A diurnal species finding refuge in forest edge, forest relics and riverine thickets, but also commonly emerging into open savanna and woodland. Heringa (1990) reported a distinct preference for riverine woodlands and thick vegetation close to permanent sources of water such as springs within rocky areas. In Nigeria it prefers places with rocky outcrops and forest outliers, even dry valleys within savanna, riparian or forest areas but never in high forest (Agbelusi 1992). In Mole N. P., Ghana, this duiker was 265

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Cephalophus rufilatus

found in open savanna woodland in which tree cover averaged 30% (range 5–65%); the main trees were Burkea africana and Terminalia avicennoides together with other typical woodland trees such as Gardenia, Lannea, Pterocarpus, Piliostigma, Grewia and Isoberlinia spp. (Schmitt & Adu-Nsiah 1993). Wilson (2001) found them to be abundant in open savanna in Central African Republic in a region where there were tens of thousands of small termite mounds. In some regions the replacement of primary forest with farm bush and other secondary vegetation has enabled this species to expand its range (East 1999). Abundance  Data are scarce, but the species is generally widespread over most of its range. Estimates of density vary: Wilson (2001) recorded densities of 3–4/km² in prime habitat in Mole N. P., while Fischer & Linsenmair (2001a) estimated densities of 0.45/km2 in 1995 in Comoé N. P., Côte d’Ivoire (though these declined to 0.14/ km² in 1998). Heringa et al. (1990) estimated densities of less than 0.1/km2 from ground surveys in Burkina Faso. This was the second most commonly observed antelope in the National Park of Upper Niger in Guinea, with a density of 2.6/km², although it had none the less declined by 50% since a previous census in 1997 (Brugière et al. 2005). East (1999) estimated total population numbers at 170,000. Adaptations  The Red-flanked Duiker is unusual in being an inhabitant of relatively open woodlands and savannas, degraded forests and riverine wooded environments. In this adaptation to drier, more exposed environments the Red-flanked Duiker has followed a similar evolutionary trajectory to the Common Duiker Sylvicapra grimmia. However, this adaptation has involved much less departure from the general red duiker morphotype. Molecular studies confirm that the Red-flanked Duiker (and its genetic sibling, the Black-fronted Duiker C. nigrifrons) diverged very much later than the bush duiker, which split away from other species quite early in the duiker radiation (Jansen van Vuuren & Robinson 2001).

The discovery of genetic affinity between the less specialized Redflanked Duiker and swamp-adapted Black-fronted Duiker offers special opportunities for the study of evolutionary history and adaptive radiation in duikers. One question concerns habitat.The Black-fronted Duiker has long legs and elongated, splayed hooves that parallel those of another swamp-dwelling antelope, the Sitatunga Tragelaphus spekii. Distribution patterns for both species suggest that their ancestral populations originally adapted to swamp forests bordering lakes and rivers running immediately north of the Equator (Kingdon 1982). As the Red-flanked Duiker is distributed along the furthermost northern tributaries of the same system both rufilatus and nigrifrons can be viewed schematically as ‘outliers’ that have been displaced onto the northern and southern margins of an ancestral range. In this construction their common ancestor was likely to have inhabited a long east–west forest belt running north of the Equator.This is a zone that currently supports a considerable diversity of duikers, all of which are ecologically partitioned by size, food and habitat type (Gautier-Hion et al. 1980, Kingdon 1982). Within this partitioned community, the Black-fronted Duiker has departed furthest from the generalized duiker type, whereas the Red-flanked Duiker has not. Instead, the latter’s distribution implies a sort of evolutionary ‘displacement’ out onto the northern margins of duiker habitats with minimal external change to show for it. None the less, both species are likely to possess inconspicuous but substantial physiological adaptations and among these is an observed difference in scent glands. The Redflanked Duiker is without glands in the groin, whereas the Black-fronted Duiker has developed large, deep inguinal glands. Understanding this divergence has the potential of throwing light on the overall functions of scent glands in antelope biology, a topic that is still deeply problematic. Antelopes belonging to several lineages have developed inguinal glands and the emission of their secretions is unequivocally directed at conspecifics where they augment other olfactory signals. Unlike facial, pedal and excretory scents, where animals betray a direct interest in the scent and may deliberately mark places or other animals, inguinal glands are generally cryptic in expression. Exactly what purposes these glands serve is uncertain, but they are known to signify excitement and presumably assist conspecifics to identify sex, age and condition and then track the individual. Although pedal glands are best suited to the last task, constant switching of the tail might help disperse inguinal scent along trails. It is resting places or ‘forms’ (possibly urine puddles too) that must be the main scent-depositories. When a previously glandless lineage develops such glands there is the distinct implication of a change in social structure or a heightened need for intra-specific contacts. The principal external cost of developing such very odoriferous scents is facilitating predators that track their prey by scent: a selective agency that is greatly reduced in densely obstructed, boggy habitats. In lacking inguinal glands, the Red-flanked Duiker resembles a majority of duiker species and it is likely that this is the original condition for these predominantly solitary animals. A careful and detailed comparison of Red-flanked Duiker social behaviour with that of Black-fronted Duikers might reveal currently unknown aspects of social behaviour in the latter. In any event, there is much still to be learned about communication and the evolution of social structures in duikers and antelopes in general. The preorbital glands are wiped on a wide variety of sites, vegetable and mineral, including termite mounds (Wilson 2001).The same site is often wiped with both sides of the face.

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Foraging and Food  Red-flanked Duikers feed on the fruit, flowers and foliage of numerous trees, shrubs and herbs. Some of their foods consist of large cumbersome fruits, others are little more than berries or seeds and every size of fruit in between. In Bangangai, Sudan, on the outermost borders of its range, Hillman (1982) noted fallen fruits, flowers, dry leaves, root tubers and the meat of dead animals. In the Toumodi area, Côte d’Ivoire, stomachs (n = 15) of duikers killed in and around forest-cleared farms contained the fruits of 21 different plants that were typical of the forest/savanna mosaic, with preference given to Phoenix reclinata, Nauclea latifolia and Ficus capensis, although other species consumed included Combretum racemosum, Griffonia simplicifolia, Alchornea cordifolia, Phyllanthus discoideus, Anthocleista djalonensis, Carapa procera, Antiaris africana, Ficus mucoso, Macuna pruriens, Canthium vulgare, Blighia sapida, Malacantha alnifolia and Solanum spp.They also contained cultivated plants such as mangoes, cassava, cocoa and pawpaw as well as raphia and oil palm dates (Hofmann & Roth 2003). In Mole N. P., these duikers have been recorded feeding on the figs of eight species of Ficus as well as fallen fruit of large trees, in particular Uvaria chamae, Xylopia parviflora, Elaeis quineensis, Nauclea latifolia, Pavetta crassipes and Anthocleleista vogelii (Wilson 2001). In Nigeria, this duiker was found to mainly browse the leaves of shrubs, such as Piliostigma thonningii, Pterocarpus erinaceous, Annoa senegalensis, Grewia arborea, Landolphia oweriensis, and Vitex domiana (Agbelusi 1992). Such flexibility in diet suggests an ability to survive fruitless periods or areas with few fruits by switching to browse and may help explain why this species thrives outside true forest. Red-flanked Duikers commonly rise onto their hindlegs to reach plant parts above their heads. They pay close attention to monkeys and exploit dropped waste beneath the primates’ food trees (Wilson 2001). Social and Reproductive Behaviour  Red-flanked Duikers are solitary, resident animals living in spaced-out territories at densities of about 3–4/km² (Wilson 2001). Male and female territories overlap, but each sex spends most of its time on its own. Of 386 observations in Mole N. P., 325 were of solitary animals, 47 were male–female pairs and ten were of // with their young. In the same area, Wilson (2001) found two animals that had 5–7 different resting places within an area of 2 ha. Middens were prominent landmarks in this habitat and Wilson (2001) found 90% of them deposited on open ground, well away from vegetation. Unlike some other antelope species, faeces were not piled up on top of each other and anything between 15 and 80 single deposits, of varying age, were found scattered over an area of 15–50 m2. The sexes have distinctively different urination postures, ?? with their hindlegs stretched out behind while // bunch up and lower their hindquarters to the ground. In captivity, ?? are almost continuously interested in // (even attempting to mount // not in oestrus), approaching the / from the rear with body stretched out and exhaling with a snort while following the / (Dubost & Feer 1988). The / utters a short moan as she exhales and ?? exhibit flehmen when sampling her vulva or

urine (Wilson 2001). A captive pair copulated four times in 40 min and oestrus lasted for a day and a half (Wilson 2001). Reproduction and Population Structure  There is no information on timing of parturition in the wild. Gestation period has been cited as 223–245 days (Schweers 1984). Based on records from captivity, birth-weights average 1.0 kg (range 0.8–1.2 kg) and weaning takes places at 93–98 days (Wilson 2001), although the latter author also notes a record of 58 days. A captive / gave birth at 26 months (Hayssen et al. 1993). Animals in captivity have survived more than 15 years (Weigl 2005). Predators, Parasites and Diseases  Martial Eagles Polemaetus bellicosus and Crowned Eagles Stephanoaetus coronatus, pythons and various carnivores, notably Lions Panthera leo, Leopards Panthera pardus, Servals Leptailurus serval and jackals Canis spp. are all likely predators. Dipeolu & Akinboade (1984) recorded the ixodid ticks Amblyomma variegatum and Boophilus decoloratus from animals in Nigeria, while Ntiamoa-Baidu et al. (2005) recorded the following ixodid tick species from Red-flanked Duikers in Ghana: Haemaphysalis parmata, Ixodes muniensis, I. moreli, Rhipicephalus simpsoni and R. ziemanni. Conservation  IUCN Category: Least Concern. CITES: Not listed. This species is declining due to hunting, but, as East (1999) remarked, has shown much resilience to both hunting and the spread of settlement and probably still occurs reasonably widely throughout much of its historical range. The species is common in many protected areas and reserves (about half of the total population occurs in and around protected areas), including Niokolo-Koba N. P. (Senegal), National Park of Upper Niger (Guinea), Comoé N. P., Haut Bandama Fauna and Flora Reserve and Marahoue N. P. (Côte d’Ivoire), Mole, Bui and Digya National Parks (Ghana), W N. P. (Benin), Bouba Ndjida, Bénoué and Faro National Parks (Cameroon), and Manovo–Gounda–St Floris and Bamingui–Bangoran National Parks (Central African Republic) (East 1999). Measurements Cephalophus rufilatus HB: 600–800 mm T: 70–100 mm Sh. ht: 300–380 mm WT: 6.0–14.0 kg Throughout geographic range (Kingdon 1997; mean and sample number not given) Maximum recorded horn length is 10.4 cm for a pair of horns from Kozo, Central African Republic (Rowland Ward) Key References  East 1999; Kingdon 1982; Wilson 2001. Jonathan Kingdon & Michael Hoffmann

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Cephalophus nigrifrons  Black-fronted Duiker Fr. Céphalophe à front noir; Ger. Schwarzstirnducker Cephalophus nigrifrons Gray, 1871. Proc. Zool. Soc. Lond. 1871: 598. ‘Gaboon’ (Gabon).

Black-fronted Duiker Cephalophus nigrifrons adult male.

Lateral and palatal views of skull of Black-fronted Duiker Cephalophus nigrifrons.

Taxonomy  Five subspecies have been recognized (see Ansell 1972), with four confined to montane areas of eastern Africa and the nominate race found in the forests of the Congo Basin.The Rwenzori Red Duiker Cephalophus rubidus has been included as a subspecies of C. nigrifrons (St Leger 1936, Groves & Grubb 1981), although Kingdon (1982) considered rubidus to be a full species (and see Grubb 1993c, Kingdon 1997). Jansen van Vuuren & Robinson (2001) presented molecular evidence supporting the recognition of the species as distinct, although the only material available for the analysis was a tooth from a specimen in the Swedish Museum of Natural History.Wilson (2001) was reluctant to consider rubidus a distinct species, and Grubb & Groves (2001) and Grubb (2005) both considered it a subspecies of C. nigrifrons. Grubb & Groves (2001) also described a new subspecies from the Itombwe Mts, west of the north end of L. Tanganyika, C. n. hypoxanthus. Cephalophus fosteri and C. hooki were considered evolutionarily distinct species by Cotterill (2003a). Synonyms: apanbanga, aureus, claudi, emini, fosteri, hooki, hypoxanthus, kivuensis, lusumbi, mixtus. Chromosome number: not known, but given their conserved karyotype it may be 2n = 60 (B. Janssen van Vuuren pers. comm.).

animal is moving and feeding. The very young have a darker pelage colour, which is lost within a few weeks of birth (J. A. Hart pers. comm.). The pedal glands are large and there are virtually no hairs blocking the opening to the glands (Wilson 2001). Well-developed inguinal glands occur in most individuals, but are sometimes missing from C. n. kivuensis. Sexes similar with the ? slightly smaller. The skull is narrower with the muzzle ‘pinched in’ so that the nasals obscure the maxillae in dorsal view (Grubb & Groves 2001). Both sexes have horns, which are black, short and straight, pointing backwards at about 45°  to the vertical.

Description  A medium-sized red duiker, similar to other red duikers, but with relatively long legs. Crown and forehead black with black or very dark red or brown frontal blaze (the blaze that gives this species its name), and a black tuft of hairs between the ears. A well-developed preorbital gland runs from the eye towards the muzzle. Pelage glossy red with coarse body and neck hair, some hairs with black or darker red tips, with a darker back and rump grading to a paler red belly. The pelage is shorter in lowland areas, but long and shaggy in high-lying areas; similarly, montane animals are darker and more grizzled. The legs are darker than the body and tend to almost black at the hocks. Some animals have a large amount of black on the shoulders. Hooves are long and thinner than in any other duiker (Wilson 2001). Tail is the same colour as the rump, red or dark red with a white underside that is constantly flicked as the

Geographic Variation C. n. nigrifrons: from the Niger Delta and Mt Cameroon, lowland forest of SE Cameroon, Gabon and Congo through the lowland forests of the Congo Basin and N Angola to the Albertine Rift Valley. In lowland forest, almost solely found in swamp forest along streams and rivers. Shining chestnut-brown, with forelegs black as far up as the elbow, and hindlegs usually up to hock; chest black. Forehead generally black; chin yellow (sometimes white) (Grubb & Groves 2001). Hooves are particularly elongated because swamp forest is the main habitat they occupy. C. n. kivuensis: mountains along the Albertine Rift from the Kahuzi Biega massif to the Rwenzori Mts. Very contrasting colouration, the limbs being grey-black and the facial blaze very black; hair thick and coarse; chin pale reddish, but more whitish in specimens from higher altitudes (Grubb & Groves 2001). C. n. fosteri: confined to Mt Elgon at 2600–3700 m. Brownish colour, with face and sides of neck reddish, not grey; chin white; coronal tuft short, entirely black; hair thick and coarse; smaller in size than other subspecies (Grubb & Groves 2001). C. n. hooki: confined to Mt Kenya at 2800–3300 m, and in the Aberdares. Chestnut or reddish-grey, with face and sides of neck greyer; chin reddish-white; face-blaze bordered by red stripe; coronal tuft short, entirely black; fur thick and coarse, and tail bushy;

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Black-fronted Duiker Cephalophus nigrifrons hooki.

smallest of the subspecies after fosteri.This subspecies is supposedly ecologically separated (by altitude and habitat) from Harvey’s Duiker C. harveyi on Mt Kenya (with Harvey’s Duikers not occurring above the bamboo line), although it is unknown whether they interbreed at all at the interface of their ranges (Grubb & Groves 2001). C. n. hypoxanthus: confined to the Itombwe massif in E DR Congo. Pale, light yellow-chestnut colour, with greyish legs rather than black; hair fairly long and very soft (more coarse in other highland races); chin white (Grubb & Groves 2001). The four montane forms generally are found between 2000 and 3800 m on their respective mountains, while C. n. nigrifrons occurs at much lower altitudes. The Rwenzori Red Duiker generally occurs above the range of C. n. kivuensis on the Rwenzori massif (up to 4200 m). Similar Species Cephalophus rubidus. Restricted to the upper reaches of the Rwenzori Mts. Dark grey underlying red tips of the hairs down the mid-line of the back and neck. Underfur of flanks cream; very pale to white belly. C. harveyi. Sympatric in the Aberdares and Mt Kenya. Black and white chin and whiter ears with black tips; belly pale cream or white; shorter legs. C. rufilatus. Allopatric. Smaller in size, this species has a grey-brown back and grey-brown legs that contrast with the red-orange on the face, neck and flanks. White on lower jaws contrasts with black nose. This species has no inguinal glands. C. ogilbyi. Sympatric in western central Africa (C. o. crusalbum) and presumably Nigeria/Cameroon (the nominate form). Paler pelage and in C. o. crusalbum with white legs; shorter legs. Distribution  Endemic to equatorial Africa. Widespread from SE Nigeria in the Niger Delta (where only recently confirmed; Powell & Grubb 2002), through the forests of S and SE Cameroon, south of the Sanaga R., mainland Equatorial Guinea, Gabon, SW and SE Central African Republic, Congo and N (Cabinda, Uige and Cuanza Norte Provinces) and NE Angola (Lunda Norte and Lunda Sul Provinces) eastwards across the Congo Basin to the Albertine Rift (East 1999, Wilson 2001, Crawford-Cabral & Veríssimo 2005). In Uganda they are known only from Mt Elgon and remnant forest on mountains along the Albertine Rift, including the Rwenzori Mts, Bwindi Impenetrable N. P. and the Virunga Mts, while in Rwanda and Burundi they are now restricted to Volcanoes N. P. and Nyungwe

Cephalophus nigrifrons

Forest Reserves, and the Nyungwe-Kibira forest on the Rwanda– Burundi border. In Kenya they occur only in three widely separated populations, on Mt Elgon, Mt Kenya and in the Aberdares (East 1999, Wilson 2001). Isolated populations of this species were recorded on Mounts Cameroon, Kupe and Manengube by Bowden (1986); however, Grubb et al. (2003) were not convinced these records relate to Black-fronted Duikers, and thought they may represent an undescribed species. Habitat  Found in lowland tropical forest in the Congo Basin from altitudes near sea level to more than 4000 m in montane forest and moorlands in East Africa.Typically occurs in swampy or waterlogged habitats along streams where its long hooves help it move around more easily. In the Ituri Forest, DR Congo, it was equally abundant in monodominant Gilbertiodendron forest as in mixed forest, although in both forest types it is confined to the courses of streams and rivers (Hart 1985, 2000). In the Virunga Mts there was a preference for wooded habitat with an open understorey dominated by dense herbs such as nettles up the sides, and in the saddle between the volcanoes (Plumptre & Harris 1995, Plumptre & Bizumuremyi 1996). On Mt Elgon it occurs in dense bamboo forest where it is probably not abundant (J. Kingdon pers. comm.), but in the Virungas this was not a preferred habitat and densities were low in bamboo (Plumptre 1991). On mountains there is less dependence on waterlogged areas than there is in the lowland forest although the long hooves are still present, implying that the original form may have occurred in lowland forest where its adaptations to waterlogged soils might have favoured its colonization of moist forested areas and spread up the mountains. There is some evidence that this species avoids predation by Leopards Panthera pardus by being better adapted to swamp forest in lowland areas where the supply of fruit is lower and animals survive on herbs (Hart 2000, 2001). Abundance  In the Virunga Mts, densities ranged from 5 to 22/km2 across a range of different habitats (Plumptre & Harris 1995). In the 269

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Black-fronted Duiker Cephalophus nigrifrons kivuensis adult male facial features and forehead.

above and right:

Ituri Forest, density was lower, at about 1.3–2.0/km2 (Hart, J. A. et al. 1996, Hart 2000, 2001). Assuming average densities of 2/km2 in high-density areas and 0.2/km2 in low-density areas, East (1999) estimated the total population at around 300,000. Adaptations  The long hooves enable this species to inhabit swampy and waterlogged areas. J. A. Hart et al. (1996) have suggested that this gives it an advantage in keeping away from Leopards, one of its major predators. The foot-stamping that is typical of other duiker species appears to be absent in this species. It is possible that speciesspecific vocal signals may compensate. When feeding or moving, animals constantly flick their tails, exposing the white undersides that may be used to advertise their presence to others. The bone of the skull on the foreheads is thickened, probably to minimize the impact of head butting between individuals. Black-fronted Duikers are primarily diurnal, but occasionally are active at night. Foraging and Food  Diets vary where they have been studied. In the Ituri Forest, 29% of the diet determined from rumen contents was leaf material (primarily herbs) and the remainder was fruit and seeds, with some flowers and fungi (Hart 1985). Those fruits consumed tend to be fairly fibrous, such as Ricinodendron heudelotti, rather than fleshy. However, it was the most folivorous of all the duikers in the Ituri Forest and the only one to eat herbs. The diet varied seasonally depending on the availability of fallen fruit, and seed content increased dramatically in most years for Gilbertiodendron dewevrei. In Makokou, Gabon, rumen analysis indicated 73% of the diet consisted of fruit (mostly 1–3 cm in diameter) with the remainder consisting of leaves, flowers, petioles and fungi (Gautier-Hion et al. 1980); the Black-fronted Duiker was again the most folivorous duiker of those studied in Makokou (Dubost 1984). At higher altitudes (2900–3700 m) in the Virunga Mts there was very little fruit available and 80.7% of the diet consisted of leaves of herbs and grasses, with 14.8% lichen and 4.5% bark/roots based on faecal pellet analyses (Plumptre 1995). The low tannin and alkaloid levels in montane herbs probably allow this species to survive on such a drastically different diet and to live at high densities (Plumptre 1995). Usnea lichen was highly preferred, with individuals competing over pieces that fell to the

ground and standing on their hindlegs to reach lichen that was within reach. The desire for this lichen may be due to its reasonably high level of soluble carbohydrates (A. Plumptre unpubl.), which may substitute for those found in fruit consumed at lower altitudes. Diet in the Virungas did not vary seasonally as there was little variation in the availability of food. Social and Reproductive Behaviour  Generally solitary with occasional sightings of pairs (either mother–infant or male–female). They are territorial, using preorbital glands to mark territories, and tending to defecate near the boundaries of their territory. Territorial defence may be aided by the use of a thumping sound that appears to be vocal (Kingdon 1982), and a loud whistle-like call that is also used as an alarm call. Very occasionally two individuals will fight, which involves rearing up and butting heads together, pausing to recover and then repeating this, reminiscent of the way goats fight. Individuals have also been seen to snap at each other (Walther 1984). Fighting usually ends with a pursuit through the vegetation. Anti-predator behaviour involves freezing, often in mid-stride, sinking down to the ground or sneaking away to hide in dense vegetation if they believe they have not been detected. If they know they have been detected they flee, giving a loud whistle, dashing through the vegetation with their head down for a short distance, often in a zig-zag manner to throw off any predator. Like many ungulates, young when born are left concealed by the mother in dense vegetation and she returns to suckle at regular intervals. Reproduction and Population Structure  Nothing is known about reproduction of this species either in captivity or in the wild. Wilson (2001) mentions two lactating // recorded in Feb and Mar

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from the Central African Republic, and two pregnant // in Dec. Longevity in captivity has been given as nearly 20 years (Jones 1982). Predators, Parasites and Diseases  Leopards are a major predator of all duiker species in the Ituri Forest and have a significant impact on their populations (Hart, J. A. et al. 1996). Leopards account for 87% of duiker mortality in areas where humans do not hunt in the Ituri Forest (Hart 2000, J. A. Hart pers. comm.). However, this species only formed about 1.7% of Leopard diet in the Ituri Forest, the lowest of any duiker species, despite being reasonably abundant, and occurred in only half of a calculated expected number of Leopard scats (Hart, J. A. et al. 1996). They also had the lowest mortality rate of any duiker species in the Ituri Forest (Hart 2000). Other large cats, such as African Golden Cats Profelis aurata, and African Rock Pythons Python sebae will also take this species. Large eagles such as the Crowned Eagle Stephanoaetus coronatus will take juveniles and very occasionally adults. Robust Chimpanzees Pan troglodytes and Nile Crocodiles Crocodylus niloticus occasionally take individuals. Helminths recorded from Black-fronted Duikers include cestodes (genera Avitinella, Stilesia) and nematodes (Bunostromum, Dipetalonema, Ochocera and Setaria) (Round 1968, Bain et al. 1978, Chabaud et al. 1978). A study of the diseases and parasites found in wild duikers in the Ituri Forest showed that most Black-fronted Duikers in this forest contain gut parasites (Karesh et al. 1995). Of the five individuals sampled all had at least evidence of one parasite species in their faeces and for most parasites the percentage of individuals infected was higher for this species than the four other duiker species tested. Evidence of parasites in the blood included: ‘bluetongue’ virus, which was at a similar level to other species; epizootic haemorrhagic disease and infectious bovine rhinotracheitis, which were generally at a lower level than other species; and leptospirosis, which was at a higher level of infection compared with most of the other species sampled (Karesh et al. 1995). Some of these infections, particularly leptospirosis, could be transmitted to humans through the bushmeat trade in this species.

can accelerate their maturation in // under low population densities but not as well as blue duikers can (Hart 2000). They are easily trapped, as there is some indication that they follow regular paths from the swamps to drier parts of the forest (Rahm & Christiaensen 1963, Wilson 2001). Data from Rwanda revealed that Black-fronted Duiker meat is significantly lower in price than domestic meat (Plumptre & Bizumuremyi 1996, Plumptre et al. 1997) and they were a favoured species for hunters. There was evidence that poaching of this species had increased in Volcanoes N. P. during and after the insecurity created by the civil war and genocide in Rwanda (Plumptre et al. 1997). Protected areas important for the continued persistence of the species include Apoi Creek F. R. (Nigeria), Dja Faunal Reserve (Cameroon), Monte Allen N. P. (Equatorial Guinea), Lopé N. P. (Gabon), Dzanga-Sangha Forest Reserve and Bangassou Forest (Central African Republic), Nouabalé-Ndoki and Odzala National Parks (Congo), Kahuzi-Biega, Maiko and Virunga National Parks and Okapi Wildlife Reserve (DR Congo), Bwindi Impenetrable and Rwenzori Mountains National Parks (Uganda) and Volcanoes N. P. (Rwanda). Measurements Cephalophus nigrifrons TL (??): 1080 (1020–1120) mm, n = 6 TL (//): 1090 (1040–1140) mm, n = 7 T (??): 155 (130–160) mm, n = 6 T (//): 150 (140–160) mm, n = 7 HF c.u. (??): 295 (281–304) mm, n = 6 HF c.u. (//): 296 (281–310) mm, n = 7 E (??): 90 (87–94) mm, n = 6 E (//): 91 (86–95) mm, n = 7 Sh. ht (??): 560 (540–570) mm, n = 6 Sh. ht (//): 560 (540–580) mm, n = 7 WT (??): 13.8 (12.7–14.9) kg, n = 6 WT (//): 14.0 (13.1–16.3) kg, n = 7 Cameroon, Central African Republic and Congo Republic (Wilson 2001) Hart, J. A. et al. (1996) recorded the mean weight of two // as 15.5 kg (range 15–16 kg) and 14.3 kg for two ?? (14.0–14.5 kg) Maximum recorded horn length is 12.0 cm for a pair of horns from Ntem R., Cameroon (Rowland Ward)

Conservation  IUCN Category: Least Concern. CITES: Not listed. Numbers of Black-fronted Duiker are probably declining across the range, mainly due to hunting, particularly with the opening up of the Congo Basin by timber extraction companies. The three subspecies confined to Mt Elgon, Itombwe Massif and Mt Kenya are probably at greater risk of extinction. This is particularly the case for C. n. Key References  Hart 1985, 2000, 2001; Kingdon 1982; Plumptre hypoxanthus on the Itombwe Massif, which is not currently protected. 1991, 1995; Plumptre & Harris 1995; Wilson 2001. In central and West Africa this species is hunted heavily wherever it occurs in proximity to man. There is some evidence that red duikers Andrew J. Plumptre

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Family Bovidae

Cephalophus ogilbyi Ogilby’s Duiker Fr. Céphalophe d’Ogilby; Ger. Ogilbyducker Cephalophus ogilbyi (Waterhouse, 1838) Proc. Zool. Soc. Lond. 1838: 60. Equatorial Guinea, ‘Fernando Po’ (= Bioko).

Taxonomy  Ogilby’s Duiker, as described here, probably repre­ sents a species group that embraces three allopatric species. While acknowledging this probability we follow Wilson (2001) in provisionally continuing to treat this complex as a single species. The type of C. ogilbyi was described in 1838 from the island of Fernando Poo (Bioko); C. o. crusalbum was described in 1978 from Gabon and C. o. brookei in 1903 from Ghana. Brooke’s Duiker C. o. brookei has been recognized as a full species by several recent authors (Grubb et al. 1998, Grubb & Groves 2001, Grubb 2005). Cephalophus o. crusalbum has been recognized as a full evolutionary species by Cotterill (2003a). Molecular studies suggest that Ogilby’s Duiker is an ancient relict duiker that has ambiguous genetic links with other duiker species, notably with Peters’s Duiker C. callipygus (Jansen van Vuuren & Robinson 2001; and see Hassanin et al. 2012). Payne (1992) considered Ogilby’s Duiker and Peters’s Duiker to be particularly close, and Grubb (1978a) thought the population he named C. o. crusalbum was ‘intermediate’ between Peters’ and Ogilby’s Duikers. Wilson (2001) drew attention to the ‘intergrading’ of some Ogilbylike characteristics in individual Peters’s Duikers, at or close to their zones of contact. This raises the possibility of occasional, or even sustained, long-term hybridization, past and present, between individuals or even entire populations. As ancestral duiker populations have been replaced by later ones the nature of that replacement could include various permutations of genetic admixture. As Ogilby’s Duiker gives every sign of being a relictual form in the process of decline, the possibility of genetic admixture should be borne in mind. Unscrambling such mixed genetic backgrounds will be an interesting task for future molecular scientists. It is, therefore, appropriate to point out that the C. ogilbyi complex represents an enigmatic and puzzling taxonomic entity. Our uncertainties about the status of Ogilby’s Duiker populations challenge science, yet their rarity, their relictual nature and their vulnerability to hunting may soon deprive scientists of important clues to the nature of duiker radiation. Synonyms: brookei, crusalbum. Chromosome number: not known. Description  A trim orange to mahogany-coloured duiker with chunky hindquarters and a bold black dorsal stripe (10–60 mm wide) that tapers to a point just above the tail. Nose and crown are black, but red or orange ‘brows’ of variable width may meet on the forehead. Horns may be partially obscured by short tufts of red or black fur. Pinnae are sepia behind and have narrow white margins with tracts of short paler hair in their forward aspect. On Bioko I., necks are well furred with soft but longish fur, while mainland populations tend to have short-haired darkish necks; hair on the neck is frequently reversed. Body colour extends down relatively long, slender legs, but C. o. crusalbum has off-white legs while C. o. brookei has narrow black lines or smears down the front of the limbs. Small lateral hooves are present. Inguinal, preorbital and pedal glands are all present. Tail narrow but ends in a distinct tuft that is sometimes quite large, like a pompom, with hairs up to 75 mm long (as in C. o. crusalbum).

Ogilby’s Duiker Cephalophus ogilbyi ogilbyi.

The horns are short but peculiarly incurved and heavily corrugated and can occur on both sexes. The skull is characterized by the extreme inflation of the forehead behind the nasofrontal suture, such that the skull has a marked frontal boss, but which is reduced laterally so that the infraorbital foramina are not roofed over; the zygomatic arch is more curved than in the Black Duiker, flaring out in the middle (Grubb & Groves 2001). Geographic Variation C. o. ogilbyi (Ogilby’s Duiker): Bioko I., Equatorial Guinea, and Camer­ oon/Nigeria borderlands. Larger body size. Relatively uniform golden-rufous colouring overall but with whitish underside, throat and chin. Crisp, narrow, black mid-dorsal line. Neck pelage not noticably shorter than rest of body. Heavily bossed forehead.

Brooke’s Duiker Cephalophus ogilbyi brookei.

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White-legged Duiker Cephalophus ogilbyi crusalbum.

C. o. brookei (Brooke’s Duiker): Sierra Leone, possibly SE Guinea, Liberia, S Côte d’Ivoire and Ghana, west of the Volta R.; Kingdon (1997) mistakenly described this population occurring in Cameroon. Pale golden colour with slightly redder rump; broad dorsal stripe narrows to a point above tail; red coronal tuft; neck hair very short; forehead not obviously bossed. Probably a distinct species. C. o. crusalbum (White-legged Duiker): Gabon, mostly south of Ogooué R., and extreme NW Congo. Smaller body size; golden or orange-ochre torso, redder rump, but lighter on the flanks; somewhat greyer neck and face; neck hair very short; broad dorsal stripe; legs white from knee and hock to hoof, in contrast to all other duikers; tail orange-ochre with median stripe black and pronounced tail tuft; forehead not conspicuously bossed. Similar Species Cephalophus nigrifrons. Sympatric in Gabon and Congo (with C. o. crusalbum) and Nigeria/Cameroon (C. o. ogilbyi). A long-legged, long-hooved, dark red duiker with no dorsal band and a strong preference for swamp forest. C. callipygus. Sympatric in Gabon (e.g. in Forêt des Abeilles and Lopé N. P.) and Congo. A large species with dark legs and extensive black dorsal stripe on upper rump. Hybridization may occur and C. o. crusalbum sometimes appears to be intermediate between C. ogilbyi and C. callipygus (see Grubb 1978a). C. dorsalis. Sympatric throughout mainland range of Ogilby’s Duiker. A broad-headed, short-faced, slender-horned, heavily built and usually dark red duiker, which is readily confused with Ogilby’s Duiker. Nocturnal. C. niger. A black duiker, restricted to Upper Guinea forests from SW Guinea to the Niger R. Distribution  Endemic to Africa, Ogilby’s Duiker is distributed in four distinct populations. In West Africa it occurs in the forests of Upper Guinea, from Sierra Leone (where the only confirmed records are from the Outamba part of Outamba–Kilimi N. P. and Lalehun) through Liberia and S Côte d’Ivoire to Ghana, west of the Volta R. (Grubb et al. 1998, East 1999,Wilson 2001). May occur in SE Guinea, where reported from Ziama–Diécké F. R. by East (1999), but not by Butzler (1994). There is no recent information on the presence of this species in SW Ghana, and East (1999) noted that it had disappeared

Cephalophus ogilbyi

from the Bomfobiri and Owabi wildlife sanctuaries where it occurred formerly. There is a break in distribution until reappearance (in the form C. o. ogilbyi) in the moist lowland forest in SE Nigeria and SW Cameroon. An isolated population occurs on Bioko I., particularly at higher altitudes. The white-legged form of Ogilby’s Duiker, C. o. crusalbum, occurs in Gabon, mostly south of the Ogooué R., and is reported to occur in NW Congo (East 1999, Grubb et al. 2003).There is no record of this species from mainland Equatorial Guinea. Habitat  Found in all forest types, from sea level up to 2260 m and perhaps even higher on Bioko I. (Butynski et al. 2001), where it is the only large duiker species, and where it occurs in the Schefflera Forest Zone (ordinarily occupied by other species on the mainland). No clear specialization in forest type. Occurs in dense forest patches on the Guinea savanna edge in Sierra Leone (Grubb et al. 1998). Reported from a mixture of undisturbed high and also logged forests in Liberia (Wilson 2001). Found in forest patches within forest–savanna mosaic in northern part of Lopé N. P. in Gabon (East 1999). A different permutation of other duiker species is found in every one of the areas where Ogilby’s Duiker occurs but there is virtually no overlap in range with ‘main line’ broad-spectrum red duikers (exclusive of Bay Duikers) (Kingdon 1982). The absence of any clear specialization in habitat or diet suggests that duikers of the C. ogilbyi complex represent relictual populations of an early lineage that may be in the process of succumbing to competition and genetic contamination from other forest duikers.The most notable of these is Peters’s Duiker, which is almost allopatric and a very similar species. This similarity is most marked along the borderlands between the two species where occasional ‘intermediate’ specimens suggest that the ‘relationship’ between the two species may be complicated by long-term hybridization. In any event, the situation calls for further investigation. In Upper Guinea, where Black Duikers and Bay Duikers are the commonest large duikers, C. o. brookei was formerly widespread but never common (Grubb et al. 1998). 273

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Lateral view of skull of Ogilby’s Duiker Cephalophus ogilbyi ogilbyi.

Abundance  Data are scarce, but during a survey conducted in 1990 in the Gran Caldera on Bioko I., Butynski et al. (2001) found evidence for their presence at the rate of 0.24 per linear transect km in suitable habitat. Figures from bushmeat markets and hunter interviews suggest an annual off-take of 4000 Ogilby’s Duikers on Bioko I.; about 2/km² of forest habitat. A more localized estimate was 2250 from 160 km2 (Fa et al. 1995). While it is not known how sustainable such off-takes are, East (1999) assumed an average density of 10/km2 on Bioko I., 2/km2 in other areas where the species is known to be common, and 0.2/km² elsewhere, and estimated a population of 12,000 animals on Bioko I. and in the Nigeria/ Cameroon area. Elsewhere, he estimated 5000 individuals for C. o. brookei and 18,000 for C. o. crusalbum. It is worth noting that Wilson (2001), who conducted extensive survey work over a period of ten years in various parts of West Africa, only saw C. o. brookei in the wild once, in Kakum N. P., Ghana. Adaptations  A very generalized, but variable type of duiker; the possibility of a relictual status has been discussed above. There is also the possibility, raised by several authors (Grubb 1978a, Kingdon 1982, Wilson 2001), that hybridization may complicate our understanding of the duiker radiation, and of this species in particular. This possibility is central to any attempt at defining the adaptive niche of Ogilby’s Duiker. There are several reasons to suspect that this duiker is a relict species in active decline and in the process of replacement by other species of the same generalized red duiker lineage. First is the lack of any distinct specializations. Second are peculiarities of its disjunct distribution, which lack any clear correspondence with vegetation or climatic zones. Third is a peculiar interdigitation in its distribution with that of the commoner Peters’s Duiker, which implies competition, decline and retreat from the latter. Fourth, Ogilby’s Duiker, from the margins of this ‘frontier’ zone, shows several characteristics that imply previous long-term hybridization. Likewise, some Peters’s Duikers show variations in the form of their dorsal stripes that could be taken to imply hybridization between the two species. Fifth, current molecular evidence allies this species with several other species, including Peters’s Duiker, and such ambiguities of affinity could imply both/either archaic status and/or a history of hybridization. The sixth reason to suspect an overall decline in the face of competition from other duiker species is the fact that the Bioko form is very common on the island, yet naturally rare in many of its remaining mainland sites. Unlike the long-haired island population, mainland duikers have short-haired necks that could imply genetic admixture because neck pelage is generally a clear distinguishing characteristic between two separate duiker lineages (Kingdon 1982). These and many other unanswered questions about the C. ogilbyi complex preclude any easy definition of the adaptive niche of this species.

The apparent restriction of bossed foreheads to Bioko duikers, combined with known recent high densities, suggest that this isolated species/subspecies lives in smallish, frequently defended territories. It is interesting to speculate whether frontal bossing once occurred in the ancestral populations of C. o. brookei and C. o. crusalbum. The anomalous occurrence of this physiologically expensive characteristic in just one population implies that reduced densities and less frequent conflict among mainland populations might have caused rapid atrophy in what might otherwise have been taken as a relatively immutable diagnostic characteristic (as tends to be the case in descriptions of the skulls of C. weynsi, C. callipygus and C. niger, all species that live at high densities). Foraging and Food  The ability of this duiker to range through all forest zones on Bioko I. implies a catholic diet and ability to adapt to different vegetation and rainfall zones. Like other duikers, it is known to feed on fruit, flowers and, presumably, foliage from the forest floor. Gautier-Hion & Gautier (1994) observed a subadult C. o. crusalbum eating the hard fruits of Klainedoxa gabonensis. The stomach contents of a specimen of C. o. brookei examined by Newing (2001) comprised 92% fruits and seeds, 7% vegetative parts and 1% flowers. Social and Reproductive Behaviour  A single adult ? that was radio-collared in Korup N. P., Cameroon, provided valuable information on the movements and activity (as determined from a mercury activity switch) (Payne 1992). The lucky capture of this individual followed a large number of fruitless drives and capture efforts in an area where Ogilby’s Duikers were already acknowledged to be rare. The home-range of this ? was estimated to be 10.6 ha (as determined by the minimum convex polygon method). Within this home-range there was a small central area that was consistently and almost exclusively used for sleeping by this strictly diurnal duiker. By contrast, diurnal resting areas, mainly used between mid-day and 15:00h, were on the peripheries of the home-range. Peak activity was immediately after dawn (06:30–11:00h) and between 16:00h and 19:00h. Of 50 sightings, only three, thought to be mother–offpring pairs, were not solitary; this is in agreement with the findings of Gautier-Hion & Gautier (1994), all of whose observations were of solitary individuals or pairs. In contrast to mainland species, which have been consistently described as rare and shy, Bioko duikers, when surveyed between Jan and Mar 1986, were abundant and vocal, implying a more complex social structure. They were frequently heard to make a loud ‘wheet’ call (Butynski et al. 2001) and East’s (1999) estimate of 10/km2 was probably realistic for the time of Butynski’s surveys (and partly also due to the general absence of predators on duikers on Bioko; see Predators, Parasites and Diseases). Reproduction and Population Structure  Currently, there is no published information, but the dynamics of population structure are likely to differ radically between high-density C. o. ogilbyi on Bioko and very low density populations on the mainland. There is no indication of a breeding season. Predators, Parasites and Diseases  Leopards Panthera pardus may prey on Ogilby’s Duikers in some mainland habitats, and African Golden Cats Profelis aurata are also likely to take young and possibly

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adults, as are African Rock Pythons Python sebae. Young animals are probably taken by Crowned Eagles Stephanoaetus coronatus. Mandrills Mandrillus sphinx probably prey on young and infirm duikers (Lahm 1986, T. Butynski pers. comm.). Butynski et al. (2001) suggested that the main predators likely include Drill Mandrillus leucophaeus on Bioko, where hunters imitate the cry of an Ogilby’s Duiker young to bring in Drill close enough to shoot. Conservation  IUCN Category: Least Concern (C. o. crusalbum – Least Concern; C. o. brookei – Vulnerable C1; C. o. ogilbyi – Vulnerable C1). CITES: Appendix II. The main threats to Ogilby’s Duikers are habitat degradation and overhunting, and on current trends these duikers will become extinct or will survive in a few well-protected areas. The effective protection of the Gran Caldera de Luba Scientific Reserve is crucial to the survival of this species on Bioko I. On the mainland, Upper Guinea populations of C. o. brookei have few remaining strongholds. Among them might be Sapo N. P. (Liberia) and Taï N. P. (Côte d’Ivoire), although hunters resident in Tai N. P. have been reported harvesting 1500–3000 tonnes of wild animal meat per year in parts of this park (Caspary et al. 1999) and heavy exploitation of bushmeat has been observed around Sapo N. P. (R. Hoyt pers. comm.).This form has also been recorded recently in Gola N. P. in Sierra Leone (Lindsell et al. 2011). Cephalophus o. ogilbyi occurs in Korup N. P. (Cameroon), where it is very rare (Payne 1992), and in Cross River N. P. (Nigeria); they are also present in several forest reserves in SW Cameroon, such as Mone and Ejagham (Forboseh et al. 2007).The distinctive white-legged form is now known to be relatively widespread and numerous in Gabon, including protected areas such as Lopé N. P. and the Gamba complex of National Parks, as well as Odzala N. P. (Congo). This species is hunted throughout its range, but with particular intensity in Bioko, where the bushmeat trade has been well studied over a period of years (Colell et al. 1994, Fa et al. 1995, 2000, Juste et al. 1995, Butynski et al. 2001). Of 94,616 wild animal carcasses brought to the bushmeat market in Malabo between the last quarter of 1997 and the first 10 months of 2006, 5.3% were Ogilby’s Duikers and this species ranged between the fourth and seventh most common in the market over this period (W. Morra & G. Hearn

pers. comm.). This is similar to the 6% recorded between October 1990 and October 1991, when Ogilby’s was the fifth most common species in the market at that time (Juste et al. 1995; and see Fa et al. 2000). These 1991 market harvest levels were clearly unsustainable (Fa et al. 1995), yet availability increased to 89% in 1996; in 1996, estimated daily abundance was about 3.4 carcasses per day (Fa et al. 2000). Since then, however, the availability of this species at the Malabo market has undergone a steady decline reaching 1.9 carcasses per day in 2006 (W. Morra & G. Hearn pers. comm.). Colell et al. (1994) found that of animals taken by hunters in south-east Bioko, Ogilby’s Duiker was the fourth most frequently collected species; 71% of these were shot and 29% were trapped. Less than 10% were consumed locally, the rest being taken to market, where prices as high as US$14 per carcass were paid in 1986 (Butynski et al. 2001). As of 2006, the average price paid had increased to US$94 (W. Morra & G. Hearn pers. comm.). The top priority for the conservation of Ogilby’s Duiker on Bioko is to stop openly illegal hunting of this species in the two nominally ‘protected’ areas (T. Butynski pers. comm.). Measurements Cephalophus ogilbyi HB: 850–1150 mm T: 120–150 mm E: 88 mm* HF c.u.: 260 mm Sh. ht: 550–650 mm WT: 14.0–20.0 kg Kingdon (1997); *single adult (sex not noted) from Bioko I. (T. Butytnski pers. comm.) Payne (1992) gave the weight of three adult ?? from Korup N. P., Cameroon, as 18.0, 19.0 and 20.0 kg Maximum recorded horn length is 12.3 cm for a pair of horns from Grebo, Liberia (Rowland Ward) Key References  Payne 1992; Wilson 2001. Jonathan Kingdon

Cephalophus weynsi Weyns’s Duiker Fr. Céphalophe de Weyns; Ger. Weynsducker Cephalophus weynsi Thomas, 1901. Ann. Mus. Congo Zool. 2 (1): 15. ‘district des Stanley-Falls’ (DR Congo, near Stanley Falls).

The duiker is named for Lt. Col. Auguste F. G. Weyns, a Belgian explorer for whom Weyns’s Weaver Ploceus weynsi is also named.

barbertoni, centralis, ignifer, johnstoni, leopoldi, lestradei, rutshuricus. Chromosome number: unknown, but likely 2n = 60.

Taxonomy  Previously treated as a subspecies of Peters’s Duiker Cephalophus callipygus (Kingdon 1982, East 1999, Wilson 2001), or even lumped variably in the past with the Natal Red Duiker C. natalensis (Ansell 1972) and Harvey’s Duiker C. harveyi (Haltenorth & Diller 1980). Considered a distinct species by East et al. (1990), Groves & Grubb (1974), Grubb & Groves (2001) and Grubb (2005). Specific designation is supported by genetic analysis (Jansen van Vuuren & Robinson 2001; but see Hassanin et al. 2012). Three subspecies were recognized by Grubb & Groves (2001). Synonyms:

Description  A medium-sized, coarse-haired, rufous-coloured duiker, distinguished by its lack of a black dorsal back stripe.The species is variably darker, especially on the limbs, shoulders and underparts, especially in some eastern populations. Crown and forehead red to reddish-brown with a coronal tuft of thick, bright, rufous hairs located between the horns, and covering the horn base in both sexes.The crown is rounded and uplifted in a distinct vaulted boss. Sides of face and ears variably brownish-rufous. The lips are thickened, giving the animal a coarse appearance. Back, sides and abdomen bright rufous with hairs 275

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Weyns’s Duiker Cephalophus weynsi adult female. Lateral view of skull of Weyns’s Duiker Cephalophus weynsi adult male. left:

above:

longest on the rump. Neck and shoulder rufous, mixed with darker hairs, especially on the shoulder. Neck and shoulder hairs shortened and hair on the neck reduced, markedly in some populations, exposing the pale-coloured skin beneath. Pelage on limbs uniformly dark below the hocks, extending variably onto the upper limbs, chest, upper abdomen and shoulders in some populations and individuals. Lower abdomen covered with sparse pale hairs. Tail rufous at base, dark distally, with small dark tuft containing scattered white hairs. Preorbital glands are large, running from the eye towards the muzzle. Inguinal glands are well developed, producing a reddish secretion, which, in populations in the Ituri Forest in NE DR Congo, is sweet-smelling. Pedal glands are present on all four feet (Wilson 2001). Sexes are similar in colouration, and while ?? are smaller than //, as is true in all forest duikers, sexual dimorphism is reduced; adult // average less than 3% larger body weight than adult ??. Pelage of newborns is overall brown, darker above and liberally grizzled with red hairs. The coronal tuft is present, even when horns occur only as buds. This pelage is replaced by adult colouration before the animal is weaned. The skull has an elongated rostrum and is distinguished by the vaulted forehead and robust horn bases. Both sexes have horns, which are black, short, thickened and strongly annulated at the base and over two-thirds of their length. Horns, distinctly longer in ? than /, are often curved downward and inward at the tips. Geographic Variation  A variable species, with the taxonomic status of a number of forms, in particular east of the Albertine Rift and south of the Congo R., still uncertain (Wilson 2001). Cephalophus w. weynsi (including leopoldi and centralis), the most rufous form, ranges across DR Congo, on the right bank of the Congo R. east into the sub-montane forests of Kivu, south through Maniema and south Kivu to the forest limit. The forms in the montane forests of the Itombwe Massif, west of L. Tanganyika appear to be this race. Weyns’s Duikers from the forest patches of SE Sudan and Uganda (west of L. Victoria) may also be attributable to this form, though some populations of the latter have been lumped as C. w. johnstoni. C.

w. lestradei, characterized by overall darker pelage, is reported from the sub-montane and montane forests along the eastern Albertine Rift, including Rwanda (Groves & Grubb 1974), and in the forests east of L. Tanganyika; however, the occurrence of this form, as well as the probably now extirpated C. w. rutshuricus, named from a now deforested area of eastern Kivu, and their relations to the nominate race on the west side of the Albertine Rift, are not well defined. The taxonomic status of a number of eastern forms, characterized by their smaller size and lumped as C. w. ignifer (Grubb & Groves 2001, Grubb 2005), and including duikers from Mt Elgon (C. w. barbertoni) through the forest patches of W Kenya, is also poorly known. Possible hybrids between C. weynsi and C. harveyi are reported from the Mau escarpment (Kingdon 1982). Duikers south of the Congo R., ranging through the central cuvette to Kasai, and Sankuru have been attributed to this species; however, these forms have a black dorsal patch, and may be better grouped with C. callipygus. Similar Species Cephalophus callipygus. Allopatric, occurring only west of the Congo and Ubangui Rivers. All forms have a distinctive dark patch on the back and rump, which distinguishes them from Weyns’s Duiker. C. leucogaster. Sympatric east of the Ubangui R. Distinguished by paler colouring, and dark dorsal patch. C. dorsalis. Sympatric in the east of its range, east of the Ubangui R. Broader-headed, shorter-muzzled, darker rufous, and with a distinct dorsal black patch. C. harveyi. The species occurs mainly east of the Rift Valley in Kenya, but may hybridize with Weyns’s Duiker along the Mau escarpment (Kingdon 1982, Wilson 2001). Black and white chin and whiter ears with black tufts; belly is pale cream or white; shorter legs. Distribution  Endemic to equatorial Africa. Widespread in DR Congo, including large areas of contiguous range north of the Congo R., from the Ubangui R. in the west, through the Ituri Forest to the limits of continuous forest on the western flanks of the Albertine

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Rift, then south to the forest limits in Maniema, Kivu and possibly into northern Katanga province. The northern and southern range limits appear to be defined by the forest/savanna boundary on both sides of the Equator. The species’ eastern distribution is more fragmented, with populations recorded historically from a number of lowland, sub-montane and montane forest islands in W Uganda,W Rwanda and Burundi, W Tanzania (Mahali Mts and Gombe), S Sudan (Imatong and Dongotona Mts), east to Mt Elgon and the forests of Kakamega and the Mau Escarpment in W Kenya. This species has not been recorded east of the Rift Valley in Kenya. Some eastern populations may now be seriously reduced or extinct. Habitat  In its lowland range in DR Congo the species is restricted to large blocks of closed forest and does not range far out into gallery forests or forest islands in the savanna ecotone. However, in East Africa, Weyns’s Duiker occurs in relatively small and isolated forest remnants. Its limited occurrence at the forest edge in the north of its range may be due to the presence of Red-flanked Duiker C. rufilatus, which has specialized on the forest/savanna ecotone. In the Ituri Forest, Weyns’s Duiker occurs widely in primary and older secondary forests, avoiding recent clearings and deep swamps. The species is common in mixed canopy forests, but is rare or absent from large stands of monodominant Gilbertiodendron dewevrei (Hart 2001). The species ranges from altitudes of 400 m to about 2000 m in forests of the Albertine Rift highlands, while east of the Albertine Rift, it occurs locally in sub-montane to montane forests up to 3000 m (reportedly on Mt Elgon; Haltenorth & Diller 1980). Abundance  In the Ituri Forest, Weyns’s Duiker is the second most common duiker species after Blue Duiker Philantomba monticola. Densities of unhunted populations averaged 15 animals/km2 (Hart 2000). East (1999) estimated a total population 188,000. Adaptations  Weyns’s Duiker is a robustly built duiker with a strongly developed head and neck. Relatively large male body size, in association with the distinctive bony vaulted crown and reinforced horn bases, present on both sexes, but most developed in the ?, are indicative of regular intra-specific combat. In the Ituri Forest, preorbital glands of Weyns’s Duiker ?? are often strongly swollen with visible exudates, suggesting frequent marking behaviour. Males, and possibly //, also mark by rubbing the coronal area against the stems of small trees. Animals that are excited or stressed show a red flush on the pale skin of the neck (which is exposed because the hair is so sparse). Weyns’s Duiker has an elongated rostrum, and the tooth arch is not markedly widened, but the dentition is robust, and suggests the capacity to handle coarse foods. Relatively high rumen volume to body weight ratios also confirm the species’ ability to digest a broad range of food quality (Hart 1985). Weyns’s Duiker is diurnal, like the closely related Peters’s Duiker (Wilson 2001). Foraging and Food  Ripe and unripe fruits are the most frequent foods in the diet of Weyns’s Duiker in the Ituri Forest, comprising between 18% and 100% (mean = 69%) of a sample of rumens across the seasonal fruiting cycle (Hart 1985). In this same sample, foliage, primarily mature leaves fallen from the canopy, comprised the second most frequent food item (annual mean = 23%), with seeds, fallen flowers and fungi comprising the remainder of the diet. Many

Cephalophus weynsi

rumen contents contained small numbers of ants. It is not certain if these were ingested separately from other foods. Kingdon (1982) observed the species feeding on Spathodea flowers in W Uganda. In the Ituri Forest, fruits selected tend to be large to intermediate sized. Important plant families include: Irvingeaceae, Sapotaceae, Euphorbiaceae, Apocynaceae, Annonaceae, Ulmaceae, Sapindaceae and, seasonally, the mast seed-fall of dominant caesalpinaceous canopy species, Julbernardia seretii and Cynometra alexandri. The composition of Weyns’s Duiker diets suggests that the species has a generalized foraging strategy, feeding on a range of species and food patch sizes in proportion to their availability (Hart 1985). Animals captured by hunters during periods of mast fruit seed-fall had layers of fat around their kidneys that were absent at other times of year. Several preferred fruit species (notably, Klainedoxa gabonenis, Ricinodendron heudelotii and Irvingia grandifolia) have large armoured seeds that Weyns’s Duiker regurgitates intact during rumination. These species are adapted to ungulate and elephant dispersal (Feer 1995). Social and Reproductive Behaviour  Radio-collared Weyns’s Duikers in mixed forest in the Epulu area, in the Ituri Forest, occurred in stable, highly cohesive and presumably related parties of 3–4 individuals. Two groups consisted of an adult male–female pair, accompanied by a subadult, and one group of four consisted of a pair accompanied by a subadult and another older animal of undetermined sex. The group of three (adult ? and /, with a subadult /) had individual core home-ranges that overlapped nearly completely over an entire year, and the animals often moved in spatial synchrony. Individual home-ranges in two other social groups also widely overlapped. These synchronized movements suggest strong territorial behaviour, with both sexes taking part in territorial defence. Radio-collared animals had average core home-ranges of 13 ha (adult ??, n = 4) and 11 ha (adult //, n = 5), with total annual occupancy of 26–30 ha (adult ?? and //, respectively) (J. A. Hart pers. obs.). East of the Albertine Rift, populations of Weyns’s Duikers are most often observed as solitary 277

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area (Hart 1979, 2000, J. A. Hart pers. obs.). Pregnant // were recorded in nearly every month. Seven of a total of ten lactating // were recorded during the period Sep–Dec. This is the period of late rains, when fruit supplies are generally at their most abundant. One young is born at a time, and gestation period is not known. Two full-term embryos (1500 g and 1510 g) were recorded in Jun. Weigl (2005) reports longevity at 15 years in captivity. In the Ituri Forest, ?? represented 42% of a total of 55 individuals captured. Of the 23 ??, 52% (n = 12) were adults; in contrast, adult // (n = 24), comprised 75% of a total of 32 // captured (n = 32). The combination of skewed sex ratio, and younger cohort age, suggests higher mortality in ?? in this population.

Weyns’s Duiker Cephalophus weynsi.

animals (Kingdon 1982) or occasionally pairs; however, detailed information on home-range use and social behaviour are not available. Development of group living in at least some populations of Weyns’s Duikers is unusual in forest duikers, and separates this species from the closely related Peters’s Duiker (Feer 1989a). In the Ituri Forest, the small individual home-ranges and defence of shared space, in combination with a generalized diet, may permit Weyns’s Duikers to control areas of relatively higher overall food availability. However, intensive mate-guarding by adult ?? is also not excluded as a driving factor. Group living may also contribute to anti-predator defence. Both ?? and // frequently utter whistled contact calls, in addition to loud bleating calls that they, as well as other duikers, utter when distressed. In the Ituri Forest, where Weyns’s Duiker co-occurs with five other duiker species and the frugivorous Water Chevrotain Hyemoschus aquaticus, group living may provide this species with an advantage in inter-specific competition. In the Ituri Forest,Whitebellied Duikers C. leucogaster occur in reduced numbers in areas where Weyns’s Duikers are abundant in comparison with areas where they are uncommon or absent. Radio-collared White-bellied Duikers avoid areas controlled by Weyns’s Duiker parties. The commitment to social living has restricted this otherwise highly successful species from occupying some habitats. In the Ituri Forest, Weyns’s Duikers are almost entirely absent from large areas of monodominant Gilbertiondendron dewevrei forest. Fruit resources in this forest are rare and widely dispersed over most of the year due to the low floristic diversity. This limitation is alleviated only during brief irregular periods of mast flower and seed production by G. dewvrei. The small, stable home-ranges of Weyns’s Duiker, coupled with their reduced mobility, do not permit the species to take advantage of high-quality ephemeral food sources, unless these occur on the animal’s home-range. Reproduction and Population Structure  In the Ituri Forest, 50% (11 of 22) of adult // examined were pregnant over the course of a two-year study in a moderately to intensively hunted

Predators, Parasites and Diseases  Leopards Panthera pardus are the most important predator of Weyns’s Duikers in the Ituri Forest; however, numbers killed are lower than would be expected given the species’ abundance in the community (Hart, J. A. et al. 1996, Hart 2000). Diurnal habits and group living may contribute to reducing the extent of predation. African Golden Cats Profelis aurata are reported to capture this species occasionally (Y. Sugiyama pers. comm.). One carcass was reported from the nest of Crowned Eagles Stephanoaetus coronatus in Uganda (Skorupa 1989). High frequencies of seropositive titres were recorded for bluetongue, leptospirosis and viral haemorrhagic fevers in a sample of animals tested in the Ituri Forest (Karesh et al. 1995). None of these individuals exhibited outward signs of disease; however, stressed animals frequently succumb to a herpes-like condition when brought into captivity, and latent infections may reduce overall fecundity. Helminths were found in the body cavity, often on the rumen surface of almost all freshly killed, butchered animals observed in the Ituri Forest (J. A. Hart pers. obs.). Conservation  IUCN Category: Least Concern. CITES: Not listed. Weyns’s Duiker is among the primary species hunted by the Mbuti net hunters in the Ituri Forest and populations may be severely reduced where hunting pressure is high (Hart 1979, 2000). Overall, however, this species, like the Blue Duiker, is among the more resilient species to human hunting pressure. Populations in some East African locations are at higher risk, especially in the isolated forest islands at the eastern edge of the range; at least one population in Nyungwe Forest, Rwanda, is suspected to be locally extinct. Otherwise, populations of the species occur in a number of protected areas, including Okapi Faunal Reserve and Maiko and Salonga National Parks (DR Congo), Kibale N. P. (Uganda), Mount Elgon N. P. (Kenya) and Mahale Mountains N. P. (Tanzania) (East 1999). Measurements Cephalophus weynsi WT (??): 17.1 (15.5–19.5.5) kg, n = 12 WT (//): 17.5 (16.0–20.0) kg, n = 22 Ituri Forest, DR Congo (J.A. Hart pers. obs.) Key References  Hart 1985, 2000, 2001; Kingdon 1982; Wilson 2001. John A. Hart

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Cephalophus callipygus  Peters’s Duiker Fr. Céphalophe de Peters; Ger. Petersducker Cephalophus callipygus Peters, 1876. Monatsb. K. Akad. Wiss. Berlin, p. 483. ‘Africa occidentalis (Gabun)’ (Gabon, Gabon River).

Peters’s Duiker Cephalophus callipygus adult male. Lateral view of skull of Peters’s Duiker Cephalophus callipygus adult male. left:

above:

E DR Congo, east of the Cogo and Ubangui Rivers, to the western Taxonomy  Monotypic. Associated with the Natal Red Duiker C. side of the Eastern Rift Valley. Darker pelage with longer dorsal natalensis, Aders’s Duiker C. adersi and Weyns’s Duiker C. weynsi, as a stripe extending in a dark zone on withers; face with blackish tones. potential superspecies (Ansell 1972). Considered to be conspecific with the Natal Red Duiker (Wilson 1987) or not related (Groves C. ogilbyi. The White-legged Duiker (C. o. crusalbum) is sympatric with C. callipygus in Gabon (e.g. Forêt des Abeilles) and NW Congo. & Grubb 1974). Weyns’s Duiker has been described as a subspecies Generally brighter colour with black dorsal line, black muzzle of Peters’s Duiker (Kingdon 1982, 1997) or as a distinct species and marked dark brows; C. o. crusalbum has conspicuous white (Grubb & Groves 2001, Grubb 2005). The latter conclusion has lower hindleg (Grubb 1978b, Gautier-Hion & Gautier 1994). J. been followed in this work. Molecular data have consistently placed Kingdon (pers. comm.) suggests that some individual specimens Weyns’s Duiker as a sister taxon to Peters’s Duiker, within the ‘West of Peters’s Duiker with apparent ogilbyi-like characteristics might African red duiker’ lineage (Jansen van Vuuren & Robinson 2001; and be wild hybrids between the two species. see Hassanin et al. 2012). Synonyms: none. Chromosome number: C. leucogaster. Broadly sympatric west of the Congo and Ubangui not known, but likely 2n = 60. Rivers. Smaller in size, pale body with thick black dorsal stripe, underside sharply white. Description  Medium-sized duiker, with overall pale tawny pelage becoming red on the loins. The forehead and frontal crest C. dorsalis. Sympatric nocturnal species, occurring in forests from Senegal to L. Tanganyika. Similar in size, but shorter-legged and is rich reddish in colour; the crest hair is forwardly curved. Sides smaller-headed, rich dark red body with black dorsal stripe and legs. of face greyish-fawn, and lips, chin and throat white. A fine black dorsal line begins in the middle of the back and widens on the base C. niger. An entirely allopatric duiker from West Africa that resembles this species in morphology and size, but has glossy black pelage of the tail and croup. Underparts lighter coloured than the flanks. and lacks a black dorsal stripe. Legs darker than body, and there is a large blackish tail tuft. Females are about 12% heavier than ??. There are large preorbital glands, covered with short rufous hair, present in both sexes, although those Distribution  Endemic to western central Africa in moist lowland in ?? are larger and more active; the slit is about 40 mm long, forests of S and SE Cameroon, SW Central African Republic, mainland with 13–16 distinct large pores (Wilson 2001). Inguinal pouches not Equatorial Guinea, Gabon and N and SW Congo (East 1999, Wilson present.Well-developed pedal glands present.The newborn is darker 2001). The Sanaga R. appears to be the northerly limit of distribution in colour than adults, and Wilson (2001) notes that a newborn from in Cameroon, although there are apparently museum specimens from the Central African Republic had the dark brown almost black dorsal north of the river (P. Grubb, in Lamarque et al. 1990). It is unlikely that this species occurs in extreme W DR Congo, east of the Ubangui stripe present extending down the back to the tail. Horns present in both sexes, striated with massive transverse R. and north of the Congo R., since there are large collections from ridges, tips slightly converging and curving upward. Skull with this region in the Tervuren Museum in Belgium, which include other strongly developed frontal boss as in Weyns’s Duiker. Preorbital antelopes but not this species (P. Grubb, in von Richter et al. 1990). fossae deeper in ??. Habitat  Peters’s Duikers inhabit moist lowland forest. They prefer primary forest, but are also present in logged forest, probably because Geographic Variation  None recorded. they prefer heterogeneous undergrowth (Feer 1989a). In Gabon, Peters’s Duikers mostly occupy mature forest, but have also been Similar Species Cephalophus weynsi. Allopatric sibling species, occurring in forests from recorded in remote old secondary forest, riverine forest and seasonally 279

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Cephalophus callipygus

inundated terrain (Lahm 1993). In S Cameroon, Peters’s Duikers occur in secondary forest and farm-bush (Fotso & Ngnegueu 1997). Abundance  Peters’s Duiker is the second or the third most abundant duiker in undisturbed forests of NE Gabon, where densities can reach 25/km2 (net capture Dubost 1979), 13–15/km2 (net capture Feer 1988) and 6.7/km2 (line transect Lahm 1993). In forests of S Central African Republic they rank third among duikers and densities are 0.9/km2 (line transect) and 0.9–1.2/km2 (net count Noss 1999). As a result of hunting, densities decrease near villages in NE Gabon (Lahm 1993), in S Cameroon (Muchaal & Ngandjui 1999) and S Central African Republic (Noss 1999). East (1999) estimated a total population size of about 382,000. Adaptations  When alarmed, Peters’s Duikers may freeze before running for cover; they flee quickly at short distances, occasionally with barking. The median and distal segments of the hindleg are relatively longer than in the Bay Duiker Cephalophus dorsalis and correspond to the ‘semi-runner’ type of duiker (Feer 1988). Living under a canopy with numerous fruit-plucking and fruitdropping primates, bats and birds, this species is able to sustain a more frugivorous diet than other species of duiker. Dependence on a year-round fruit supply may explain why this species is mainly restricted to the consistently fruit-rich main forest block in westcentral Africa while its very closely related sibling, Weyns’s Duiker, occupies equivalent areas to the east of its range and the somewhat less closely related, but very similar Black Duiker C. niger occupies the main medium-sized duiker frugivore niche to the west of its range (J. Kingdon pers. comm.). Peters’s Duikers are active during the day, resting from around mid-day to late afternoon. As in other diurnal duiker species, the orbits are less dorsally oriented than in the nocturnal Bay Duiker. Females mostly prefer dense undergrowth, both for diurnal rest and for activity (Feer 1989a).

Foraging and Food  One of the most frugivorous of duikers (Dubost 1984, Feer 1989b, Gagnon & Chew 2000, Wilson 2001). In two studies in Gabon (Dubost 1984, Feer 1989b), fruit comprised between 82.7% and 89.6% of the diet by dry weight based on examination of stomach contents. Leaves comprised between 7.9% and 10.0% of the diet, with petioles and stems following at 6.2%. Fruit and leaves were found in all stomachs in both studies. Flowers, fungi and animal matter (mostly insects) are found in fewer than 50% of samples and comprise less than 1% of the diet (Dubost 1984). Unweaned animals eat much less fruit than adults (52.7%) and more leaves (37.0%) (Dubost 1984). Fruit consumption is lowest from Mar to May (short wet season) whereas leaves are eaten significantly more throughout the rest of the year (Feer 1989b). Most of the fruit consumed are drupes or berries (77%), 1–5 cm in size (80%). Dubost (1984) identified 55 fruit species (n  = 20 stomachs) and Feer (1989b) recorded 110 species (n  = 68).The favoured species of fruits are Xylopia hypolampra, Cylindropsis parvifolia, Canarium schweinfurthii, Klainedoxa gabonensis (Dubost 1984) and Dacryodes büttneri, Santiria trimera, Polyalthia suaveolens and Irvingia gabonensis (Feer 1989b). During rumination, Peters’s Duiker spit out intact protected seeds of ten plant species (Feer 1995).While foraging, the daily mean ranging distance in subadults and adults varies between 2600 m and 4090 m in ?? and between 1870 and 4290 m in // and decreases during the long wet season (Feer 1989a). Peters’s Duiker increases its daily range when food resources become rare. Wilson (2001) mentions an interesting observation of this duiker deliberately hunting and capturing a Hartlaub’s Duckling Pteronetta hartlaubi. Social and Reproductive Behaviour  Peters’s Duikers are most often solitary (67% of observations, Feer 1988). Adult // have stable home-ranges of ca. 40 ha. Male home-ranges are of similar size and include several overlapping ranges of // and young. Adult ?? are very intolerant of other ?? in captivity and their homeranges do not overlap. Female home-ranges are completely separated or overlapping when family bonds exist between them. Both sexes preferentially use locations situated near the perimeter of homerange. Males use a larger portion of their home-ranges on a daily basis than do // (Feer 1989a). Males and // frequently mark by scraping twig bark with their horns and depositing preorbital secretions (Dubost & Feer 1988). Middens or ritualized defecation were not observed. Social play, reciprocal allogrooming and marking of the partner were observed in captive animals (Dubost & Feer 1988). Reproduction and Population Structure  Histology suggests a sexual maturity at 17 months of age for ?? and ca. 20 months for //. In Gabon, reproduction occurs all year round, although births peak twice a year at the beginning of the dry seasons when fruit availability or food qualitative richness is most favourable (Dubost & Feer 1992). Around 29% of // over 20 months old are pregnant. This figure seems underestimated possibly because of a bias of the sample related to lower mobility, and thus less capture of some pregnant animals (Feer 1988). Two near-term foetuses measured by Wilson (2001) had masses of 2.7 kg and 2.9 kg. Weaning takes place when the maximum amount of fruit resources or leaves is available (Dubost & Feer 1992). In Gabon, a sample of 339 individuals from a moderately hunted population comprised 35% adults; ??

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accounted for 43.5% of adults. However, in a sample captured by nets, the proportion of ?? was 27.3%, corresponding more to the observed social life pattern. Mortality of subadult and young adult // is higher than in ?? (Feer 1988). Predators, Parasites and Diseases Peters’s Duiker is potentially preyed upon by Leopards Panthera pardus (Hart, J. A. et al. 1996); hairs of red duikers were observed in a felid scat in Gabon (F. Feer pers. obs.). Diseases and parasites in the wild are unknown. Conservation  IUCN Category: Least Concern. CITES: Not listed. The primary threats to the survival of the species are habitat loss due to human settlement and hunting (Lahm 1993, East 1999). Because Peters’s Duiker prefers undisturbed primary forest, its future survival may be increasingly dependent on protected areas. At present the protected areas within which it is common, for example Dja Wildlife Reserve and Lobéké N. P. (Cameroon), Dzanga-Sangha Special Reserve and Dzanga-Ndoki N. P. (Central African Republic), Odzala and Nouabalé-Ndoki National Parks (Congo), Lopé, Ivindo, Minkébé, Loango and Moukalaba-Doudou National Parks (Gabon) and Monte Alén N. P. (Equatorial Guinea), receive unequal levels of protection and management. Peters’s Duiker is frequently the most abundant medium-sized duiker species in undisturbed areas, but its populations are heavily harvested. Peters’s Duiker is especially affected by snare hunting. In the Central African Republic, Peters’s Duikers accounted for 29% of all animals captured in snares, a level that appears to be unsustainable (Noss 1998a). In Gabon, the proportion of Peters’s Duikers among duikers captured in snares is five times the proportion harvested with guns (Lahm 1993). In N Congo, they comprised 21% of all animals

killed with shotguns (Mockrin et al. 2011). In Dja Reserve they are the most frequently captured species and accounted for 48% of duikers hunted (Fotso & Ngnegueu 1997), but at another site in the same area this species accounted for 20% of duikers killed (Muchaal & Ngandjui 1999). In Gabon, Peters’s Duikers comprised 9% of duikers caught by village hunters and trappers (Lahm 1993). In Congo, Peters’s Duikers represent only 4% of duikers for sale in Brazzaville bushmeat markets (F. Feer pers. obs.) and is rarer in Pointe Noire (Wilson & Wilson 1991). Peters’s Duiker ranks between third and fifth among the game species proposed in Libreville (Gabon) markets, with a mean number of animals sold ranging from 2.0 to 6.6 per day (Ntsame Effa 2005). Measurements Cephalophus callipygus HB (??): 977 (940–1060) mm, n = 11 HB (//): 1025 (970–1090) mm, n = 14 T (unsexed): 120–150 mm HF c.u. (??): 251 (240–270) mm, n = 7 HF c.u. (//): 250 (240–260) mm, n = 7 Sh. ht (??): 532 (490–550) mm, n = 12 Sh. ht (//): 549 (510–570) mm, n = 13 WT (??): 19.6 (17.5–21.5) kg, n = 10 WT (//): 21.9 (18.6–25.2) kg, n = 7 Gabon (Feer 1979, F. Feer pers. obs.) Maximum recorded horn length is 14.9 cm for a pair of horns from Lomié, Cameroon (Rowland Ward) Key References  Dubost & Feer 1988; Feer 1988, 1989a, b; Wilson 2001. François Feer & Miranda Mockrin

Cephalophus niger  Black Duiker Fr. Céphalophe noir; Ger. Schwarzducker Cephalophus niger (Gray, 1846). Ann. Mag. Nat. Hist., ser. 1, 18: 165. ‘Guinea’, but apparently Ghana, Shama.

Taxonomy  Seven years after Gray first named this species, Temminck described it under the name Antilope pluto. According to Grubb (2005), the type ‘came from the Leiden Museum, one of a series from Chama (= Shama) and Dabocrom, Ghana, including the syntypes of pluto. Only the specimens from Dabocrom were retained in Leiden (Jentink, 1892) so presumably the type is from Shama.’ In morphology, size, behaviour and ecology this duiker is the Upper Guinea equivalent of Peters’s Duiker C. callipygus and Weyns’s Duiker C. weynsi. In colour and in a smooth neck and swollen nostrils it resembles the larger Abbott’s Duiker C. spadix and Yellow-backed Duiker C. silvicultor. In some formulations of their mtDNA studies, Jansen van Vuuren & Robinson (2001) suggested that the callipygus/ weynsi and spadix/silvicultor lineages were widely divergent whereas in other, more plausible, topologies they associated all these large, smooth-necked duikers in a single but complex cluster of species. Synonyms: pluto. Chromosome number: 2n  = 60; the X chromosome is a submetacentric (Hard 1969). Black Duiker Cephalophus niger.

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Lateral view of skull of Black Duiker Cephalophus niger.

Description  A heavily built, long-bodied, long-headed, glossy black duiker with swollen nostrils and relatively short stocky legs. Short hair on the face is often very thin or absent, probably abraded by frequent rubbing of the face and preorbital glands. Bridge of the nose black or dark brown graduating to a coronal tuft of dense reddish hair; rest of the face dull grey or, in some individuals, red. Lower jaw and upper throat off-white, dull cream or grey. Backs of ears dark brown, anterior surfaces with short, sparse off-white hairs. Neck black or brown, covered in short hair, and with thickened skin. Back intensely black, long and coarsely haired; legs, flanks and underside black or very dark brown with rufous tinge on upper, inner surfaces and on buttocks. Individual hairs have straw-coloured bases (Grubb & Groves 2001). Wilson (2001) comments that of 186 adults examined across the range of the species, only two exhibited any signs of small patches of white on them: a ? from Kumasi had a circle of white hair on the left hindleg near the tail, and another ? from Cape Coast in Ghana had three irregular patches of white hair on the left side between the front- and hindlegs. Tail brown and thinly haired except for a terminal tuft, which has a whitish tip. Juveniles similar in colour, but with less red on face and paler below (Grubb & Groves 2001). Sexes of similar size, although // are on average larger, especially in mass (Wilson 2001); ?? have longer, more robust horns. Wilson (2001) notes that the average preorbital gland of an adult ? was 500 mm long, with a single line of 35–50 pores, from which a white secretion could be squeezed; preorbital glands are smooth and the area surrounding the slit and pores is naked. Pedal glands are present on both fore- and hindfeet. No trace of inguinal glands has been found. Grubb & Groves (2001) include among distinguishing charac­ teristics of the skull: preorbital pits extending forward to premaxillae, and downward to molar alveoli; a well-defined groove between frontals, which are thickened and slightly convex; zygomatic arch straight; and supraorbital foramina often partly roofed over by lateral extension of frontal thickening. Geographic Variation  No significant variation has been recorded, but individuals vary in intensity of black and in the extent of rufous colour on head. Similar Species Cephalophus callipygus. An entirely allopatric duiker from western

central Africa that resembles this species in morphology and size, but has rich red colouring and a black dorsal stripe. C. ogilbyi. Brooke’s Duiker (C. o. brookei) is sympatric from Sierra Leone to Ghana. A smaller, almost extinct sympatric duiker of reddish colour. C. spadix. A larger, allopatric duiker from East Africa that closely resembles this species in colour, but is distinctly larger.

Cephalophus niger

C. silvicultor. A much larger sympatric species with prominent yellow back. Distribution  Endemic to Africa, occurring in forested and formerly forested areas from near Kindia in SW Guinea and eastwards through Sierra Leone to the Niger R.; there are no confirmed records from Benin or from Burkina Faso (East 1999, Wilson 2001). There is no indication that the species has ever occurred east of the Niger R., and reference to the species occurring in Cameroon (Jeannin 1936) are in error; similarly, records from Boshi–Okwango forests on the Nigerian border with Cameroon cannot be accepted (Anadu & Green 1990). It is particularly common and successful in the central parts of its range, from Liberia to Ghana, but is rare or declining both east and west of this heartland. Habitat  Primarily found in disturbed and secondary forest, but also inhabits forest galleries, thickets and is especially common in secondary ‘farm-bush’. Black Duikers are also found, albeit more rarely, in primary rainforest (Jeffrey 1974). In Liberia, Peal & Kranz (1990) described them favouring secondary bush and riparian habitats. In Côte d’Ivoire, the Black Duiker was one of the two most frequently observed species in mixed farmland mosaics (along with Maxwell’s Duiker Philantomba maxwelli), and the least frequently observed of the five small- and medium-sized species present in closed-canopy forest (Newing 2001). In a study of 251 carcasses in Ghanaian markets and 41 sightings, Wilson (2001) recorded the majority from various categories of secondary vegetation; less than 10% came from unlogged high forest. Of a smaller sample of 46 records, Anstey (1991) found only 13% in high forest.Wilson (2001) attributes these habitat preferences to a greater variety of food plants and to the denser cover growing under a felled or partial canopy. Of special interest is the role of an American invasive hemp-like species, Chromolaena odorata, which forms dense thickets after forests have been cleared. For Black Duikers the attraction of these ‘Akyempong

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weed’ thickets seems to be the cover they provide as the plant did not feature in an exhaustive study of Black Duiker diets (Wilson 2001). Abundance  The Black Duiker’s preference for regrowth after forest felling probably led to initial increases in abundance throughout the former forest zones of Upper Guinea, but intensification of agriculture and the spread of vehicles, road networks, markets, firearms, spotlights and snares, all in the service of an increasing human population, eventually offsets initial increase. Once regarded as extremely abundant in Sierra Leone, where eight species of duiker were known, Stanley (1925) reported ‘by far the commonest are the black duiker and Maxwell’s duiker (the bushgoat and fritambo of the Creoles), which are found everywhere in Sierra Leone where there is plenty of cover, preferably forest regrowth’. By 1990 this species had become restricted to isolated pockets where it continued to decline (Teleki et al. 1990). In Liberia, Anstey (1991) considered it the most widely sighted duiker, and the second most frequently recorded in bushmeat markets in Liberia. In Côte d’Ivoire, Newing (2001) recorded the species as being very common in secondary habitats. Wilson (2001) suggested that the Black Duiker was more common in Ghana than anywhere else in West Africa, and the second most common forest duiker in the country after Maxwell’s Duiker. This species is still abundant in many other parts of its range and a density of 2/km2 in the more favourable localities has been suggested by East (1999), who estimated the total population to be about 100,000. Adaptations  There are interesting questions raised by a switch from predominantly red to black colouring in several lineages of duikers. Within the callipygus/weynsi/niger complex, melanic colouring occurs in the far west and far east of their range. Both Peters’s Duiker and Weyns’s Duiker are vividly coloured red duikers, but one race of the latter, C. w. lestradei, and occasional individual variants from other parts of this species’ range are very dark brown or nearly black (a change in colouring that seems to derive from extensions of the very variable black dorsal stripe). It has been reported that the Black Duiker is nocturnal (Sclater & Thomas 1899) or crepuscular (Wilson 2001), and it could be argued that dark colouring might represent a type of crypsis appropriate to this activity, with predation by Leopards Panthera pardus providing the principal selective pressure. However, captives in Monrovia Zoo were found to be active at night only 24% of the time and 69% during the day (Newing 2001), and other authors (Dunn 1991, Anstey 1991, and see Hoppe-Dominik et al. 2011) have recorded diurnal activity, so the link between dark colouring and nocturnal habits seems unlikely. An alternative explanation could be that visual communication has some intra-specific utility for bright red duikers that live at high densities in relatively open undergrowth, whereas equally large numbers of duikers living in denser vegetation rely more on auditory, olfactory or indirect clues to regulate intra-specific contacts. If this is so, then dark colouring could be characterized as a sort of ‘cancelling out’ of the visual channel, except at very close quarters, where ear and face markings can enhance expressions and intention movements. This species shares with both Peters’ andWeyns’s Duikers a strongly reinforced forehead that is especially thick in ??. This peculiarity implies adaptation to exceptionally severe concussion during intraspecific fighting. The special development of this reinforcement in

?? makes it less plausible that it evolved to break open large, hard fruits or even to butt fruit-bearing trees. These might, of course, be secondary benefits of hard-headedness. To date neither aspect of behaviour has been confirmed, but there are widespread hunter’s stories of this species felling banana Musa paradisiaca and pawpaw Carica papaya trees to get at the fruit (Wilson 2001). Assuming that the selective pressure favouring reinforcement of the forehead was the ability to withstand head-butting, especially among ??, the frequency of confrontations implies either a naturally high density of territorial ?? or a high level of competition for //. What little is known of all three species supports the former explanation. A significant difference, however, is that Peters’ and Weyns’s Duikers inhabit well-developed, multi-species mature high forest whereas the Black Duiker prefers thick secondary growth, a preference that places it closer to Abbott’s Duiker and the Yellow-backed Duiker. This suggests that the Black Duiker occupies an ecological position in between the two giant duikers and the larger red duikers of the main forest block. This conclusion has some support from the molecular trees of Jansen van Vuuren & Robinson (2001) and there is the implication that with phylogenetic enlargement of duiker body sizes there may have come a point where high-quality diets might have had to give way to less selective, broader-band feeding habits, especially in disturbed or secondary forests. If that is the case, the Black Duiker’s biology might illustrate some of the conditions that favoured increase in the size of duikers. Black Duikers are well known for digging up subterranean plant parts and for employing their sharp hooves to slash and break up edible roots. Among the tubers regularly excavated and eaten by Black Duikers are those of the introduced exotic cassava Manihot esculenta. As these tubers are known to harbour strong toxins in their unprocessed state it is clear that the Black Duiker has a natural resistance to certain plant poisons (Wilson 2001). Foraging and Food  Fallen fruits, flowers and leaves from the forest floor, as well as bulbs, rhizomes, fungi, bark and occasional animal matter, make up the diet. In Liberia, Black Duikers have been recorded following Western Pied Colobus Colobus polykomos for canopy waste, presumably mostly leaf and stem material. The bulk of their diet is fruit, and every one of 131 stomachs retrieved from Ghanaian markets and examined by Wilson (2001) contained pieces of fruits or whole fruits and seeds. Figs Ficus spp. were a major food throughout the year, as were Nauclea latifolia, Canthium vulgare, Lecaniodiscus cupanoides, Blighia sapida and Solanum spp. Other more seasonal fruits were Cola gigantea (May, Jun and Sep– Dec), Anchomanes difformis (Nov–Mar), Griffonia simplicifolia (Oct– Mar), Musanga cecropoides (Feb–Mar), Alchornea cordifolia (Feb–Apr), Phyllanthus discoideus (Jun–Aug), Antholeista djalonensis (Mar–May) and Canavalia enisformis (Sep–Nov). This spread of wild fruits was augmented by visits to cultivated crops; in 35 of the stomachs, the bulk of the food was from cultivation. The most important of these were: oil nut palm dates Elaeis guineensis, avocados Persea americana, cassava, coffee Coffea cane­ phora, new cocoyam Xanthosoma sp., cocoa Theobroma cacao, pawpaws, bananas and cow-itch Mucuna pruriens. As a very high proportion of these food plants are exotic to Africa, it is clear that Black Duikers have a naturally catholic diet and, in spite of intensive hunting, have found some benefits from partial forest clearance and low-intensity 283

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agriculture.Wilson (2001) found animal matter in 15% of his sample and he also recorded Black Duikers feeding on an open sea-side beach where he thought minute shells (perhaps salt-retaining types?) were the attraction. In total, Wilson (2001) recorded some 27 species of fruit or cultivated crops eaten. In a similar study, Hofmann & Roth (2003) examined 57 stomachs and recorded a total of 33 different types of fruit eaten. Their study also confirmed that Black Duikers frequent cultivated lands for feeding and are not dependent on primary forest habitat for food; leaves of cassava were found in 53% of stomach samples, and leaves of Alchornea cordifolia in 18% of stomachs. Black Duikers are believed to play an important role in the dispersal and germination of small seeds, such as Solanum verbascifolium (Alexandre 1982). Newing (2001) found that Black Duiker mouths and teeth could break open food items up to a diameter of 6 cm.

Servals Leptailurus serval, African Golden Cats Profelis aurata, Eagleowls Bubo spp., Martial Eagle Polemaetus bellicosus and Crowned Eagle Stephanoaetus coronatus. Ntiamoa-Baidu et al. (2005) recorded the following ixodid tick species from animals in Ghana: Haemaphysalis parmata, H. leachi, Ixodes muniensis, I. moreli, I. aulacodi, Rhipicephalus ziemanni, R. simpsoni and Amblyomma compressum.

Social and Reproductive Behaviour  A territorial and mainly solitary species: of 53 sightings made by Wilson (2001), 39 were of single animals, ten were of male–female pairs (three accompanied by an infant) and four were of // with young. Duikers in general are known to respond to bleat-like calls and this susceptibility is widely exploited by hunters. The Black Duiker appears to be particularly responsive to artificial lures blown on whistles that are cut from fern stems in Liberia (Wilson 2001). What attention-getting calls could signify in the social and spatial structures of a solitary species remains to be elucidated.

Conservation  IUCN Category: Least Concern. CITES: Not listed. Their adaptability to degraded and secondary forests has enabled Black Duikers to withstand settlement better than other mediumsized forest duiker species in West Africa, and while they have been eliminated from the more densely settled parts of their range they still occur relatively widely within their historical range. They also show resilience to hunting and remain locally common; they have been recorded as extirpated from several reserves in Ghana (e.g. Digya N. P. and Kalakpa Resource Reserve; Wilson 2001), but in fact do still persist in these areas (R. J. Dowsett pers. comm.). Dunn (1991) noted that long after all primates had been shot out in parts of Liberia, the Black Duiker persisted. None the less, East (1999) has warned ‘If current trends continue, including a complete lack of effective protection and management over most of its range, its status will eventually decline to threatened.’ Black Duikers are well represented, generally in stable numbers, in protected areas such as Sapo N. P. (Liberia), Western Area F. R. (Sierra Leone), Taï N. P. and Comoé N. P. (Côte d’Ivoire) and Bia, Nini-Suhien and Kakum National Parks (Ghana).

Reproduction and Population Structure  Although preg­ nant // have been recorded in every month of the year, Wilson (2001) recorded a very marked birth peak between Nov and Jan (the wet season) based on reproductive tracts of 106 sexually mature // collected mainly in Ghana. Of the // examined, 72.6% were either pregnant or lactating and of these nearly 10% were both pregnant and lactating. Females are capable of conceiving at one year of age (Wilson 2001). Such fecundity helps explain how Black Duikers have managed to survive very high mortality rates from the bushmeat trade in West Africa. Although ovulation had taken place in both ovaries, Wilson (2001) found that single foetuses were always implanted in the right horn of the uterus; he found that neonates weighed between 1.65 and 2.31 kg at birth (with no difference between the sexes). Captive Black Duikers weigh 1.4–2.2 kg at birth, doubling in weight after the first month (Barnes et al. 2002). Weaning occurs at between 80 and 108 days (Wilson 2001). Captive animals have lived to more than 14 years (Weigl 2005).

Measurements Cephalophus niger TL (??): 1060 (1020–1100) mm, n = 30 TL (//): 1080 (1020–1160) mm, n = 44 T (??): 100 (90–120) mm, n = 30 T (//): 110 (100–140) mm, n = 44 HF c.u. (??): 250 (240–260) mm, n = 30 HF c.u. (//): 250 (240–260) mm, n = 44 E (??): 94 (91–99) mm, n = 30 E (//): 95 (90–100) mm, n = 44 Sh. ht (??): 460 (450–480) mm, n = 30 Sh. ht (//): 470 (440–500) mm, n = 44 WT (??): 21.0 (19.0–23.0) kg, n = 30 WT (//): 24.0 (17.0–26.0) kg, n = 44 Ghana (Wilson 2001) Maximum recorded horn length is 17.46 cm for a pair of horns from Ghana (Rowland Ward)

Predators, Parasites and Diseases  Lions Panthera leo, Leopards and African Rock Pythons Python sebae are known predators (Wilson 2001, Bodendorfer et al. 2006). Young may be predated upon by

Key References  Newing 2001; Wilson 2001. Jonathan Kingdon & Michael Hoffmann

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Cephalophus spadix ABBOTT’S DUIKER Fr. Céphalophe d’Abbott; Ger. Abbottducker Cephalophus spadix True, 1890. Proc. U.S. Natl Mus. 13: 227. ‘High altitudes on Mt. Kilima-njaro, frequenting the highest points’ (Tanzania, Mt Kilimanjaro; at 2400 m according to Grimshaw et al. 1995).

Lateral view of skull of Abbott’s Duiker Cephalophus spadix.

Abbott’s Duiker Cephalophus spadix.

The common name is after W. L. Abbott, an American naturalist and explorer who spent 1888–89 in the Kilimanjaro district, collecting zoological specimens. Taxonomy According to Sclater &Thomas (1899), F.True suggested that ‘C. spadix was closely allied to C. niger and even possibly identical with it’. Roosevelt & Heller (1915) suggested that it was more closely related to the group of giant duikers of which theYellow-backed Duiker C. silvicultor is typical, and Haltenorth (1963) considered Abbott’s Duiker a subspecies of silvicultor. Kingdon (1982) indicated that Abbott’s Duiker was closely related to theYellow-backed Duiker and probably represented a relic of an ancestral population.Wilson (2001) interpreted the skull morphology to suggest that Abbott’s Duiker is primitive with respect to both the Yellow-backed Duiker and the Bay Duiker and considered it a near-perfect morphological ancestor. A cladistic analysis that considered 41 morphological characters highlighted the monophyly of the silvicultor group and, within this, a split between dorsalis versus jentinki/silvicultor/spadix (Grubb & Groves 2001). Molecular phylogeny of seven duiker species representative of four supposed adaptive lineages has confirmed the existence of a distinct, giant duiker lineage that includes silvicultor, spadix and dorsalis (Jansen van Vuuren & Robinson 2001). Synonyms: none. Chromosome number: not known. Description A large, stocky duiker with a glossy, dark brown pelage, especially on the back, flanks and upper legs. Neck and face are paler, greyer brown. A prominent russet hair tuft between the ears is dense, tinged with black and usually lighter basally.Throat, ventrum and inner thighs are sometimes paler brown, with white fur in the genital area. The extent of the paler ventral hair can vary between individuals within the same population (T. Davenport & S. Machaga unpubl., F. Rovero pers. obs.). The wedge-shaped head ends with a broad, flat-

fronted nostril pad that overhangs the mouth (as with Yellow-backed Duikers). The rhinarium is shiny black. Ears are rounded with naked and grey-pinkish inner surfaces. Lower parts of the legs are slightly darker brown to almost black and end in black hooves, the front ones longer than the hind ones. Tail short, dark grey to black on the outer, dorsal portions and paler grey to white on the inner surfaces and tip. A six-month old juvenile from West Usambara appeared uniformly paler brown than adults, including head and neck, and had a darker brown dorsal stripe. The skin of a juvenile from Mount Rungwe exhibited longer fur than adults (T. Davenport & S. Machaga unpubl.). Kingdon (1982) mentioned ‘a hint of the yellow-back of silvicultor is visible in the form of a small grey patch above the root of the tail’, and Grubb & Groves (2001) mention a ‘vague dark dorsal stripe’ and note that two skins in the British Museum show what may be rudiments, or precursors, of the silvicultor dorsal markings at the base of the tail. Wilson (2001) found no signs of such markings in two adult !! examined and no markings were evident on at least four individuals camera-trapped in the Udzungwa Mts (F. Rovero pers. obs.). However, in three of five skins from Mt Rungwe, a very slight, approximately 5 cm-wide, darkening of hair along the spine from the lower neck to the rump is visible, although there is no obvious grey patch above the root of the tail (T. Davenport & S. Machaga unpubl.). Both sexes possess horns (8–12 cm long), which are mostly hidden in the hair tuft. Horns are without any conspicuous thickening at their base. GeographicVariation No subspecies have been described and no geographical variation is reported. The coat colour of a captive adult pair from West Usambara appeared remarkably darker brownish, almost black, with less evident, contrasting paler underparts, than several individuals camera-trapped in the Udzungwa Mts (F. Rovero pers. obs.); the same is true of Mt Rungwe animals (T. Davenport & S. Machaga unpubl.). It is likely that variation in coat colour occurs within populations as much as between populations. Similar Species Cephalophus niger: Allopatric species from West Africa. Considerably

smaller, the body is black with more contrasting paler neck and head, and similar reddish hair tuft. C. silvicultor. Allopatric species from the Guineo-Congolian forests. Slightly larger, greyish-brown with a vivid cream-coloured patch on the back. 285

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Habitat Typical habitat is montane and sub-montane moist forest; in Mt Kilimanjaro it is reported as commonest between 1300 and 2700 m (see Grimshaw et al. 1995) in forest and high-altitude swamps, scrub and moorland. In the Udzungwa Mts, Abbott’s Duiker has been recorded as low as 300 m in Matundu Forest, a large, lowland and semideciduous forest (Rovero & Marshall 2009), as well as on the highest peak (Mt Luhombero, 2600 m; Rodgers & Swai 1988). It has been camera-trapped on several occasions in semi-deciduous to evergreen forests with dense understorey in Matundu and Mwanihana forests at 500–800 m in Udzungwa Mountains N. P. (Rovero et al. 2005, F. Rovero pers. obs.). On the Uzungwa Scarp it has been sighted on bamboo-dominated ridges at 1700 m (J. Fjeldså pers. comm.). It is known from disturbed and secondary montane forest and bamboo forest to 2500 m and occasionally plateau grassland to 2800 m on Mt Rungwe and in Livingstone–Kitulo in the Southern Highlands (T. Davenport & S. Machaga unpubl.). These records suggest that Abbott’s Duiker, if undisturbed, occurs also in lowland, semi-deciduous forest with clearings and large areas of secondary vegetation.

Cephalophus spadix

Distribution Endemic to Africa, being confined to a few isolated forested massifs in N, E and SW Tanzania (Kingdon 1982, 1997, East 1999). It is currently reported from Mt Kilimanjaro, W Usambara Mts, Rubeho Mts, Udzungwa Mts and Mt Rungwe (Wilson 2001, Moyer 2003, F. Rovero pers. obs.), as well as the forests of Livingstone (now part of Kitulo N. P.), Irungu, Irenga, Ndukunduku and Madehani in the Southern Highlands (Wilson 2001, Machaga & Davenport 2004, T. Davenport & S. Machaga unpubl.). Surveys in the Udzungwa Mts have extended their known distribution to nine discrete forests (Jones & Bowkett 2012). However, recent work in Madehani failed to record the animal (T. Davenport pers. obs.). In the Uluguru Mts, the most recent published record is from Swynnerton & Hayman (1951), and no signs were obtained during extensive surveys in the last few years (Frontier-Tanzania 2005), including a camera-trap survey in Uluguru North F. R. in 2005 (F. Rovero & A. Bowkett unpubl.). A previously unknown population of Abbott’s Duiker, estimated at a maximum of 50 individuals, was discovered in 2006 in a montane forest locally called ‘Ilole’, in the southern Rubeho Mts (Rovero et al. 2008), while no records are reported from other forests of this area (Doggart et al. 2006, F. Rovero pers. obs.). Among other highland forests of Tanzania, Abbott’s Duiker was not reported to occur from recent surveys in North Pare, South Pare, Nguru, Nguu, Ukaguru and Mahenge Mts (Tanzania Forest Conservation Group unpubl., Frontier-Tanzania unpubl.). The historical distribution also included forests on the Rift Wall near Babati in C Tanzania, forests above L. Manyara, the Poroto Mts and Mfrika Scarp in the Southern Highlands (Rushby & Swynnerton 1946, Swynnerton & Hayman 1951), as well as Mporoto, Mpara, Sawago and Njombe Forest Reserves and possibly Mbeya Range (T. Davenport & S. Machaga unpubl.). However, it is probable that they have been extirpated from all these sites (Rovero et al. 2005). Reports of this species being present in Sierra Leone and Ghana are clearly in error, and likely refer to Black Duiker C. niger (see references in Grubb et al. 1998).

Abundance Abbott’s Duiker is one of the rarest duiker species (East 1999). However, there is very little information to infer reliable estimates of population densities. It is sighted too infrequently to be counted using diurnal, line-transect methods (Rovero & Marshall 2004), while camera-trapping is proving a potentially useful technique (Rovero et al. 2005). East (1999) used comparative data to derive a conservative estimate of 1 individual per km2 in optimal habitat. Camera-trapping data from the Udzungwa Mts fed into a regression equation relating sighting rates to estimated densities obtained for Harvey’s Duiker Cephalophus harveyi suggest maximum densities of 1.3 Abbott’s Duikers per km2 (F. Rovero pers. obs.). Albeit preliminary, this figure matches the estimation given by East (1999). From in-forest follows, tracking, habitat suitability analyses and hunting surveys, it is estimated that there are fewer than 40 individuals throughout Mt Rungwe and Livingstone (T. Davenport & S. Machaga unpubl.). East (1999) and Moyer (2003) gave total population size in the range of 1500–2500 individuals based on estimated area of occupancy. However, these did not account for the dramatic decline of Abbott’s Duiker due to hunting in potentially suitable habitat, especially over the last decade (e.g. Uzungwa Scarp F. R., Mt Rungwe and Livingstone, West Usambara Mts and West Kilimanjaro). Therefore, although the total population size is unknown, it is probably less than 1500 individuals (Rovero et al. 2005). Adaptations This is an extremely secretive species, occurring at low densities and very rarely seen even where it is considered relatively common. Furthermore, it appears to be mainly nocturnal and crepuscular (F. Rovero pers. obs.), and capture times of 25 photographs from several sites in Udzungwa Mts N. P. show that 52% were taken at night, 16% from 06:00h to 07:00h and from 18:00h to 19:00h, and 32% in the day (F. Rovero pers. obs.). Foraging and Food Kingdon (1997) indicates that the diet of this species includes fruits, flowers, green shoots and herbage, and that it has been recorded browsing the leaves of a balsam (Impatiens elegantissima). In the Udzungwa Mts, it has been seen browsing both on understorey leaves in closed forest and on marshy vegetation in forest clearings (T. Jones & F. Rovero pers. obs.), and one individual was camera-trapped

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with a large frog in its mouth (Rovero et al. 2005). Wilson (2001) reports Abbott’s Duikers on the lower slopes of Mt Kilimanjaro cropraiding and feeding on sweet potato leaves and tubers, bananas, cassava leaves and cowpeas; he also saw an adult Abbott’s Duiker eating pieces of green moss from rocks. In the Southern Highlands on Mt Rungwe, Abbott’s Duikers have been reported eating various balsams Impatiens spp., the climbing herb Begonia meyeri-johannis and beans from local shambas (Machaga & Davenport 2004). Social and Reproductive Behaviour There is an almost complete lack of behavioural notes on Abbott’s Duiker. All sightings and camera-trap records are of single individuals except for one mating pair (see below) and a ! with a two-month-old juvenile (Wilson 2001). It is likely that Abbott’s Duiker is mainly solitary, as claimed by all hunters of the species in the Southern Highlands (T. Davenport & S. Machaga unpubl.) and as reported for the Yellow-backed Duiker (Kingdon 1997). When disturbed, no alarm vocalization is usually emitted (T. Jones pers. obs.), in contrast to other forest antelope species. In the Udzungwa Mts, an adult Abbott’s Duiker was briefly observed following a large group of Sanje Mangabeys Cercocebus sanjei, which were foraging on the forest floor (T. Jones pers. obs.). Reproduction and Population Structure A camera-trap photograph of a mating pair was taken on 27 Jan in Mwanihana Forest, Udzungwa Mts (F. Rovero & T. Jones pers. obs.). Wilson (2001) reports two births that occurred in Aug/early Sep and suggests that as Abbott’s Duikers are found mainly in wet and fairly moist habitats they may not have a fixed breeding season, and as claimed by hunters in the Southern Highlands (T. Davenport & S. Machaga unpubl.). A pair kept in captivity was observed mating on 28 May, and a birth occurred on 13 Jun, these observations being recorded in different years (J. Beraducci pers. comm.). Predators, Parasites and Diseases The main non-human predator of Abbott’s Duiker is the Leopard Panthera pardus (Wilson 2001, Rovero et al. 2005, D. Moyer pers. comm.). In the Udzungwa Mts, Lions Panthera leo and Spotted Hyaenas Crocuta crocuta are also potential predators. Juveniles are probably predated by Crowned Eagles Stephanoaetus coronatus and pythons Python spp. Conservation IUCN Category: Endangered C2a(i). CITES: Not listed. Wilson (2001) reports Abbott’s Duikers as having been quite common until the early 1960s and mentions frequent records of hunted animals from several sites until the late 1980s.There is little doubt that increasing hunting, habitat destruction, alteration and fragmentation have caused a dramatic decline in the last few decades in the abundance of most populations and probably their extirpation from many areas. This is the case of small/marginal forests in the Udzungwa Mts (Kigogo, Mufindi Scarp East and Mufindi Scarp West Forest Reserves; Moyer 2003), many sites in the Southern Highlands, and Chonwe Forest in the Uvidunda (Bismark) Mts. Near-extinction is probably occurring in the Uluguru Mts, Uzungwa Scarp F. R., Kising’a-Lugalo F. R., New Dabaga-Ulang’ambi Forest in the Udzungwa Mts and throughout the Southern Highlands (Bowkett et al. in press). A comprehensive survey of duiker hunting was carried out in Mt Rungwe and the adjacent Livingstone forest in 2003 (Machaga

& Davenport 2004, T. Davenport & S. Machaga unpubl.). All 114 hunters interviewed reported having seen Abbott’s Duiker. Some 77% hunt for domestic purposes, whereas 23% hunt for commercial purposes. Abbott’s Duiker is not sought after for medicinal properties, but its skin is used in making drums. In 2003, an Abbott’s Duiker carcass was valued in the villages at $US20–30, although only 3% of hunters successfully trapped one that year. The vast majority used snares and the number of snares laid per hunter ranged from 300 to 1000. All hunters had noted a dramatic reduction in the number of Abbott’s Duikers caught, especially over the last decade, and only 5% said hunting was still worthwhile. The heavy hunting pressure in unprotected forests has probably left the Kilimanjaro and Udzungwa populations as the two strongholds of this species, while in other areas it most likely persists at very low densities. The current distribution on Mt Kilimanjaro is not known, but the area is mainly protected as a national park. Udzungwa Mountains N. P. holds a healthy population of Abbott’s Duikers, while the species’ presence in adjacent, unprotected forests such as Uzungwa Scarp F. R. is severely threatened by recent and alarming increases in levels of poaching (Moyer 2003, Rovero et al. 2005, 2010). However, a recent, preliminary study on the genetic structure of the Udzungwa population found that genetic diversity values are low relative to Harvey’s Duiker in the Udzungwa Mts or published values for other mammal species (Bowkett et al. in press). This may be the result of habitat fragmentation and the resulting isolation of small subpopulations. In the West Usambara Mts, heavy hunting is seriously threatening the survival of this population, and recent interviews of hunters indicate that Abbott’s Duikers might be left only in Shume-Magamba F. R. (F. Rovero pers. obs.). The situation in the East Usambara Mts is reported to be even more critical (Moyer 2003), and Abbott’s Duiker is very likely to have been extirpated there already. Livingstone Forest, now part of Kitulo N. P., is the only Southern Highland habitat that is currently being effectively managed, although moves to upgrade the protected area status of Mt Rungwe are under way (T. Davenport pers. obs.). Unless major conservation efforts are urgently applied within the next few years, Abbott’s Duiker will probably become restricted to the Udzungwa Mts and Mt Kilimanjaro (Moyer 2003). Thus, coordinated survey work should be carried out throughout its range to assess current status and distribution. Conservation and education initiatives should also be instigated as a matter of priority. Current work in the Southern Highlands employing hunters in environmental education initiatives in exchange for stopping hunting has met with some success. It is too early to say if this will have a significant positive impact on Abbott’s Duiker populations, although it may prove a valuable model for conservation at other sites. Major conservation management measures that would enhance the protection of Abbott’s Duiker are increased protection to include important forests in the Udzungwa Mts currently poorly protected, in particular Uzungwa Scarp, Iyondo and Matundu (Sumbi et al. 2005). The expansion of Kilimanjaro N. P. to annex lower altitude forests has been recently gazetted. Also necessary is the inclusion of Mt Rungwe within the new Kitulo N. P. and greater law enforcement enacted in those areas that are currently not adequately protected (such as Southern Highland forests, Usambara and Uluguru Mts). Critical forest connections, such as the degraded Bujingijila corridor linking Mt Rungwe to Livingstone Forest in Kitulo, must be adequately 287

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protected.The ca. 25 km2 of moist montane forest in the Rubeho Mts, where a new population of Abbott’s Duikers was discovered in 2006, does not benefit from any legal protection, and therefore measures to give protected status to this forest should be urgently considered. Measurements Cephalophus spadix HB ("): 1240 mm, n = 1 T ("): 110 mm, n = 1 HF (slot) length ("): 50 mm, n = 1 E ("): 1190 mm, n = 1 Mt Rungwe, Tanzania (T. Davenport & S. Machaga unpubl.) HB ("): 1330 mm, n = 1

T ("): 135 mm, n = 1 E ("): 112 mm, n = 1 Sh ht. ("): 710 mm, n = 1 WT: 58.0 kg, n = 1 Mt Kilimanjaro, Tanzania (Wilson 2001) Kingdon (1982) gave the following measurements: HB: 970–1400 mm; T: 80–130 mm; Sh. ht: 660–740 mm; WT: 50.0–60.0 kg Maximum recorded horn length is 11.1 cm for a pair of horns from the Usambara Mts, Tanzania (Rowland Ward) Key References Wilson 2001.

Machaga & Davenport 2004; Moyer 2003;

Francesco Rovero, Tim R. B. Davenport & Trevor Jones

Cephalophus silvicultor YELLOW-BACKED DUIKER Fr. Céphalophe à dos jaune; Ger. Riesenducker Cephalophus silvicultor (Afzelius, 1815). Nova Acta Reg. Soc. Sci. Upsala 7: 265, pl. 8, fig. 1. ‘Habitat in montibus Sierrae Leone & regionibus susuensium fluvios Pongas & Quia adjacentibus frequens’; since restricted to Sierra Leone, vicinity of Freetown (Grubb et al. 1998).

Yellow-backed Duiker Cephalophus silvicultor adult female myology and skeleton.

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Yellow-backed Duiker Cephalophus silvicultor.

Taxonomy The type species of the genus, this species was described at a surprisingly early date for a tropical forest duiker. The reason was that the botanist Adam Afzelius, a pupil of Linnaeus, provided an illustrated account to the Swedish Royal Scientific Society on his return to Sweden after serving in Sierra Leone between 1792 and 1794. Ansell (1972) recognized three subspecies (the nominate form, C. s. ituriensis and C. s. ruficrista), but noted that their validity was in some doubt. St Leger (1936) had earlier only considered ituriensis as distinct, though Hill & Carter (1941) and Ansell (1960b) both recognized ruficrista (and which they considered to include the form coxi, which St Leger included in the nominate form). Haltenorth (1963) recognized ruficrista, but not ituriensis, and included coxi in the nominate form. More recently, a comprehensive review by Grubb & Groves (2001) recognized four subspecies, including the description of a new subspecies. The correct spelling of the species name is silvicultor, not sylvicultor, which is an incorrect subsequent spelling (Grubb 2004). Synonyms: coxi, curticeps, ituriensis, longiceps, melanoprymnus, punctulatus, ruficrista, sclateri, silvicultrix, sylvicultor, thomasi. Chromosome number: 2n = 60; the X chromosomes are submetacentric, while the Y chromosome is assumed to be a small acrocentric (Hard 1969, Robinson et al. 1996b). Description The largest of the duikers, dark brown, with a vivid cream-coloured patch on the back. The long wedge-shaped head has a grey muzzle and cheeks ending in a shiny black rhinarium. Pale offwhite colouring on the throat and lips is very variable in extent but tends to blend into the darker body colour without sharp demarcation. Eyes and ears are relatively small. Facial vibrissae are not obvious, nor do they grow out of light-coloured patches. Preorbital glands are large and prominent and can measure 44 mm with as many as 15 large pores (Wilson 2001). A coronal tuft of red or maroon hair, of variable length and conspicuousness, sticks up between the horns. Neck covered in short hair, variable in extent of pale throat

colouring. The skin in this area forms a thickened ‘neck shield’. The yellow dorsal triangle is variable in its extent, tonality and colour. Although there are no obvious glands beneath this triangle, the entire coat, which is sleek but softly haired, is slightly oily, presumably due to secretions by sebaceous and apocrine glands. The triangle begins just behind the shoulder blades and widens, ending on the croup. Behind this long-haired triangle is a short-haired light coloured area, which Grubb & Groves (2001) call a ‘haunch spot’, and which varies, both individually and regionally, in its colour (yellow to pale grey or white) and extent. Body and legs are very dark brown, almost black in some individuals. Tail very short and thinly haired except for a whispy terminal tuft. Sexes of similar size and colour, but "" tend to have thicker horns and larger preorbital glands. Pedal glands are present on both fore- and hindfeet. Inguinal glands absent. Horns slightly arched, with no or very small inner ridges, and possessed by both sexes, measuring up to 21 cm. Relative to Jentink’s Duiker C. jentinki and the Bay Duiker C. dorsalis, the skull has a more convex frontal region; the supraorbital foramina are not sunk into channels, and the lateral nasal processes are usually conspicuous. The rostrum is narrower, but shorter than in Jentink’s Duiker. Median pillars are commonly developed on the buccal sides of the cheekteeth (Grubb & Groves 2001). Geographic Variation C. s. silvicultor (including silvicultrix, punctulatus, sclateri): formerly from Senegal and Gambia to Niger R. Very large; brown with greyish tones, and with broad yellow dorsal triangle; no ‘haunch spot’. C. s. longiceps (including melanoprymnus, thomasi, ituriensis): from Niger R. to Congo R. and eastwards to extreme S Sudan and Western Rift Valley. Somewhat smaller than C. s. silvicultor, and darker to blackish-brown, with narrower dorsal triangle; ‘haunch spot’ absent, rare or weakly developed. 289

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Dorsal and lateral views of skull of Yellow-backed Duiker Cephalophus silvicultor.

C. s. curticeps: predominantly montane form between Western and Eastern Rift valleys, including in E DR Congo, Rwanda, Burundi, Uganda and SW Kenya. Smallest of the subspecies, very dark brown coat (similar to C. s. longiceps), dark golden-ochre dorsal triangle; ‘haunch spot’ present, usually well developed. C. s. ruficrista (including coxi): extensive area south of lower Congo R. to N Angola, and Zambia; some intergradation with C. s. longiceps. Similar in size to C. s. longiceps, but colour lighter (light to dark brown) and dorsal triangle broader; ‘haunch spot’ always present. Similar Species Cephalophus spadix. A large allopatric duiker that resembles this species in colour and conformation but lacks a dorsal triangle. C. jentinki. Sympatric in Sierra Leone, Liberia and SW Côte d’Ivoire. A large, grey-bodied duiker of similar size but much stockier build, with black head and neck and white shoulder halter. C. niger. A smaller, partially sympatric black duiker that lacks a dorsal triangle. C. dorsalis. Broadly sympatric. A squat red duiker with short broad head; smaller than C. silvicultor. Distribution Endemic to Africa, theYellow-backed Duiker has the widest distribution of the forest duikers (Ansell 1972, Wilson 2001). Originally, the species ranged from SW Senegal through all West African countries to extreme S Chad, the Congo Basin and east to SW Sudan and SW Uganda, from the northernmost extensions of gallery and riverine forests (almost up to the Sahel) to equivalent forest extensions in N Angola (including Cabinda) and Zambia, east of the Zambezi R. and north of the Muchinga Escarpment (East 1999,Wilson 2001). It is rare in primary forest of DR Congo, but is present in 800– 1000 m montane forest in the Bélinga mountains, NE Gabon (G. Dubost pers. comm.). Its equivalent habitats east of the Gregorian Rift are occupied by its close relative Abbott’s Duiker C. spadix. It has also been recorded from montane forest and bamboo in the Mau Forest in SW Kenya (Kingdon 1982, Hillman et al. 1988, East 1999), and is reportedly present on Mt Elgon (Kingdon 1982), although no museum specimens are available to corroborate this (Grubb et al. 2003). Within their range Yellow-backed Duikers are still common in some localities, but in other parts they have undergone significant

Cephalophus silvicultor

declines or are now absent.They are now considered extinct in Gambia (East 1999), although whether they actually ever formerly occurred is uncertain (Grubb et al. 1998, 2003). Malbrant & Quijoux (1958) recorded Yellow-backed Duikers from S Chad, but there is no recent information on their occurrence in this country. They also were thought to have been extirpated from Rwanda (East 1999), but have been confirmed as surviving in Nyungwe N. P. (F. Mulindahabi & A. Vedder pers. comm.). In Uganda, East (1999) recorded them surviving with certainty only in Bwindi Impenetrable Forest N. P. Habitat An animal of lowland and montane primary and secondary forest, riverine galleries and many permutations of forest–savanna mosaics, including secondary swidden ‘farmbush’, the borders of plantations and, in montane areas, bamboo groves and steep forested valley slopes. It is typically an ecotonal species and survives well in narrow riverine strips and fragmented woods. Often found close to deep swamps, perhaps seasonally. During very dry periods small aggregations have been reported around salt-licks. Throughout its range it lives only in strips or pockets of suitable habitat, where it can be quite numerous. Yellow-backed Duikers in densely forested NE Gabon exhibited strong associations with riparian habitat, and to a lesser extent with old secondary forest and advanced fallowing fields (Lahm 1993). In Taï N. P., Côte d’Ivoire, Newing (2001) observed this species only in secondary forest. Abundance The density of these large duikers varies enormously, partly due to habitat preferences and natural constraints that limit them to quite specific localities and partly to very high levels of predation by human hunters over the greater part of their range. In a particularly choice locality in forest–savanna mosaic in Lopé N. P., Gabon,Tutin et al. (1997) estimated a density of 2.1/km2. Elsewhere, in the National Park of Upper Niger, Guinea, Brugière et al. (2005) recorded individual and group densities of 0.79/km2 and 0.69/km2, respectively; Prins & Reitsma (1989) estimated 0.26/km2 in coastal

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Yellow-backed Duiker Cephalophus silvicultor.

forest/savanna of SW Gabon, while in the heavily poached Comoé N. P., Côte d’Ivoire, Lauginie (1975) estimated 0.09/km2. Lahm (1993, 1997) had encounter rates of 0.005/km and 0.03/km for this species in hunted and non-hunted areas, respectively, of densely forested NE Gabon, and 0.32/km in an unexploited logging concession now part of Lopé N. P. In the Ituri Forest, DR Congo, where these duikers are naturally rare (but even so opportunistically hunted on a regular basis), densities ranged from 0.1–0.7/km2 near settlements and 0.5– 1.6/km2 in remote areas (Hart, J. A. et al. 1996, Hart 2000). As the largest duiker, it consumes a greater proportion of widely dispersed large-sized fruits than smaller species (Dubost 1984). Comparing the relationship between the body weights and known home-ranges of some duiker species, Feer (1988) suggested that its home-range size may attain 200 ha. East (1999), assuming a density of 1.0/km2 where Yellow-backed Duikers were common and 0.1/km2 elsewhere, arrived at an overall population estimate of 160,000 animals. Adaptations Among the most striking peculiarities of theYellowbacked Duiker are its pale dorsal triangle, its considerable size (up to 70 kg), its ability to make an explosive and intimidating thumping noise, and its rather bovine appearance. Dubost (1979) estimated that its body is proportionally heavier than the diurnal duikers, but less than the strictly nocturnal species. Especially interesting is the fact that it has a black dorsal stripe at birth, as do young and adults of its closest relative, Abbott’s Duiker. Nocturnal duikers tend to be dark-coloured and it is possible that the common ancestral population of Abbott’s Duiker andYellow-backed Duiker was predominantly nocturnal. However, both species are essentially crepuscular, being most active both before and after dawn and dusk but intermittently active at other times of the day and night as well. When quietly browsing or resting back hairs lie flat and are not particularly conspicuous (even while fleeing).This is partly due to the freckled forest light and the existence of other light streaks, such as stems and trunks and partly because the surrounding dark pelage ‘encroaches’ on the dorsum when the coat is sleeked down. However, the dorsal hairs are elongated, erectile and in this condition are able to catch the light and make the back the single most arresting feature of the animal. Evolution of the near-white dorsal triangle is clearly linked to sending a signal to other forest dwellers, the most susceptible being members of the same species. What sort of information could such a vivid pattern have in a mainly solitary species?

Dorsal hair is raised whenever there is a disturbance that has put an animal on the alert. This is often described as ‘alarm’ behaviour, but the observations of J. Kingdon (pers. obs.) and those of Wilson (2001) suggest that an animal on the alert is generally static and closely monitoring its surroundings. It may even approach to investigate. It is during this phase of active interest that the back is hunched in readiness to run and the hair is at its most erect and conspicuous; S. Lahm (pers. obs.) observed this behaviour during both day and night surveys in Gabon. There are four likely main sources of disturbance: the first, and probably the most frequent, are neutral agents such as falling branches or monkeys’ noisy leaps. The second is any animal or event with a potential for harm, such as a hunter or Leopard Panthera pardus.The third is anotherYellow-backed Duiker. The fourth is any potential competitor, such as duikers of other species, Red River Hogs Potamochoerus porcus or Mandrills Mandrillus sphinx. Species-specific signals of this sort are only likely to have selective value when received by animals belonging to the last two categories (and a conspicuous back can easily address either). Because an alert animal typically bunches its back, the yellow patch effectively enlarges and advertises the highest part of a big black body mass (thereby emphasizing the signaller’s height and bulk). This can simultaneously identify and intimidate, particularly if accompanied by the ‘humphing’ sound peculiar to Yellow-backed Duiker. Competing duikers and some smaller carnivores can immediately see that they are dwarfed by aYellow-backed Duiker, while conspecifics have the choice of approaching or fleeing.There is, as yet, no evidence that the dorsal triangle overlies a glandular area but the entire animal is faintly greasy and the hairs may well serve as dispensers for a generalized body odour with all the information that entails. Overall the most likely function of this pattern is to invite retreat from an animal that has been selected to advertise its height and bulk and back this with an alarming ‘thump’. Dubost (1980) suggested that this species uses fewer visual signals than strictly diurnal duikers, but more than nocturnal species. Morphology and genes unite in suggesting that the Yellow-backed Duiker is an end-of-the-line species, one of the most recently evolved of duikers. That conclusion is also consistent with the idea that all duikers have derived from a diminutive ancestor and that the living radiation of duikers represents a protracted adaptation to the very varied opportunities offered by fruit-fall in the diverse forest habitats of Africa (Kingdon 1982). Where duikers belong within the larger radiation of bovids is a contentious topic that has a bearing on the strikingly ‘bovine’ appearance of this large duiker. Duikers have been seen as occupying an ambiguous position in the primary divide between bovines and antilopines. Most authors have placed them within the latter (as in this volume), but Gentry (1992), Groves & Schaller (2000) and Grubb & Groves (2001) noted the resemblance between duiker skulls and those of the Asian Four-horned Antelope Tetracerus quadricornis and other Asian boselaphines. They suggested that duikers might be as close to the bovines as to any other antilopine group and therefore proposed recognition of a new subfamily Cephalophinae to reflect a supposed intermediate position and relative distinctness from both Bovinae and Antilopinae. As cattle and buffalo have evolved into ever larger forms they have acquired that complex of characteristics and appearance that we sum up in the vernacular term ‘bovine’. It is therefore interesting that the 291

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Yellow-backed Duiker Cephalophus silvicultor.

evolutionary enlargement of this duiker species seems to parallel that transformation in the Bovinae and that this has led some scientists to doubt the duikers’ affinity with other antelopes. Foraging and Food Fallen seeds, fruits, berries and the bark of shrubs, fungi, ground moss and many herbs. In montane areas, waterberry Syzygium, dog plum Ekebergia and yellow-wood Podocarpus are favoured fruits (Kingdon 1982). Wilson (2001) emphasized that this species shows a marked preference in lowland forests for dry fruits, pods and seeds, which accounted for 79% of the diet, leaves, shoots and traces of animal matter making up the balance of his central African sample (three stomachs). Wilson (2001) recorded them having ingested large pieces of Piliostigma thonningii pods (and leaves), Gardenia sp. fruits and Swartzia madagascariensis. Gautier-Hion et al. (1980) recorded 71.3% fruit, 28.6% leaves and 0.1% animal matter in four stomachs in Gabon. Dubost (1984) found ten fruit species and two Eremospatha spp. as preferred foods in four stomachs. Feer (1995) also found them feeding on mature fruits, noting Irvingia spp. and Detarium macrocarpum. He recorded 30 species in 23 stomachs with large quantities of pulp, and recorded duikers swallowing fruits of up to 4.7 cm diameter. Small seeds passed through the digestive system unharmed, but several fruits were chewed so thoroughly that their seeds were actually destroyed: these were Chrysophyllum spp., Mammea africana, Pseudospondias longifolia and Ricinodendron heudelotii. The most important species by order of decreasing frequency were: Panda oleosa, Polyalthia suaveolens, Irvingia spp. and Gambeya beguei (F. Feer pers. obs.). In the south of its range, forest/savanna ecotonal species such as Pseudolachnostylis dekindtii, Oncoba spinosa, Swartzia madagascariensis and Canthium venosum are recorded foods (Ansell 1960b); other records are of African Mangosteen Garcinia and Duiker-tree Sapium fruits. It was mentioned as a crop raider of maize shoots and cassava leaves in five of 218 villages sampled in Gabon (Lahm 1994). Yellow-backed Duikers may also take carrion (Dekeyser & Villiers 1955), and one captive duiker was observed to capture, kill and eat pigeons (Kranz & Lumpkin 1982).Wilson (2001) recorded the remains of a lizard from one stomach, those of a chameleon from another in Congo, and reported that they would readily eat newly hatched tortoises.This species has also been observed feeding on a Forest Buffalo Syncerus caffer nanus carcass (L.White pers. comm.) and fresh Leopard Panthera pardus scat (P. Henschel, pers. comm.) in Gabon, and a Whitenosed Guenon Cercopithecus nictitans carcass in Congo (N. Shah pers. comm.). Mbuti pygmies in DR Congo described its diet as comprising fruits, leaves, insects and dead animals (Carpaneto & Germi 1989).

Of 23Yellow-backed Duiker stomachs, four contained meat, bones and hair, and two contained the remains of pangolins (F. Feer pers. comm.). Being both nocturnal and diurnal, there is no clear period of activity (Gautier-Hion et al. 1980, Newing 2001, Wilson 2001). It is physically adapted for rapid flight, unlike strictly nocturnal species such as the Bay Duiker and Water Chevrotain Hyemoschus aquaticus, which are relatively slow (Dubost 1979). The greater proportion of more diffuse large-sized fruits in its diet (compared with other duikers) may necessitate a 24-hour activity rhythm in a large homerange (Feer 1989b). Hart (2000) inferred that it is mainly nocturnal near human settlements. This may be a behavioural adjustment to avoid contact, as suggested by Lahm (1993) for this species and other large ungulates in hunted areas of Gabon. Social and Reproductive Behaviour Although this appears to be a mainly solitary species and most records have been of single animals, Wilson (2001) found what he called ‘bonded pairs’ living in discrete home-ranges centred along a riverine forest strip in Mole N. P. in Ghana.The gallery was about 3.5 km long, but a 600 m dry break separated the two home-ranges. Both sexes mark their home-ranges with large preorbital glands but "" mark three times more often (Wilson 2001). Lying-up on termitaries or slight elevations on the forest floor suggests that they regularly monitor their surroundings and broken horns on female skulls suggests that !! may be actively intolerant of other !!. Individuals lie up singly (very occasionally in pairs) between the buttresses of large trees, under fallen trunks or dense tangles, in ‘forms’, which show signs of regular use and where the duiker may shelter from rain (even entering the remnants of pitsawyers’ or hunters’ lean-tos). As many as six such resting places can be found within a square kilometre. Given the absence of marked ridges on the inner side of the horns, this species probably does not scrape the bark of small trees, as do some other duiker species (G. Dubost pers. comm.). Courtship resembles that of other duikers: the " tests the condition of the ! with flehmen and, if she is in oestrus, pursues her relentlessly, licking her but also butting and biting, and striking her with a foreleg. Meanwhile the ! grunts or bleats (Wilson 2001). Reproduction and Population Structure Ansell (1960a) thought there was no fixed breeding season in Zambia, but reported a lactating ! in Mar and young in Oct and Dec. Ionides (1964) noted young in Kenya during Jan, and J. A. Hart (pers. comm.) recorded very young lambs in Ituri Forest in Jan and May. In Gabon, where births occur year-round in duikers, bats, rodents and other forest antelope, there are two peaks that differ among species (Brosset 1986, Feer 1988). The frugivorous duikers and Water Chevrotain generally had more births during the two annual dry seasons compared with rainy months in NE Gabon (Dubost & Feer 1992). S. Lahm (pers. obs.) saw small lambs in Jan (dry month) and Oct (wet month) in the same region and in Jun (wet–dry transition) in SW Gabon. Wilson (2001) found that even when ovulation originates from the left ovary, the foetus implants in the right horn of the uterus. There has been much discussion about the length of gestation. Most authenticated captive records are between 210 and 282 days, but extremes of 151–310 days have been published (see Wilson 2001, and references therein). A single young is born. Captive birthweights range between 2.3 and 6.1 kg (Wilson 2001); the young is born a dark umber colour with strong reddish tint, particularly on

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Yellow-backed Duiker Cephalophus silvicultor.

the underparts and freckled flanks.The dorsum is jet black but begins to pale at 40 days, visibly whitening at five months and fully developed by seven months (Schürer 2002). It begins to eat plant material at about two months and is weaned by five months (Wilson 2001). In captivity, birth intervals range from 213 days to three years (Kranz & Lumpkin 1982). A captive ! at Miami Metro Zoo produced several young, the last of which was born when the ! was 18 years old. The " of the pair was still alive in 2005 at age 23, the oldest knownYellow-backed Duiker in captivity (L. Rohr pers. comm.). Predators, Parasites and Diseases Known to be taken by Leopards and pythons Python spp. and young are probably taken by Servals Leptailurus serval,African Golden Cats Profelis aurata and Crowned Eagles Stephanoaetus coronatus. In N Congo, a considerable number ofYellow-backed Duikers and other bovids succumbed to an epidemic of a pneumonia-like disease possibly transmitted by biting Stomoxys flies (Stockenstroom et al. 1997; and see Elkan et al. 2009). A health evaluation of 77 duikers representing five species in the Ituri Forest revealed the presence of strongyles, trichurids, coccidia, Moniezia sp. and occasionally ticks (Rhicephalus spp., Haemaphysalis parmata, Ixodes cumulatimpunctatus). Many animals had positive antibody titres to leptospirosis serovars, bluetongue virus, infectious bovine rhinotracheitis and epizootic haemorrhagic disease (Karesh et al. 1995). The sample did not include any Yellow-backed Duikers, although the species is part of the local duiker community, and thus must be exposed to the same parasites and diseases. Dekeyser (1955) recorded the tick Ixodes rasus. Conservation IUCN Category: Least Concern. CITES: Appendix II. This species was formerly subject to strict taboos that once protected it in some parts of its range (Kingdon 1982). It is still considered non-preferred game in parts of central Africa, owing to its dark colour, scavenging habits and aggressive temperament when hunted (Carpaneto & Germi 1989, Lahm 1993). In Gabon, where it is also believed to be a ‘were-animal’, 17% of 144 villagers

sampled declared it to be non-edible (Lahm 2002). However, the Yellow-backed Duiker is now declining over most of its range due to intensive and unregulated hunting and the disappearance of taboos. It has disappeared from Gambia and is rare in Senegal, Guinea, Sierra Leone, Nigeria, Kenya and Uganda and over much of the peripheries of its range (Anadu & Green 1990, Anstey 1991, Wilson 2001). Important protected areas include Ziama and Diécké Forest Reserves (Guinea), Gola Forests (Sierra Leone), Sapo N. P. (Liberia), Mbam Djerem N. P. and Lobéké N. P. (Cameroon), many of those in Gabon, including Odzala N. P. and Nouabalé-Ndoki N. P. (Congo), Monte Alén N. P. (Equatorial Guinea), Bwindi Impenetrable N. P. and Queen Elizabeth N. P. (Uganda), Kafue N. P. and Kasanka N. P. (Zambia), and Okapi Faunal Reserve (DR Congo). Comoé N. P. was once a stronghold for this species, but with 100–500 poachers operating on a daily basis in this park it had already become rare by the mid-1990s (Fischer 1997). Wilson (2001) regarded Yellowbacked Duikers as extinct in at least seven protected areas in which they formerly occurred, although subsequent surveys have confirmed that they do still occur in Digya N. P., and that they are present in Bui N. P. and Gbele Resource Reserve (R. J. Dowsett pers. comm.). Although the species was described as easy to breed and hand-rear in captivity (Kranz & Lumpkin 1982), there were only 65 animals in about 23 North American zoological parks and eight known in international institutions in 2005. Within these captive groups a birth rate of 10 per year has been stable, and improved husbandry and population management practices in recent years have increased the Yellow-backed Duiker’s survival potential in captivity (L. Rohr pers. comm.). Measurements Cephalophus silvicultor TL (""): 1420, 1460 mm, n = 2 TL (!): 1620 mm, n = 1 T (""): 120, 125 mm, n = 2 T (!): 115 mm, n = 1 HF c.u. (""): 295, 330 mm, n = 2 HF c.u. (!): 350 mm, n = 1 E (""): 116, 121 mm, n = 2 E (!): 115 mm, n = 1 Sh. ht (""): 748, 775 mm, n = 2 Sh. ht (!): 730 mm, n = 1 WT (""): 66.5, 68.6 kg, n = 2 WT (!): 71.0 kg, n = 1 SE Central African Republic (Wilson 2001) A ! from NE Gabon was subadult, yet weighed 74.5 kg (G. Dubost pers. obs.), so maximum weights (and other measures) probably exceed those shown here. It should be remembered that regional populations differ in body size, although the extent of this variation remains to be quantified. Maximum recorded horn length is 21.2 cm for a pair of horns from Tchibota, DR Congo (Rowland Ward) Key References Dubost 1979, 1984; Dubost & Feer 1992; Feer 1988, 1989b, 1995; Kingdon 1982; Kranz & Lumpkin 1982; Wilson 2001. Jonathan Kingdon & Sally A. Lahm 293

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Cephalophus dorsalis BAY DUIKER Fr. Céphalophe bai; Ger. Schwartzruckenducker. Cephalophus dorsalis Gray, 1846. Ann. Mag. Nat. Hist., ser. 1, 18: 165. ‘Sierra Leone’.

Bay Duiker Cephalophus dorsalis. Lateral view of skull of Bay Duiker Cephalophus dorsalis. LEFT:

ABOVE:

Taxonomy Named from a menagerie specimen from Sierra Leone; Gray named another specimen C. badius, which is now regarded as a synonym. Rode (1943) mistakenly lumped leucogaster and ogilbyi as subspecies of C. dorsalis. St Leger (1936) separated out dorsalis, based on horn characters. Cephalophus dorsalis is sympatric with both Brooke’s Duiker C. ogilbyi brookei and the White-bellied Duiker C. leucogaster. Ansell (1972) included the form arrhenii as a subspecies, but this form is actually attributable to the Whitebellied Duiker (Grubb & Groves 2001). Synonyms: badius, breviceps, castaneus, kuha, leucochilus, orientalis, typicus. Chromosome number: 2n = 60; the X chromosome is a metacentric (Hard 1969, Robinson et al. 1996b). Description Heavily built duiker with a red or yellowish-brown glossy coat, black or dark brown legs and a black dorsal mid-line. Also a dark brown mid-line down the chest and belly. Muzzle is reduced and strongly tapered. Eyes are larger, more prominent and higher in the skull and the entire head is broader and flatter than in any other duiker. The black dorsal line, which varies greatly in length and width and darkness, usually extends up the neck, over the crest and down the nose, especially in western populations, where much of the head can be black; however, the extent of black on face and back is very variable. Frontal crest reduced to some long hair at the base of horns. In common with some other duiker species, there are vibrissae on the muzzle and above the eye. These have black bases set in distinctive white flashes on the upper lip and upper eyelid. Chin and throat also white. Backs of the pinnae are black or brown; their fronts have white margins and some very sparse off-white hair inside. Hair on the shoulders is somewhat shorter than on the back and hindquarters. Legs black or brown, but sometimes show rufous streaks. Tail black with a white underside and somewhat tufted towards the tip. Deeply brown or black at birth,

rufous colouring appears between 5 and 6 months of age; the black dorsal stripe is only partly visible on the rump. Preorbital glands are relatively small compared with some other medium-sized duikers (Wilson 2001), although in Gabon the preorbital glands of "" were twice as large as in Peters’ Duiker C. callipygus. Preorbital glands have 11–14 pores set in a straight line about 21 mm long (Wilson 2001). The inguinal glands are very distinctive, comprising deep, flask-like sacs in the lower flanks that exit from a narrow tube immediately inside the fold between thigh and abdomen. Their odour is said to be very disagreeable to the human nose. Pedal glands are large and active. Sexes are of similar size, although !! are slightly heavier. Preorbital glands are larger in "" than in !!, as are the slender, cylindrical, parallel, spike-like horns. Geographic Variation C. d. dorsalis: Guinea, Guinea-Bissau and Sierra Leone to the Niger R. Grubb & Groves (2001) considered the boundary between the two forms to be the Adamawa Highlands in Cameroon. Redder body colour, black facial blaze and crest; dorsal stripe comparatively narrow. Smaller body size. C. d. castaneus: east of the Niger R. to the Rwenzori Mts and Albertine Rift Valley and south to NE Angola. Darker body colour, redder face and crest. Larger body size. Grubb & Groves (2001) remark that there is considerable geographical variation in this subspecies. Hornless adult !! have been observed east of the Congo R. Similar Species Cephalophus ogilbyi. Sympatric across most of the range. A diurnal red

duiker of similar size, and often confused with the Bay Duiker, but of paler red colour with a long narrow face and lighter-coloured legs.

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C. leucogaster. Sympatric in central Africa north of the Congo R. A more gracile, pale-cream-coloured duiker with prominent white belly and two-toned legs. C. callipygus and C. weynsi. Red duikers of similar size with long faces, prominently swollen foreheads and thick annulated horns. C. silvicultor. Broadly sympatric. A larger, dark greyish-brown duiker with cream-coloured patch on lower back. Distribution Endemic to Africa and closed-canopy areas within the rainforest block from Guinea-Bissau to the Rwenzori Mts, Albertine Rift Valley and L. Tanganyika, and south to NE Angola to about 11° S (Grubb et al. 1998, East 1999, Wilson 2001, CrawfordCabral & Veríssimo 2005). The westerly limit is Guinea-Bissau, where East (1999) reported Bay Duikers from forest patches in five restricted areas of the mainland, including Cacheu R. and L. Cufada. In Guinea, they are recorded from the south-west and south-east, including Mount Nimba Biosphere Reserve and Ziama and Diécké Forest Reserves, and then eastwards to Togo, where they were recorded in the forests of the Fazao and Togo mountains (East 1999). However, there are no confirmed records from Gambia (Grubb et al. 1998, 2003) or from Benin. There are also no records from Nigeria, west of the Cross R. (Grubb et al. 2003), their distribution continuing again east of the Cross R. in SE Nigeria, and then through S Cameroon, SW and SE Central African Republic, mainland Equatorial Guinea (being absent from Bioko I.), Gabon, Congo, east to the montane forests in E DR Congo. Fay et al. (1990) mentioned a doubtful record from Manovo–Gounda–St Floris N. P. in the north of Central African Republic, and another record from east of Bamingui–Bangoran N. P., but considered these as doubtful; however, Grubb et al. (2003) subsequently report on a specimen from N Central African Republic from Kabo, just south of the Chad border, and to the west of Gribingui-Bamingui Faunal Reserve.There is a single record from Uganda, below the north-western foothills of the Rwenzori Mts, but this area is now densely populated, and the

Bay Duiker probably no longer occurs in Uganda (Kingdon 1982, East 1999). Habitat An inhabitant of the entire equatorial lowland rainforest block with a marked preference for high primary rainforest (Wilson 2001); also, if undisturbed, may occupy patches of forest within savanna mosaics. Within the rainforest zone Bay Duikers may visit edges of clearings and well-diversified areas with both dry and seasonally waterlogged areas (Dubost 1979). They generally avoid montane forests, although they are found in moist lowland forests on the slopes of the montane regions in E DR Congo. Very slow to recolonize felled or disturbed forest, but does occur in old farmbush and old secondary forest (Fotso & Ngnegueu 1997, Wilson 2001). They seek out the most densely vegetated areas, piles of dead branches in forest gaps, the spaces between buttresses and large hollow logs to lie up in during the day, refuges that they freeze in and only leave when discovered (J. Kingdon pers. obs.).

Frontal view of Bay Duiker Cephalophus dorsalis (right) compared with Yellowbacked Duiker C. silvicultor to show former’s larger eyes and shorter muzzle.

Abundance Estimates of home-range data in Makokou, Gabon, suggested densities of 7.5–8.7/km2 in prime habitat (Feer 1988). In Makokou, Dubost (1979) calculated a density of about 19/km2 (net capture). Lahm (1993) estimated by transect a density of 5.8/ km2. In S Central African Republic, the species ranks second among duikers, with densities of 0.3–6.6/km2 (line transect and net count; Noss 1998b). Densities are 1.5/km² in the Ituri Forest (DR Congo) (Hart 1985) and 1.9/km2 in Taï N. P. based on night transect counts (Hoppe-Dominik et al. 2011). Using modest densities of 0.2–2.0/ km2, East (1999) calculated an overall population of 725,000 animals.

Cephalophus dorsalis

Adaptations Striking features of this species are the enlarged eyes and orbits, adaptations that are clearly related to improved night vision in an animal with very strictly nocturnal habits. Protrusion of the eyes on either side of a rather flat-topped head and dorsally oriented orbits are also likely to improve bifocal vision (Kingdon 1982, Feer 1988). However, the most significant peculiarity lies in the broad, flattened skull.When Bay Duiker skulls are compared with those of other duikers they show inflation of the zygomatic bones, outward bowing of the zygomatic arches, lateral expansion of the molar toothrow (without equivalent modification of the premolars), widening of the mandibular condyles, and significant narrowing of the choanae so that the pterygoid bones are closer together. Taken all together this combination of 295

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Palatal views of Bay Duiker C. dorsalis skull (above) and Black-fronted Duiker C. nigrifrons (below) to show broader molar teeth, greater mandibular traverse and enlarged masseter and pterygoid muscles.

Bay Duiker Cephalophus dorsalis skull (above) compared with Blue Duiker Philantomba monticola (below) to show proportions in relation to toothrow (not to same scale).

characters signifies a significant amplification of the lateral element in chewing by introducing a wide lateral swing at the maxillary condyles (Kingdon 1982). This lateral action means that the duiker can grate or rasp fruits without cutting them open with a sharp vertical bite: a dangerous risk when seeds are protected by poisonous compounds. This risk might have exerted significant selection on anatomy and physiology during this duiker’s evolution. The estimated mass of the masseter and zygomatic-mandibular muscle is large compared with duikers of similar size (such as C. callipygus).This, combined with broad upper and lower molars (Feer 1988), signifies more powerful jaws able to process hard fruits and seed. The maxilla has also contracted, and this shortening of the maxilla, combined with a wide, elastic mouth and lips, allows the gape to widen to the point where a 7 cm fruit can easily be taken into the mouth and a 5 cm fruit can be bitten open. Few other duikers can cope with fruits that have large hard seeds and most prefer easily managed ones with abundant soft pulp and small fruits/seeds such as figs. It has been calculated that fruits measuring less than 4 cm make up about 94.5% of the fruit-fall in a representative forest in Gabon (Dubost 1979), so small fruits represent the primary food source for most duikers. However, any animal that can process larger seeded fruits has fewer competitors (although the resource is more dispersed because there are fewer trees bearing such fruit). The Bay Duiker has specialized in stripping away pulps (that are often quite thin) from relatively large, hard (and sometimes poisonous) seeds.The majority of such seeds are high forest, closed-canopy species. Feer (1989b) and Wilson (2001) have documented at least 15 larger forest fruits that are preferred by Bay Duikers. These fruits range from 2 to 20 cm in size, but most are about 2–7 cm, dimensions that the Bay Duiker can easily cope with. The hard seeds within these fruits range from 1.5–5 cm and Bay Duikers swallow and regurgitate seeds that are up to 2.8 cm (Feer 1989b). Seeds over this size are not swallowed but are stripped within the

mouth and spat out. It is this ability to process relatively large seeds that distinguishes the Bay Duiker and explains its cranial anatomy. These animals can combine mitigating competition and avoiding predators by consuming large numbers of fruits quite rapidly at the fruit-fall and the Bay Duiker has been recorded as being active for as short a period as 5–6 h per night (Wilson 2001). Once a duiker has fed it removes itself to a secure, sheltered place to rest and ruminate; here it regurgitates and spits out undamaged seeds, thus helping disperse them away from the parent tree (Feer 1995). It would be interesting to investigate the possibility of co-evolution between the Bay Duiker and some of its food trees. Many forest tree seedlings need some light during their early growth spurt and the Bay Duiker does sometimes deposit seeds in tree-fall gaps; however, these duikers also spit out large numbers of seeds from closed-canopy, high forest species in very densely shaded (but possibly less predated) sites. Under such circumstances only very shade-tolerant seedlings have any chance of growing to maturity. It is therefore likely that the duikers’ regurgitation habits would only favour those seeds best able to survive under deep shade. If so, Bay Duikers might be actively selecting and propagating those food trees that contribute most to the maintenance of a closed canopy.Theoretically, and on an evolutionary time-scale, these habitatspecific duikers could be favouring precisely those tree species that combine food supply with the maintenance of habitat. If these duikers are exterminated it is possible that many major closed-canopy tree species could also decline and even die out. Another peculiarity of the Bay Duiker skull is an enlarged olfactory region and this can be correlated with observed ‘tracking’ behaviour, and more generally with a nocturnal mode of life.The more dispersed a food supply the greater the demands on finding it and olfaction has to be a major sense guiding this task. Captives have been seen to sniff out and excavate well-buried fruits (Wilson 2001) by scraping soil with front hooves or incisors (Dubost 1983).

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There are occasional records of Bay Duikers stalking and killing birds (they discard legs and wings) and eating embryos from aboutto-hatch eggs (Kurt 1963, Wilson 2001). They eat carrion, and the remains of African Porcupines Atherurus africanus (F. Feer pers. obs.) and cusimanses (Crossarchus spp.) (Hart 1985) have been found in stomach contents. Termites, beetles and ants have also been recorded in their diet. Live prey are stalked and struck with a slash of the forefoot.

Head of Bay Duiker Cephalophus dorsalis female indicating vibrissae embedded in white ‘signal’ patches above eye and behind rhinarium.

Because the Bay Duiker is nocturnal, has large, prominent eyes and lives most of its life in dense vegetation, its facial vibrissae play an important role in avoiding potential snags. The positioning of such vibrissae above the eyes and around the nostrils is therefore appropriate, but it is interesting that in both instances the vibrissae are advertised by being embedded in small patches of vivid white fur. As with many other exclusively nocturnal mammals it would seem that unique combinations of white flashes on an otherwise dark facial ‘mask’ serve as intra-specific signal devices. Foraging and Food The Bay Duiker has very precise preferences for fruit species and the characteristics of these fruits have been extensively studied by Feer (1988, 1989b, 1995) and by Wilson (2001). Of the larger, mango-like species, ‘Gaboon chocolate’ Irvingia spp. and the Tallow tree Detarium macrocarpum are favourites; another is the globular, latexy, fibrous fruit (7–10 cm) of Oboto Mammea africana. These large seeds are rasped for their pulp or ‘flesh’ and the seed spat out, not swallowed. The commonest seeds to be found in stomachs and at resting sites are from the yellow fruits of two Antrocaryon species, the indented, button-like seeds of which are appropriately known as ‘antelope buttons’. Also eaten are the oval yellow fruits of climbers Hugonia spp. and the bilobed 6×3 cm fruits of Ricinodendron heudelotii (a forest relative of the Manketti tree Schinziophyton rautanenii that produces mugongo nuts) and oil-palm dates Elaeis guineensis, and the orange plums of milkwood Chrysophyllum beguei (4–6 cm), the olive-like, turpentine-scented fruit of Pseudospondias longifolia (3.5 cm), the green, strongly fibrous plums of Panda oleosa (5–7 cm) and the cocoa-pod-like Cola rostrata, which has a shell about 8×20 cm, but contains many small pulpcovered seeds. Softer seeds without a hard shell or rind tend to be chewed up, and Feer (1989b) never found the seeds of such species at resting places but did record them being eaten. They included the sweet, yellow, apple-like Drypetes gossweileri (8–14 cm), the oil-seed Staudtia gabonensis (2–6 cm), blue ozigo olives Dacryoides buettneri (2–4 cm), Anguek Ongokea gore (2–4 cm), ebap, red beans of Santiria trimera, the African soursop Annonidium mannii (20–50 cm) and owala oil seeds (flat discs from the giant pods of Pentaclethra macrophylla).

Social and Reproductive Behaviour Usually solitary, but occasionally observed in pairs. Two !! and their offspring have been found to occupy the home-range of a single " (Feer 1989a), so as many as six or seven animals may live within a recognizable male territory or home-range of 79 ha. The existence of very pungent inguinal glands and exceptionally large pedal glands suggests that intra-specific contacts are probably regulated by scent. Males mark tree and shrubs with preorbital secretion (Dubost 1983). A lightly built skull and short, slender spike-like horns imply that fights do not involve head-clashing. Stabs into the haunches or sides of contestants are the most likely outcome of male competition. Nocturnal habits and intense wariness has hidden much of this duiker’s social behaviour from view. The " pursues an oestous ! with great persistence, meanwhile humming continuously and occasionally striking out with a foreleg (a rudimentary form of laufschlag) (Wilson 2001). On accepting to be mounted, the ! squats and holds her tail to the side. Reproduction and Population Structure A wide spread of birth and pregnancy records implies that some breeding goes on throughout the year and no birth peak is currently known to occur. In Gabon, birth peaks were observed during and before maximum availability of fruit (Dubost & Feer 1992). About 57% of !! over 20 months old are pregnant. Oestrus only lasts about 18 hours (Wilson 2001). Females may conceive at about 18 months and always implant in the right horn of the uterus although both ovaries ovulate; gestation is cited as about 238 days (Wilson 2001). At birth the young weighs 1600–1690 g, and begins to take solid food within a few weeks; it is weaned by 3.5 months and is unusual in eating fruit almost exclusively from the beginning (Wilson 2001). In Gabon, a sample of 151 individuals from a moderately hunted population was composed of 20% adults. Males accounted for 58% of adults, a sex ratio most probably biased in favour of "" (Feer 1988). Longevity in captivity has been given as 17–18 years (Weigl 2005). Predators, Parasites and Diseases Data from scats have confirmed that Leopards Panthera pardus are important predators (Hart, J. A. et al. 1996, Wilson 2001). Kudo & Mitani (1985) observed a Mandrill Mandrillus sphinx preying on a young Bay Duiker.Young animals are also known to be taken by Crowned Eagle Stephanoaetus coronatus (Wilson 2001), and African Golden Cats Profelis aurata are also likely to take young or subadults. The failure of an attempt to reintroduce Bay Duikers to Azagny N. P., Côte d’Ivoire, was blamed on large numbers of African Rock Pythons Python sebae in the park (Roth & Hoppe-Dominik 1990). In a study investigating the health status of Bay Duikers, bacterial and viral serology tests returned positive for bluetongue disease, epizootic haemorrhagic disease, infectious bovine rhinotracheitis and Leptospira (which causes leptospirosis) (Karesh et al. 1995). Helminths recorded 297

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include nematodes (genera Cooperia, Strongylus, Cecropithifilaria, Filaria, Setaria, Skrjabinodera, Pygarginema) and cestodes (Moniezia, Avitinella) (Round 1968). Infection with coccidia (Karesh et al. 1995) and trypanosomes (e.g. Trypanosoma brucei) (Njiokou et al. 2004b) has been recorded. Ntiamoa-Baidu et al. (2005) recorded the following ixodid tick species from Bay Duikers in Ghana: Haemaphysalis parmata, H. leachi, Ixodes muniensis, I. moreli, I. cumulatimpunctatus and Rhipicephalus ziemanni. Beaucournu & Bain (1982) recorded a new species of flea (Ctenocephalides chabaudi) from the Bay Duiker in Gabon. Conservation IUCN Category: Least Concern. CITES: Appendix II. The combination of systematic habitat destruction and the commercialization of hunting and trapping likely will ensure the eventual elimination of this species outside protected areas. Even then they are unlikely to survive in smaller protected areas because of habitat degradation and poaching: in Ghana, for example, this species has been extirpated from seven parks and protected areas (Wilson 2001; although it does still occur in Kalakpa Resource Reserve; R. J. Dowsett pers. comm.), and Van Vliet et al. (2007) recorded its likely extirpation from Ipassa Natural Reserve in NE Gabon. Important populations survive in areas such as Sapo N. P. (Liberia),Taï N. P. (Côte d’Ivoire), Bia and Kakum National Parks (Ghana), Campo-Ma’an N. P., Lobéké N. P. and DjaWildlife Reserve (Cameroon), Dzanga–Sangha Special Reserve and Bangassou F. R. (Central African Republic), Monte Alén N. P. (Equatorial Guinea), Lopé, Ivindo, Minkébé, Loango and Moukalaba-Doudou National Parks (Gabon), Odzala N. P., Nouabalé– Ndoki N. P. and Léfini Faunal Reserve (Congo) and Kahuzi–Biega, Maiko and Salonga National Parks (DR Congo). This is a very popular quarry for hunters because it is compact, muscular and good-eating. It is hunted across its entire range, together with all other forest fauna, year-round and at increasingly intense, unregulated levels. In the Central African Republic, Bay Duikers accounted for 6.8% of all animals captured with nets (Noss 1998b). In Gabon, they accounted for 12% of duikers caught by village hunters and trappers (Lahm 1993). In the Dja area (Cameroon) they represent 17–25% of duikers killed (Fotso & Ngnegueu 1997, Muchaal & Ngandjui 1999). In Congo, Bay Duikers represent 15% of duikers for sale in Brazzaville bushmeat markets (F. Feer pers. obs.) and only 3.7% in Pointe Noire (Wilson & Wilson 1991). They rank between sixth and ninth among the game species on offer in Libreville (Gabon) markets (Ntsame Effa 2005), and were among the top five most hunted species in a bushmeat study in Nigeria (Fa et al. 2006). Studies in Equatorial Guinea, W Cameroon and Central African Republic have shown that current harvest rates are entirely unsustainable (Fa et al. 1995, Noss 1998b, Muchaal & Ngandjui 1999); in the Monte Mitra region of mainland Equatorial Guinea, Fa &

Garcia Yuste (2001) studied the offtake patterns of 42 hunters over a period of 16 months. The principal hunting method employed was cable snaring, and the Bay Duiker was by far the most heavily exploited species. As mentioned earlier, the extermination of this species could have much wider repercussions in the long-term ecology of those tree species (several of them highly desired commercial timbers) that are dispersed by Bay Duikers. Measurements Cephalophus dorsalis TL (""): 1040 (1030–1074) mm, n = 5 TL (!!): 1031 (1011–1049) mm, n = 4 T (""): 97 (89–107) mm, n = 5 T (!!): 100 (91–110) mm, n = 4 HF c.u. (""): 226 (201–232) mm, n = 5 HF c.u. (!!): 219 (202–231) mm, n = 4 E (""): 75 (75–76) mm, n = 5 E (!!): 76 (74–77) mm, n = 4 Sh. ht (""): 472 (448–486) mm, n = 5 Sh. ht (!!): 479 (454–489) mm, n = 4 WT (""): 19.9 (18.1–23.0) kg, n = 5 WT (!!): 20.6 (19.1–22.6) kg, n = 4 Liberia (Wilson 2001) TL (""): 1041 (1019–1064) mm, n = 8 TL (!!): 1044 (1021–1073) mm, n = 7 T (""): 100 (87–116) mm, n = 8 T (!!): 99 (84–112) mm, n = 7 Sh. ht (""): 480 (460–497) mm, n = 8 Sh. ht (!!): 478 (453–491) mm, n = 7 WT (""): 21.2 (17.9–22.7) kg, n = 8 WT (!!): 20.9 (18.3–22.6) kg, n = 7 Ghana (Wilson 2001) HB (""): 948 (880–1000) mm, n = 8 HB (!!): 963 (880–1050) mm, n = 13 WT (""): 19.0 (16.2–23.0) kg, n = 6 WT (!!): 22.2 (20.0–24.5) kg, n = 5 Gabon (Feer 1988, F. Feer pers. obs.) Maximum recorded horn length is 12.3 cm for a pair of horns from Yakadouma, Cameroon (Rowland Ward) Key References Dubost 1979, 1983; East 1999; Feer 1988, 1989a, b, 1995; Kingdon 1982; Wilson 2001. Jonathan Kingdon & Francois Feer

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Cephalophus jentinki JENTINK’S DUIKER Fr. Céphalophe de Jentink; Ger. Jentinkducker Cephalophus jentinki Thomas, 1892. Proc. Zool. Soc. Lond. 1892: 417. ‘Liberia’; since identified as Junk R. opposite Schieffelinsville, Sharp-Hill (Kühn 1965).

Jentink’s Duiker Cephalophus jentinki. Lateral view of skull of Jentink’s Duiker Cephalophus jentinki. LEFT:

ABOVE:

Taxonomy Monotypic. Rode (1943) mistakenly suggested this species was conspecific with theYellow-backed Duiker C. silvicultor. A multiple analysis of mtDNA sequences was consistent in always showing that C. jentinki was most closely allied to the Bay Duiker C. dorsalis and, more distantly, to Abbott’s Duiker C. spadix and the Yellow-backed Duiker (Jansen van Vuuren & Robinson 2001). Synonyms: none. Chromosome number: 2n = 60; the X chromosome is a submetacentric (Hard 1969, Robinson et al. 1996b).

is longer and narrower; orbit larger; rostrum relatively longer but bowed anteriorly; forehead almost flat with grooves in which the supraorbital foramina are located; frontonasal suture more pointed; nasal and mandibular profiles more convex; mesopyerygoid fossa much narrower; bullae less angular; zygomatic arches more suddenly contracted in ear region; and no or rudimentary lateral nasal flanges.

Description A very robust, short-legged duiker with a bold pattern of black, white and grey. Very darkly coloured head and neck offset by a vivid white halter over the shoulders and lower chest and a white border surrounding lips, muzzle and nose. This colouring involves both skin and fur, the latter being extremely short and fine. In contrast to the fore-end, the hindquarters are grey agouti. There is an inconspicuous dark crest on the forehead.Young Jentink’s Duikers are almost completely brown, similar to young Yellowbacked Duikers, and apparently have long lateral hooves. The tail is short. Preorbital glands are very large. Inguinal glands also are large, and often contain quantities of soft secretion (Wilson 2001). Pedal glands are present. The local inhabitants have many different names for them, and they are known as ‘forest goat’, ‘white woman’, ‘white antelope’ or as the duiker ‘onto which the dew has fallen’. Both sexes bear long (up to 21 cm) black horns, which are straight and smooth, oval in cross-section, extending backwards from the head in the plane of the face and curve very slightly downward towards the tips. The frontal outline of the skull is flat and the facial region above the toothrow and below the anteorbital fossa is markedly swollen out laterally, so that the teeth and their alveoli are quite hidden in an upper view of the skull. Grubb & Groves (2001) list a number of features that differentiate the skull of this species from the similar-sized Yellow-backed Duiker, including: the cranium

Similar Species Cephalophus silvicultor. Sympatric and similar in size, but dark greyishbrown with a vivid cream-coloured patch on the back; horns less depressed.

Geographic Variation

None recorded.

Distribution Endemic to West Africa, being found only in the western part of the Upper Guinean forest block, from Sierra Leone through Liberia to W Côte d’Ivoire. In Sierra Leone, Jentink’s Duikers were only positively reported from the country in 1989, from Western Area F. R. (southern part of the Freetown Peninsula). There is photographic evidence of their presence in the Gola F. R. complex (Ganas & Lindsell 2010), and their presence has been reported in areas such as the Mokanji Hills, Loma Mts and Tingi Hills (Davies & Birkenhäger 1990, Wilson & Wilson 1990, Grubb et al. 1998). Kranz & Glumac (1983) indicated that the species occurred widely in E Liberia. It was found to be thinly distributed along the Sehnkwehn R., possibly present along the Sinoe R. and common at the Buto Oil Palm Plantation. Probably occurs locally throughout Liberia in areas of suitable habitat. The 1989/90 WWF/FAD survey confirmed its presence in the south-eastern forests including Grebo N. P. and surrounding forests. Also present in north-western forest blocks as well as in North Lorma National Forests. Presence in Sapo N. P. was confirmed in 1997 (East 1999). During a survey of hunters’ 299

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Wilson 2001), and is even known to visit the sea-shore, presumably for salt. It is a ‘hider’ choosing hollow trees, fallen trunks and the buttress bays of kapok (Ceiba), Bombax and mututu trees (Klainedoxa) for shelter. It is so secretive that it continued to survive less than 30 km from Freetown, a city of half a million people, hiding on steep, densely forested slopes in the city’s water catchment area (Davies & Birkenhäger 1990). Its most basic requirements appear to be a diversity of fruiting trees and very dense shelter rather than a specific forest type (Kingdon 1997).

Cephalophus jentinki

quarry in the area north of Sapo N. P. and along the logging road through the Krahn-Bassa Forest, Jentink’s Duikers were readily recognized throughout the survey (R. Hoyt pers. comm.). A bushmeat study from Caspary et al. (1999) confirmed the presence of the species in the forests on Liberia’s eastern border. In Côte d’Ivoire, Jentink’s Duiker, like the Zebra Duiker Cephalophus zebra, has apparently never occurred further east than the Niouniourou R., its historical distribution therefore having been limited to the south-western part of the Guinean forest zone (Roth & Hoppe-Dominik 1990). Widespread forest destruction and expansion of agriculture in this region during the last 25 years have confined the species to the remaining areas of primary forest: Haut Dodo, Rapid Grah, Hana, Cavally-Gouin and Scio Forest Reserves. Its main stronghold is Taï N. P., where it is found in primary forest and rarely in secondary forest. It is observed relatively frequently in the well-protected western part of Taï N. P., including the isolated Mt Nienokoué, and the IET scientific study area near the city of Taï (Hoppe-Dominik et al. 1998). It is seen less in the central part of the park and the heavily poached eastern part of the park. Jentink’s Duiker is sometimes reported as occurring in Ziama and Diécké forests in Guinea (e.g. Brugiere & Kormos 2009), based on a report by Butzler (1994). However, Jentink’s Duiker is not recorded or mentioned at all by Sournia et al. (1990), Grubb et al. (1998), East (1999), or Wilson (2001), and was not reported by Barrie & Kanté (2006) in a rapid survey of Diécké in 2003. However, recent records from Mont Nimba Strict N. R. and the nearby Déré Classified Forest in 2009 (N. Granier pers. comm.) would lend support to their presence in SE Guinea. Habitat Jentink’s Duiker is only found in the high primary forest zone between Sierra Leone and the R. Niouniourou, a distribution that broadly coincides with many primate populations and also that of the Zebra Duiker.Within this zone, it does enter secondary growth, scrub, farms and plantations for brief periods (Davies & Birkenhäger 1990,

Abundance Jentink’s Duiker appears to be uncommon to rare throughout its range. East (1999), assuming an average population density of 0.1/km2, estimated the total population at about 3500, although Wilson (2001) doubted whether there were even more than 2000 animals left throughout the range. Peal & Kranz (1990) give densities of 1.0/km² in Liberia. Results from a survey in Taï N. P. in 2009 and 2010 covering 365 km recorded only a single sighting (N’Goran et al. 2009). In 2000, most hunters described Jentink’s Duikers as plentiful, but when pressed for information they reported only killing one or two individuals in their lifetime, even among hunters in their 60s. During a 10-month survey of hunters (representing 3% of the village populations) in three villages along the Juarzon–Pynes Town road near Sapo N. P., Jentink’s Duikers were the ninth most commonly killed species (n = 10) (R. Hoyt pers. comm.). The population trend of the species is downwards except for a few remote areas where forest destruction and forest hunting pressures are lower, for example Sapo N. P., and the few areas where there is better protection, such as the western section of Taï N. P. (Hoppe-Dominik et al. 1998, 2011). Adaptations As Jentink’s Duiker is very shy and secretive, few direct observations exist. In Taï N. P., the species has been observed mostly during the day (Hoppe-Dominik et al. 2011), and animals held in captivity in Monrovia Zoo were active day and night (Newing 2001). It has been reported that Jentink’s Duikers use their horns to uproot cassava plants so they can feed on the leaves, bark and tubers. Like Bay Duikers, they bolt from these daytime refuges with great speed if discovered, but have no stamina and do not go far. They are very residential and supposedly territorial, but make nocturnal forays out of thick forest, especially during periods when fruit is scarce (Davies & Birkenhäger 1990, Kingdon 1997). Foraging and Food Fruits and foliage. Known to enter plantations to eat palm nuts, mangoes and cocoa pods. The growing stems of tree seedlings are eaten, including Chlorophora and Hannoa klaineana (Davies & Birkenhäger 1990). Hunters familiar with the animal’s habits have identified many fruits with hard seeds or shells in its diet, notably kola nuts, erimado Ricinodendron sp., cherry mahogany Tieghemella sp., sand apples Panaria sp. and tallow tree Pentadesma sp. Jentink’s Duiker has also been reported chewing roots after exposing them with its hooves (Kingdon 1997). Social and Reproductive Behaviour Occasionally observed in pairs (Kranz & Glumac 1983), but most field observations are of lone animals, suggesting a primarily solitary social system. Reproduction and Population Structure Observations in Liberia from Sapo N. P. (Kranz & Glumac 1983) indicate that young

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are usually born between Mar and Jun, but young have been born in captivity in Monrovia between Nov and Jan. In captivity, young weigh 3–6 kg at birth (Pfefferkorn 2001). General recommendations of management and husbandry of captive duikers including the Jentink’s Duiker are given by Barnes et al. (2002). A captive individual lived to 21 years (Weigl 2005). Predators, Parasites and Diseases D. Jenny (pers. comm.) only once found hair from Jentink’s Duiker amongst 200 examined scats of the Leopard Panthera pardus in Taï N. P. It is possible that young animals are occasionally taken by the Crowned Eagle Stephanoaetus coronatus; however, they have not been found in prey remains examined in Taï N. P. (S. Shultz pers. comm.). The extent to which African Golden Cats Profelis aurata and the African Rock Python Python sebae are to be regarded as predators is unknown. There is no information on diseases or parasites. Conservation IUCN Category: Endangered C1. CITES: Appendix I. Due to the continued demand for bushmeat and unsustainable hunting, the risk that species dependent upon primary rainforests such as Jentink’s Duiker will die out is high. According to an examination by Caspary et al. (1999) in the region of Taï N. P., the influence of hunting on these species is considerable. Subsistence hunting provides inhabitants of the park with an estimated 1500–3000 tonnes of wild animal meat per year. In this way the rural population consumes about 28–55 g of meat/inhabitant/day. Jentink’s Duiker is poached both in the east and the west of Taï N. P. For example, a market examination of a stand in the east of the park counted about 3.3% Jentink’s Duiker (ten of a total of 2171 animals killed here).The share of Jentink’s Duiker killed in the forests to the west of the Cavally R. in Liberia and smuggled into Côte d’Ivoire was 2.5%. Of a total of 11,215 dead animals, 55 Jentink’s Duikers were counted. During a survey conducted in Liberia in 2002 by the Philadelphia Zoo, Jentink’s Duikers were not common in the Monrovia bushmeat markets and no preference for Jentink’s Duiker meat was identified, but carcasses were observed during the survey (R. Hoyt pers. comm.).

Although poaching still has a highly negative influence on the population, the long-term survival of Jentink’s Duiker is closely linked to the future of the remaining blocks of primary forest of the Guinean forest zone. At present only the western part of Taï N. P. receives effective protection. Rehabilitation of Sapo N. P. and development and implementation of protection and management programmes for the forests and wildlife as key areas are important. Better protection of areas such as Cestos–Sehnkwen, Sapo–Putu–Range, Grebo, Upper Krahn–Bassa in Liberia, Haute Dodo and Cavally Goin Forest Reserves in Côte d’Ivoire, and Western Area F. R. and Gola Complex Forest Reserves in Sierra Leone could result in a dramatic improvement of this species’ current conservation status (East 1999). Measurements Cephalophus jentinki HB: 1300–1500 mm T: 120–160 mm WT: 55–80 kg Throughout geographic range (Kingdon 1997) There are no reliable body measurements available, but weights reported by Wilson (2001) for animals from captivity ("": 56.5– 79.4 kg, n = 3 and !!: 60.3–76.7 kg, n = 4) compare favourably with those measures given above. Haltenorth & Diller (1980) give a shoulder height of 750–850 mm Wilson & Wilson (1990) recorded horn lengths ranging from 14.4 to 21.2 cm, and Davies & Birkenhäger (1990) recorded a pair 21.5 cm in length, which is slightly longer than the maximum horn length recorded by Rowland Ward of 21.3 cm from NE Liberia Key References East 1999; Kingdon 1997; Kranz & Glumac 1983; Wilson 2001. Bernd Hoppe-Dominik

Jentink’s Duiker Cephalophus jentinki.

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Tribe RAPHICERINI Grysboks, Steenbok, Beira Raphicerini Knottnerus-Meyer, 1907. Arch. Naturgesch. 73: 49.

Many of the small antelopes formerly allocated to the taxon Neotragini have long been thought to be basal to the antelope radiation, but so long as genealogical trees were not attempted the issue scarcely arose that these antelopes might not be monophyletic. Contemporary efforts to construct phylogenetic trees on the basis of relatedness (Hennig 1966) and newly available genetic information have challenged the monophyly of Neotragini as traditionally defined (Georgiadis et al. 1990, Gatesy et al. 1997, Hassanin & Douzery 1999, Matthee & Robinson 1999a, Rebholz & Harley 1999) and that taxon is now redundant for all of its former members barring Neotragus and Nesotragus. This dissolution of a convenient ‘waste-paper basket’ has raised, once again, the position of grysboks Raphicerus (and the Beira Dorcatragus megalotis) in relation to other antelopes and revived, in an entirely new context, the validity of assigning Raphicerus an exclusive tribe (including, very tentatively, Dorcatragus as a species that may have more complex affinities than are currently supposed). Haltenorth (1963), who used the tribe Raphicerini to accommodate Raphicerus as well as the Oribi Ourebia ourebi, included Dorcatragus in the tribe Dorcatragini. The nomenclatural history of this taxon has been summarized by Grubb (2001a): ‘Knottnerus-Meyer (1907) used the form Raphicerotinae; this was based on “Raphiceros Hamilton Smith, 1827”, a misquotation of Raphicerus Hamilton Smith, 1827.The correct form, Raphicerinae, was used by Frechkop (1955) (“justifiable emendation of Raphicerotinae”); Haltenorth (1963) used Raphicerini, which of course is just the tribal form of Raphicerinae and so counts as the same name under the Principal of Coordination. The Code, Art.35.4.1, says “A family-group name based on an unjustified emendation … or an incorrect spelling of the name of the type genus must be corrected”.’ In a composite tree of ruminant phylogeny, calibrated against a molecular clock, Hernández Fernández & Vrba (2005) estimated a 20 mya divergence for the Raphicerus lineage (they included the Beira as a sister species; see also Price et al. 2005). The dik-diks Madoqua spp. and the Oribi lineages were estimated to be equally old. Although these authors lumped the three lineages within Antilopini, 20 mya is between 5 and 10 millions years earlier, according to the same methods and authors, than the emergence of reduncines, hippotragines, alcelaphines and caprines, all well-defined and long accepted taxa. Aside from the issue of divergence times (according to Goodman et al. 1998, a 20–14 mya divergence is the acceptable timerank for a tribe), there are also many unsolved puzzles about details of the antelopes’ earliest radiations. The excessive lumping or glossing over of differences implicit in the neotragine ‘waste-paper basket’ has deterred recognition of polyphyly. Uncritical transfer of basal or near-basal taxa from a ‘Neotragini’ that is now known to have been polyphyletic (and see Gentry 1992) to similarly polyphyletic groupings under an enlarged ‘Antilopini’ merely perpetuates the problem. We have, therefore, resuscitated Raphicerini, hoping it will help underline the complexity and subtlety of the antelopes’ evolutionary radiation and provoke further research into the earliest radiations of antelopes.

Lateral view of skull of Steenbok Raphicerus campestris female.

Lateral view of skull of Cape Grysbok Raphicerus melanotis male.

Upper-right toothrow of Cape Grysbok Raphicerus melanotis.

The cranial morphologies, ecology and distribution of contemporary Raphicerus and Dorcatragus manifest two divergent trends in temperature regulation: the former species are adapted to relatively moderate temperatures, whereas the Beira tolerates much higher levels of heat. The ecological and geographic distributions of the two genera help provide a proximate explanation for such differences but any (very ancient) common ancestor was probably closer to living at the hotter end of the spectrum (although it is unlikely that any such primitive ancestor was as highly adapted to heat as the living Beira).The principal indication of divergent heat-tolerances lies in the sizes and proportions of the nasals and premaxilla, well developed in Raphicerus, much reduced in Dorcatragus. An opened-up nasal area allows for a flexible ‘nasal bellows’ mechanism to develop a cooling system that is illustrated in its most extreme form in the dik-diks. Sorting out the precise genetic relationships between the provisional tribes that are presented here (formerly all included in Neotragini), including Raphicerini, Madoquini, Ourebiini (and perhaps other lineages), is a task for the future. Details of behaviour and ecology in the four very different species that have been allocated to Raphicerini appear in the generic and species profiles. Jonathan Kingdon

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GENUS Raphicerus Grysboks, Steenbok Raphicerus C. H. Smith, 1827. In: Griffith et al., Anim. Kingd. 5: 342.

Living Raphicerus species are the Cape Grysbok R. melanotis and Sharpe’s Grysbok R. sharpei, constituting what Ansell considered a superspecies (Ansell 1972), and the Steenbok R. campestris. The Cape Grysbok is endemic to South Africa, being restricted almost entirely to the Cape Floristic Region; Sharpe’s Grysbok occurs in the savanna woodland from Tanzania to Swaziland, with a particular association with Brachystegia woodland; and the Steenbok has a disjunct distribution in East Africa (S Kenya, N and C Tanzania) and southern Africa. Earliest fossil records are of R. paralius, from 5.0 to 2.6 mya in South Africa (Gentry 1980, Vrba 1995); Steenboks and Cape Grysboks are known from Pleistocene deposits in South Africa. The three species are generally small, but absolutely larger than species of the genus Neotragus or Nesotragus. The pelage is uniform or streaked, with white underparts.The rhinarium is naked. Ears are large or very large; the tail is short and not tufted. Preorbital glands open through single small orifices, and pedal glands are present (the Cape Grysbok " also has a preputial gland); however, there is no subauricular gland, and no inguinal glands. Females have two pairs of inguinal nipples. Horns are present in "" only, rising nearly vertical, slender, cylindrical and smooth with almost no indication of annuli. The skull is stout with the rostrum much shortened and tapered compared with Ourebia. Other characteristics of the skull include: small but deep lachrymal depressions with upper and lower margins not forming sharp ridges; secondary deposition of bone on frontals and parietals; well-marked, widely spaced temporal ridges; ethmoid vacuity present; and the dentary with rounded angle. The first incisors are less broadened than in Ourebia; other incisiform teeth are narrow, but the second incisors are broader than the rest. The cheekteeth are small (though the premolars are relatively large) and low-crowned; the occlusal pattern is simple with styles not well marked. Cape Grysboks were formerly assigned to their own genus Nototragus (Thomas & Schwann 1906), based on their retention of small lateral or false hooves and substantial differences in limb proportions and stance. Subsequently, both Shortridge (1934) and Roberts (1951) included both the Cape and the closely related Sharpe’s Grysbok in Nototragus to distinguish the two shorter-legged, broader-mouthed grysboks from the longer-legged, smaller-mouthed and nominate Steenbok. Kingdon (1982) pointed out the differing lips and mouth structure between R. melanotis/sharpei and R. campestris and correlated these differences with diet and habitat. Haltenorth (1963) even considered the two grysbok species conspecific. However, Ellerman et al. (1953) and Ansell (1972) thought there were no grounds for recognizing Nototragus, considering both R. sharpei and R. melanotis as distinct enough to warrant species status; subsequent authors have followed this treatment (e.g. Meester et al. 1986, Grubb 1993c, 2005, Bronner et al. 2003). Species are distinguished by colour of pelage, size of ears, length of horns, relative length of limbs, and presence or absence of lateral hoofs.

Steenbok Raphicerus campestris myology. Steenbok Raphicerus campestris skeleton. BELOW: Facial myology of Raphicerus spp. compared, showing mouths predominantly adapted to browsing, Sharpe’s Grysbok R. sharpei (above) and to grazing, Steenbok R. campestris (below). TOP:

ABOVE:

Peter Grubb

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Raphicerus melanotis CAPE GRYSBOK Fr. Grysbok du Cap; Ger. Kaapgreisbock Raphicerus melanotis (Thunberg, 1811). Mem. Acad. Imp. Sci. St. Petersbourg 3: 312. No locality cited, but selected as the Cape Peninsula, Western Cape Province, by Grubb (1999c) based on the records of Levaillant who encountered the species on the peninsula. Rookmaker (1988) noted that the Cape Grysbok was first named by J. R. Forster as both Antelope melanotis and A. grisea, both considered as nomina nuda.

Cape Grysbok Raphicerus melanotis. Lateral and palatal views of skull of Cape Grysbok Raphicerus melanotis. LEFT:

ABOVE:

Taxonomy Monotypic. Several authors (Haltenorth 1963, Haltenorth & Diller 1980) have considered the Cape Grysbok and Sharpe’s Grysbok Raphicerus sharpei to be conspecific (separating the species into two subspecies), but Ellerman et al. (1953), Ansell (1972) and many other authors have treated them as distinct (and see Genus profile). Synonyms: grisea, rubroalbescens, rufescens. Chromosome number: not known. Description A small reddish-brown antelope, standing only about 0.5 m at the shoulder, with a hunched or curved back, and short inconspicuous tail. The muzzle is short and with a black bridge to the nose.The common name is derived from the conspicuous white hairs visible in the overall rufous pelage on the sides and the back, which creates a greyish appearance (‘grys’ is grey in Afrikaans); the undersides are a lighter uniform brown. Ears relatively large with prominent whitish hairs on the inside whereas the outside is greyish. Supplementary or ‘false’ hooves are present above the fetlocks on all four legs (commonly absent in other Raphicerus species). Active preorbital glands are present, and pedal glands are present on all four feet; "" also have preputial glands (Manson 1974). Horns present in " only, smooth and rising vertically from the head with a slight forward curve. Geographic Variation None recorded. Similar Species Raphicerus sharpei. Allopatric in savanna woodland from Tanzania to Swaziland. Slightly smaller (mass of ±7.5 kg), and a slightly paler reddish-brown pelage, also interspersed with white hairs, whilst

the underparts are also comparatively lighter having been described as buffy-white; false (lateral) hooves absent, being reduced to areas of thickened skin (Ansell 1972). R. campestris. Broadly sympatric. Slightly heavier (±11 kg), with a relatively straight back (as opposed to the hunched or arched appearance of Cape Grysbok), and tends to live in more open areas than the scrub-loving Cape Grysbok; more uniform red-brown without interspersed white hairs, the underparts are white or nearly so; horns longer (up to 190 mm); false (lateral) hooves absent. Distribution The Cape Grysbok is endemic to South Africa. Boshoff & Kerley (2001) referred to the Cape Grysbok as nearly endemic to the Cape Floristic Region, occurring in montane and lowland habitats (with the exception of dense forest) to the south and west of the Cape Fold Mts. Skinner & Chimimba (2005) stated it occurs ‘from the Cedarberg Mts in the Western Cape to the Vredenburg area and southwards and eastwards coastally to the Albany (= Cacadu) and Bathurst (= Cacadu) districts in the Eastern Cape, with records from the Komga (= Amatole) district’. Shortridge (1934) gave the extreme north-eastern limits as Port St Johns although there appears to have been some contention as to the precise limits for the Cape Grysbok along the E South African coastline (Skead 1987). Skead (1980) cited the reports of Barrow (1804) who stated that ‘steenbok, oribi and grysbok were still plentiful in 1799 in the “Onder Bokkeveld”’ but went on to warn that doubts should be cast on the records of both Cape Grysboks and Oribis (Ourebia ourebi)for that region. Distribution extends into the mountain renosterveld of the Nieuwoudtville region and Cape Grysboks have been observed in

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Oorlogskloof N. R. (C. T. Stuart & T. Stuart pers. comm.). The distribution of the species within this vegetation transition zone between the Fynbos and Succulent Karoo biomes of southern Africa is likely to be restricted to valleys and gorges with dense vegetation. Their northern limits would appear to have been the southern parts of the Great Karoo, probably south of the Escarpment. An apparently isolated population in the Eastern Cape has been indicated by two questionnaire surveys (Bigalke & Bateman 1962, Lloyd & Millar 1983) supported by personal interviews (Lynch 1989). Records from landowners have been substantiated by additional observations from the Barkly East district (Stuart 1984) and skins obtained in the vicinity of Maclear (Feely 1992). Archaeological records from this region also point to the presence of Cape Grysboks in the area dating back some 2000–8000 years ago (Feely 1992). The species has also been recorded in the southern Drakensberg (Vincent 1962, Bourquin 1966, cited in Rowe-Rowe 1994) (for earlier records see Du Plessis 1969). There is a substantial gap between the southern limits of Sharpe’s Grysbok and the north-eastern limits of the Cape Grysbok. Habitat Bigalke (1979) stated that although it occurs in other vegetation types, the Cape Grysbok is ‘essentially an animal of the Cape fynbos’, in both winter and non-seasonal rainfall areas. Cape Grysboks prefer dense cover and closed-canopy habitats. The propensity of the species to select dense vegetation is particularly true in the western portion of its distribution, where dense fynbos (scrub) communities dominate. Towards the east of their range Cape Grysboks also utilize the dense thicket habitats associated with many of the river valleys and coastal regions. Presence in the high-altitude grasslands of the north-eastern Cape is conditional on the proximity of forest fragments and bush clumps, but, in some areas, Cape Grysboks may also use long grass for cover. Although much of the Cape Floristic Region is now cultivated, the Cape Grysbok is one of the few antelope that can survive in relatively small patches of natural vegetation surrounded by cultivated lands (Manson 1974), and Skinner & Chimimba (2005) suggest that such cultivation may sometimes actually improve habitat for Cape Grysboks provided that some dense cover is nearby.As commercial rooibos tea plantations have become established in the Cedarberg Region, the Cape Grysbok has been blamed, along with the Common Duiker Sylvicapra grimmia, for extensive damage to new shoots (C. T. Stuart & T. Stuart pers. comm.). Abundance Cape Grysboks are normally solitary and cryptic in their behaviour and therefore seldom seen.Visibility is further reduced in dense vegetation and this is exacerbated in fire-prone areas such as the fynbos of the Western Cape. Fires, bush clearing, grazing and agriculture all affect the visibility of Cape Grysboks in the field and can influence counts. Boshoff et al. (2002) estimated the spatial habitat requirements of medium- to large-sized mammals in the Cape Floristic Region and compared some of their predictions with empirical data. Predictions from the model, based on intact habitat, suggested that Cape Grysboks required between 6 and 456 ha per animal depending on the nature of the vegetation type. Based on the available habitats this would suggest that Cape Grysbok numbers could lie between 231,448 (post-habitat transformation) and 322,977 (pre-habitat transformation) in the Cape Floristic Region (Kerley et al. 2003). This is almost an order of magnitude higher than earlier estimates of their abundance in an area covering 61,000 km2 (East 1999).

Raphicerus melanotis

Cape Grysbok numbers in the Eastern Cape appear subject to local cyclical fluctuations in population numbers (Skead 1980, 1987), but this may also apply more widely to the Western Cape (P. H. Lloyd pers. obs.). Gamkaberg N. R. in the late 1990s had a notably conspicuous population of Cape Grysboks on top of the mountain (as illustrated by the fact that Norton [1986] listed Cape Grysboks, Klipspringers Oreotragus oreotragus and Grey Rheboks Pelea capreolus as the most abundant antelopes). In the early 2000s, animals were rarely seen because a large fire had intervened and it is possible that the animals were destroyed, as well as much of their cover being destroyed, necessitating movement to less accessible areas. Skead (1987) also stated that outbreaks of ophthalmia (with increased vulnerability to predation) might have been implicated in some of these declines. Scott (1991) studied the distribution of small antelopes in De Hoop (Provincial) N. R. between 1985 and 1987 and recorded density indices (number of animals per 100 km travelled) of 0.21 for Cape Grysboks compared with 2.64 for Steenboks. This might suggest that Steenboks are at least ten times more abundant than Cape Grysboks but, when habitat preferences and relative visibility are taken into account, it is likely that Cape Grysboks have been substantially underestimated. In some protected areas the abundance of Cape Grysboks appears to be linked to the density of other herbivores, particularly where such herbivores alter the structure of the vegetation. Grysbok ranges have seen local declines in numbers from areas such as Addo Elephant N. P., where escalating numbers of elephants have opened up or transformed thicket habitats (J. G. Castley pers. obs.). Adaptations As described elsewhere, the Cape Grysbok is the only member of the genus Raphicerus that consistently has false hooves, a peculiarity that implies a more primitive evolutionary state within the Bovidae. Likewise, skull structure conforms with that of early, relatively unspecialized antilopine bovids (Kingdon 1982). In general, therefore, 305

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the Cape Grysbok appears to represent the relatively unmodified descendant of an early line of antelope, rather than a more recently adapted form. Only the " is horned in this species and Novellie et al. (1984) have gone into some detail on the nature of conflict encounters between "" and suggested that damaging encounters rather than ritualized ones might be the more appropriate evolutionary strategy for the Cape Grysbok. Their arguments were based on the assertion that for small species, occupying small territories, retreats from conflict encounters put the retreating individual at a disadvantage.They argued that losing meant loss of access to forage as well as mating opportunities. Hence the unelaborated horn shape, a lack of observed rituals and retention of short, smooth, stabbing weapons in "". The need to hold and demarcate small territories has probably influenced the intensity of other activities such as scent-marking using the preorbital glands. To date a total of 85 chemical compounds have been isolated from the black aqueous secretion of the preorbital gland (Burger et al. 1981b, 1996), implying a highly complex olfactory signalling system. Cape Grysboks are small antelopes that appear to be dependent on dense, bushy or scrubby vegetation.The archaeological record suggests that their abundance has been directly related to the bushiness of the vegetation (Klein 1983). In the archaeological sites evaluated (which were southern and south-western areas of the Cape Floristic Region, as opposed to grassland areas further east), its abundance was thought to have fluctuated over the last two interglacials according to the availability of bushy vegetation. Foraging and Food Cape Grysboks are predominantly browsers, although earlier accounts reported Cape Grysboks as predominantly grazers (Dorst & Dandelot 1970, Smithers 1983). Manson (1974) recorded Cape Grysboks grazing, but he only made one such observation out of 30 in total (although he conceded that grass might play a more important dietary role in other areas of their distribution). Kigozi et al. (2008), by investigation of faecal samples, also concluded that Cape Grysboks were selective browsers and had a restricted diet of only nine species in the Acacia-dominated coastal dune fynbos of the Port Elizabeth region. Interestingly, these invasive species, notably Acacia cyclops, formed a significant part of the diet; no monocotyledonous plants were identified in the grysbok faecal samples. A wider variety of species is eaten in the western portions of their range where Cape Grysboks have become a problem to farmers as more and more of the habitat has been taken over for vineyards, orchards (Manson 1974) and rooibos tea plantations (C. T. Stuart & T. Stuart pers. comm.). More recent evidence suggests that Cape Grysbok are able to broaden their diet in response to changing forage availability. Such dietary flexibility also influences the amount of grass in the diet, which at times can be substantial (Kerley et al. 2010, Faith 2011). Because they tend to eat fresh-growing shoots, Cape Grysboks’ water requirements are limited, but they have even been observed to utilize water with a high salt content (Manson 1974). During the period of his study Manson (1974) found that the greater part of the animals’ feeding time was at night. He found feeding periods of 2–3 hours interspersed with periods of rest during which they ruminated. Browse is mostly cut by the premolars, but the bite selection process also occasionally involves the lower incisors.

Social and Reproductive Behaviour Cape Grysboks are primarily solitary animals. Out of approximately 40 sightings in the wild, only two were of pairs: in one case a ! with her offspring, and in the other a " with a !, during Sep (Manson 1974). Cape Grysboks are territorial, particularly the "", but even the more tolerant !! exhibit strong territorial characteristics (Manson 1974). Novellie et al. (1984) noted that Cape Grysboks, particularly "", had well-defined home-ranges averaging 2.5 ha (range 1.26–4.84 ha) and suggested that pairs did not share territories. Males mark their territories by using their preorbital glands to mark twigs and grass stalks, leaving a conspicuous black sticky deposit. Females also have prominent preorbital glands with secretions clearly visible, but they appear to mark infrequently (only two observations being recorded, and those in captive animals) while juveniles of both sexes have never been observed using their preorbital glands (Novellie et al. 1984). On several occasions "" have been noted biting off twigs prior to marking the stems with their preorbital glands. Marking tends to take place at night and dominant animals mark more frequently than subordinates (Novellie et al. 1984).The effect of pedal, or interdigital, gland marking is less obvious, but animals were noted scratching the ground whenever establishing a new dung midden. Number of middens used in the wild has yet to be established, but both adults and juveniles use the same midden in captivity (Novellie et al. 1984). In captivity, adult "" appear, without exception, to exhibit aggression to other adult "", with the dominant animal deposing subordinates regardless of supposed territorial boundaries (Novellie et al. 1984). Unrelated !! also exhibit relatively little tolerance of other !!. Subordinate animals generally exhibit submissive behaviour by lying down, upon which the dominant animal approaches and makes naso-nasal contact. Occasionally, subordinate animals creep closer on their knees. At no stage can more than one adult " be kept together in an enclosure. All attempts at keeping two rams have ended in major aggression, necessitating separation. When a " discovers a midden with female excretions, he first smells it, frequently following this with flehmen behaviour (typically lip and tongue movements and outstretched horizontal neck). Once oestrus has been detected and contact is made the " marks the ! with preorbital gland secretions, thereby (according to Manson 1974) making her a part of his known environment. Once visual contact has been made the " approaches and rubs against the female’s head and neck, sometimes the rest of her body, occasionally licking her face, neck and shoulders. This sequence of behaviour is often repeated during the early courtship phase, but abates later. Manson (1974) also cited Ewen (1956) as having observed ‘butting matches’ during courtship, but did not observe it himself. Males pay particular attention to the female’s urogenital region during later stages of courtship, continually licking and following as the ! moves forward: the intensity increases and following becomes incessant. This appears to stimulate urination by the !, notably on established middens, and each time the " again exhibits typical flehmen behaviour. Unsuccessful mating attempts often follow and these are caused by sudden forward movements by the !, with the " standing on his hind legs, front legs folded to the chest, and very little, if any, actual body contact. Males often make bleating sounds while licking the !, or just prior to mounting. Females also bleat during encounters but less frequently than "". Novellie et al. (1984) also reported a greater frequency of licking and bleating behaviour by the " during

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periods of successful mating and he suggested that there are seasonal peaks (Mar–Jun) in this reproductive behaviour. Successful copulation only lasts for a few seconds and laufschlag (foreleg mating kick) was never observed in Cape Grysboks (Novellie et al. 1984). The young of territorial species are typically concealed, or conceal themselves, until sufficiently mature to emerge. Manson’s (1974) sole observation of such an event was of a Cape Grysbok young that hid itself after being suckled. Suckling apparently takes place with the mother either standing or lying down. Reproduction and Population Structure A single young is born at any time of the year, although a captive group showed a birth peak between Sep and Dec (Novellie et al. 1984); Spence (2003) notes birth peaks in captivity in Mar/Apr and Oct/Nov. Skinner & Chimimba (2005) noted that under favourable conditions !! can have up to two young in one year. Skinner & Chimimba (2005) indicated that !! have their first young at 17–26 months while "" become sexually mature at 17– 18 months. Manson (1974) observed a ! give birth at the age of approximately 18½ months, suggesting that sexual maturity in !! can occur at least as early as 12 months (which agrees with Spence [2003]). Based on subsequent observations, the gestation period appears to be approximately 192 days, or six months (Manson 1974). In captive animals at Tygerburg Zoo, Spence (2003) estimated gestation at 171– 180 days, and recorded inter-birth intervals of 177–215 days. One captive specimen lived to more than seven years (Weigl 2005). Predators, Parasites and Diseases Stuart (1981) identified Cape Grysbok remains in the stomach contents of both Caracals Caracal caracal and Leopards Panthera pardus. Skead (1980) reported that a considerable number of farmers were convinced that Cape Grysbok numbers had declined due to an alleged increase in Caracal numbers, but he noted that, at least in the Eastern Cape, these appeared to be cyclical events for both Caracals and Cape Grysboks. In West Coast N. P., Cape Grysboks are likely to have contributed to the diet of Caracals even though they are a low-density species compared with other ungulates. Even so, Avenant & Nel (1997, 1998) found that ungulates only made up about 7% of the Caracal diet (the bulk being rodents). Another probable carnivore predator is the Black-backed Jackal Canis mesomelas, whilst Ratels Mellivora capensis could potentially prey on young. Langley (1986) reported a number of attacks on Cape Grysbok young by Cape Grey Mongooses Herpestes pulverulentus, suggesting that newborns may be under predation risk, even from small carnivores. Other predators known to prey on Cape Grysboks include some of the larger raptors, such as Crowned Eagles Stephanoaetus coronatus (Boshoff et al. 1994), Verreaux’s Eagles Aquila verreauxii (Boshoff et al. 1991), and probably Martial Eagles Polemaetus bellicosus (Boshoff & Palmer 1980, Boshoff et al. 1990). A number of studies have investigated the parasite loads on Cape Grysboks and compared them with other susceptible species (Boomker et al. 1989a, MacIvor & Horak 2003, Watermeyer et al. 2003). Boomker et al. (1989a) reported that nematodes of nine species as well as a further two genera and cestodes of one species

and a further genus were recovered. Highest loads were of the nematodes Skrjabinema spp., Trichostrongylus pietersei and T. rugatus. High numbers of the nematode Skrjabinema spp. (parasites that are commonly reported to occur in high numbers in grazing antelopes) has lent indirect support for some grazing activity taking place in Cape Grysbok (Boomker et al. 1989a). The nematodes Haemonchus contortus and Impalaia tuberculata were not recorded in this survey and Boomker et al. (1989a) felt that earlier records (Mönnig 1931, 1933) should be treated with caution as the hosts may actually have been Steenboks. Watermeyer et al. (2003) identified a new parasite record for the Cape Grysbok (Setaria saegeri). One of the most frequent disease issues reported to conservation authorities in the Western Cape (P. H. Lloyd pers. obs.) and Eastern Cape (Skead 1987) is that of infectious ophthalmia in Cape Grysboks. By contrast, relatively few reports have been received of incidents of this infection in Steenboks. Since the organism causing the disease, Rickettsia conjunctivae, is frequently found in, and dispersed by, a variety of flies and other arthropods, it would appear that the Cape Grysbok’s selection of dense vegetation could be a contributory factor, since many flies use shaded areas under vegetation. Conservation IUCN Category: Least Concern. CITES: Not listed. The Cape Grysbok is officially conserved in the majority of formal conservation areas in the Western Cape, as well as in many others in the Eastern Cape. It is known from seven National Parks, including Table Mountain N. P., West Coast N. P., Agulhas N. P. and Wilderness N. P., and at least 27 Provincial Nature Reserves. The long-term viability of state parks and reserves as well as the increasing number of privately owned reserves will be contributory factors to the future persistence of the species. For many of the private reserves and even some of the state-protected areas the continued presence of undesirable species (extralimital, exotic or domestic) that outcompete or negatively impact on the Cape Grysbok should be a major concern. Measurements Raphicerus melanotis TL (""): 771 (450–810) mm, n = 4 TL (!!): 793 (720–815) mm, n = 5 T (""): 58 (40–72) mm, n = 4 T (!!): 54 (44–60) mm, n = 5 HF c.u. (""): 245 (240–250) mm, n = 4 HF c.u. (!!): 248 (235–265) mm, n = 5 E (""): 113 (105–115) mm, n = 4 E (!!): 115 (110–120) mm, n = 5 WT (""): 10.0 (10.0) kg, n = 4 WT (!!): 10.5 (8.8–11.4) kg, n = 5 Western Cape (C. T. Stuart, in Skinner & Chimimba 2005) Maximum recorded horn length is 13.3 cm for a pair of horns from the Western Cape, South Africa (Rowland Ward) Key References Manson 1974; Novellie et al. 1984. Guy Castley & Peter Lloyd

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Raphicerus sharpei SHARPE’S GRYSBOK Fr. Grysbok de Sharpe; Ger. Sharpegreisbock Raphiceros sharpei (Thomas, 1897). Proc. Zool. Soc. Lond. 1896: 796, pl. 34 [1867]. S Angoniland, Malawi.

Sharpe’s Grysbok Raphicerus sharpei.

Taxonomy Sharpe’s Grysbok has been considered synonymous with the Cape Grysbok Raphicerus melanotis (Haltenorth 1963, Haltenorth & Diller 1980), but Ellerman et al. (1953) and Ansell (1972) regarded both as distinct species (and see Genus profile). Ansell (1972) recognized two subspecies: R. r. sharpei, a supposedly smaller, paler form from the northern parts of the species’ range, and R. r. colonicus, a larger, marginally darker form from the southern parts of the range. However, the precise limits of distribution between the two have never been very clear, and the validity of these subspecies is doubtful. Synonyms: colonicus. Chromosome number: not known. Description Small antelope with a rich, reddish-fawn body colouration and a liberal sprinkling of white hairs on the dorsal parts, shoulders and flanks. Eyes surrounded by an ill-defined whitish ring. Ears with buffy-white hair inside; dark on the outside. A short, dark band on top of the muzzle extends from the rhinarium to the front of the eyes. Sides of the face, outer parts of the limbs, forehead and upperparts of the muzzle yellowish-brown, lacking the white grizzling seen on the dorsal parts. Underparts, throat and insides of the legs paler than upperparts, almost white. Body hairs fairly long (30 mm). Lateral (false) hooves are absent (present on hindlegs of Cape Grysbok). Preorbital glands are present as small patches of black naked skin, about 2 mm in diameter and 3 mm deep (Ansell 1964). Preputial gland present, the opening lying anterior to that of the urethral canal. Pedal glands on both fore- and hindfeet, opening into the interdigital cleft that is well covered with hair (in contrast to the Cape Grysbok in which it is very sparsely haired [Ansell 1964]). No obvious sexual dimorphism, although meagre data from animals taken in Zimbabwe and E Zambia suggest that ! is slightly larger. Horns present in " only, short and sloping slightly backward, tapering to sharp points. Geographic Variation See Taxonomy.

Lateral and dorsal views of skull of Sharpe’s Grysbok Raphicerus sharpei.

Similar Species Raphicerus campestris. Marginally sympatric in the northern and southern

parts of the range of Sharpe’s Grysbok, but absent from the miombo Brachystegia woodlands of N Mozambique, Malawi and much of Zambia. Slightly larger (±11 kg), with smoother coat, more uniformly coloured, and lacking white speckling; black-rimmed eyes very conspicuous against pure white margin; horns longer (max. 190 mm); back profile more horizontal and less hunched. R. melanotis. Allopatric, with distribution nearly confined to the Cape Floristic Region. Slightly larger (mass of ±10 kg), with conspicuous white hairs visible in an overall rufous pelage; false (lateral) hooves present. Distribution Endemic to Africa, occurring in savanna woodland from Tanzania to Swaziland. They are recorded from the western and southern parts of Tanzania (close to L.Victoria at their northern limits). Distribution then extends southwards through SE DR Congo, most of Zambia (though not west of the Zambezi R.), Malawi and Mozambique (not including the coastal forested regions), to extreme NE Botswana and the eastern Caprivi Strip in Namibia, much of Zimbabwe (including the dry western parts, such as Hwange) and NE South Africa (Limpopo Province, E Mpumalanga), and E Swaziland (Ansell 1978, Kingdon 1982, Ansell & Dowsett 1988, Monadjem 1998, East 1999, Skinner & Chimimba 2005). Not yet recorded from SE Angola or N KwaZulu– Natal Province. Habitat Across much of their range (Zimbabwe, Zambia and Tanzania) they appear to be associated with Brachystegia woodland where there is good undercover in the form of low-growing scrub or medium-length grass (Smithers & Wilson 1979, Kingdon 1982). In some parts of Zimbabwe and Tanzania they live in rocky terrain with low bush and grass cover, often around kopjes and stony ridges. In

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Sharpe’s Grysbok Raphicerus sharpei detail of head.

Raphicerus sharpei

Hwange N. P., Zimbabwe, they are common in riverine vegetation, but also occur in broken country where there are thickets at the base of kopjes (Wilson 1975). Although they appear to be associated with areas of good ground cover, Wilson (1975) recorded that, in Hwange N. P., they were also found in pure stands of mopane Colophospermum mopane with very little good cover and in Chobe N. P., Botswana, they were seen on a sandy plateau in open woodland with light grass and scrub cover. Wilson (1975), referring to his 1969/71 survey of Hwange N. P., indicated that Sharpe’s Grysbok was confined to the northern part of the Park, and was replaced in the southern sections by the Steenbok. At that time the species was common along the Lukozi R. where there were considerable riverine thickets. Over a distance of 10 miles, 19 Grysboks were seen on the evening of 9 October 1969. In 1996, another detailed survey of the Park was undertaken and Wilson (1997) noted: ‘the Lukozi River Marked Transect was covered on no less than 18 occasions by several different teams and at different times of the day throughout the year and not one Sharpe’s Grysbok was seen’. Wilson (1997) goes on to say: ‘I have mentioned previously … that 25 years ago the riverine vegetation along the Lukozi River was very dense and that was when grysbok were found there.Today [1996] the vegetation has thinned out considerably as a result of elephant destruction and grysbok no longer occur along the Lukozi River Drive where 25 years ago they were common. However, their distribution may well have shifted southwards into the broken country south of Lukozi and east to Shumba where they are now found.’ The species also still occurs in other areas in the northern parts of Hwange N. P. Abundance East (1999) summarized recorded population density estimates for this species (0.3–0.7 ind/km2), and estimated a total population size of about 95,000 animals. Sharpe’s Grysboks are predominantly nocturnal, exceptionally shy and secretive, and can be overlooked in areas where in reality they are reasonably common, so they may be more abundant than supposed.

Adaptations This species has remarkably robust teeth correspondingly deeply embedded in skull and mandible. The mouth has a substantially wider maw than in the Steenbok.These dental and oral peculiarities correspond to a more demanding diet, probably including more lignified and mature growth than is taken by Steenboks, which prefer newer younger shoots. The extreme caution and mainly nocturnal activity of this species also contrast strongly with the commonly diurnal and often conspicuous behaviour and appearance of the larger Steenbok. The shorter legs and cryptic colouration of Sharpe’s Grysbok imply greater vulnerability to predators, which probably include both coursers and large raptors. Flight behaviour includes a brief, stamping pronk and a fast dash for dense cover. The longer-legged Steenbok instead may race away in the open, jinking from side to side before hiding. During the game rescue operations that took place during the formation of L. Kariba, Sharpe’s Grysboks were recorded as readily taking to water, and were said to swim well for their size (Child 1968). Foraging and Food Predominantly a browser, but they also will graze, and as such are classified as browser–grazer intermediates by Gagnon & Chew (2000) in their review of the dietary preferences of African bovids. In SE Zimbabwe, stomach contents consisted of 70% browse and 30% grass, with important browse items including: Acacia spp., sand olive Dodonaea viscose, buffalo thorn Ziziphus mucronata, false marula Lannea edulis, horn-pod tree Diplorhynchus condylocarpon, beechwood Faurea saligna and bitter albizia Albizia amara (Smithers & Wilson 1979). Wilson (1975), in Hwange N. P., recorded Diospyros lycioides fruits, Grewia flavescens fruits and leaves, the leaves and soft stems of Commelina sp., the leaves and fruits of Cyphostemma buchananii, and a number of unidentified grasses. Although they feed mainly at night, Sharpe’s Grysboks may be seen foraging in the early morning or late afternoon, lying up during the heat of the day in dense cover (Smithers 1971). Social and Reproductive Behaviour Very little is known about the behaviour of this species.They usually occur solitarily, in pairs, or a ! with her single offspring. A loosely connected pair may share a territory throughout the year (Kingdon 1982). Droppings are apparently placed in small middens, which are used over long periods (Smithers 1971). The latter suggests that Sharpe’s Grysboks do not undertake extensive seasonal movements. They are notoriously secretive and reclusive, inclined to lie up very tightly hidden in the undergrowth. When they do run off they often 309

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Sharpe’s Grysbok Raphicerus sharpei.

crouch low to the ground as they run through the thick underbush, in contrast to the Common Duiker Sylvicapra grimmia, which bounds through the undergrowth, or the Steenbok (described above). They have been recorded hiding in Aardvark Orycteropus afer burrows (Shortridge 1934, V. J. Wilson pers. obs.). Reproduction and Population Structure Pregnant !! have been recorded throughout the year (Shortridge 1934, Ansell 1960a, Kerr & Wilson 1967, Smithers 1971, Smithers & Wilson 1979, V. J. Wilson pers. obs.), suggesting that they are aseasonal breeders. A single young is born following a gestation of about seven months. Birth-weight in captivity is around 830 g (range 790–863 g, n = 5; V. J. Wilson pers. obs.). There is limited information available on population structure, although during the game rescue operations on L. Kariba, of 222 sexed individuals recorded in reports, there were 113 "" and 109 !! (V. J. Wilson pers. obs.).

Banhine N. P. (Mozambique), Hwange N. P. and Gonarezhou N. P. (Zimbabwe) and Kruger N. P. (South Africa) (East 1999). Measurements Raphicerus sharpei TL (""): 751 (710–800) mm, n = 12 TL (!!): 757 (725–800) mm, n = 11 T (""): 58 (50–70) mm, n = 12 T (!!): 59 (45–70) mm, n = 11 E (""): 90 (83–98) mm, n = 12 E (!!): 91 (85–98) mm, n = 11 WT (""): 7.3 (6.8–8.9) kg, n = 12 WT (!!): 7.7 (6.4–8.9) kg, n = 11 SE Zimbabwe (Smithers & Wilson 1979)

TL (""): 760, 880 mm, n = 2 TL (!!): 763 (550–820) mm, n = 8 Predators, Parasites and Diseases Predators include Lions T (""): 60, 70 mm, n = 2 Panthera leo, jackals Canis spp., Caracals Caracal caracal, African Rock T (!!): 78 (50–90) mm, n = 8 Pythons Python sebae and large raptors. On one occasion a 10-foot WT (""): 8.9 (6.3–11.8) kg, n = 4 python, captured on the Lukozi R. in Hwange N. P., was being stretched WT (!!): 9.0 (7.2–10.8) kg, n = 19 out in order to measure it and it regurgitated a partly digested adult E Zambia (V. J. Wilson pers. obs.). Sharpe’s Grysbok ! (V. J.Wilson pers. obs.). A single animal examined In E Zambia, an additional seven adult "" had a mean mass of in the Central Province of Zambia was infected with Amblyomma 8.9 kg (range 7.1–10.7 kg), and eight adult !! 9.9 kg (range 8.5– variegatum, Rhipicephalus appendiculatus, R. kochi and R. punctatus; R. 11.9 kg). The mean masses of animals caught at Kariba during game appendiculatus constituted 94% of all ticks present (Zieger et al. 1998b). rescue operations was: !! 10.1 kg (range 8.6–11.1 kg, n = 9); "" 9.6 kg (range 9.0–10.1 kg, n = 6) (V. J. Wilson pers. obs.) Conservation IUCN Category: Least Concern. CITES: not listed. Maximum recorded horn length is 10.4 cm for a pair of horns from Although they have been extirpated from parts of their range by Massingir, Mozambique (Rowland Ward) expanding human settlement and localized hunting for bushmeat, Sharpe’s Grysbok remain relatively widespread with about one-third Key References Smithers 1971; Wilson 1975; Smithers & of the total population occurring in protected areas, including: Selous Wilson 1979. G. R. (Tanzania), Upemba N. P. (DR Congo), Kafue N. P. and North and South Luangwa National Parks (Zambia), Lengwe N. P. (Malawi), Michael Hoffmann & Vivian J. Wilson 310

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Raphicerus campestris

Raphicerus campestris STEENBOK (STEINBUCK, STEINBOK) Fr. Steenbok; Ger. Steinbockchen Raphicerus campestris (Thunberg, 1811). Mem. Acad. Imp. Sci. St. Petersbourg 3: 313. No locality cited; since selected as Western Cape Prov., Malmesbury Div., Swartland (Grubb 1999: 23).

Steenbok Raphicerus campestris.

Taxonomy The colouration of Steenboks varies widely throughout their range, and has partly contributed to the recognition of so many subspecies. Ansell (1972), in his revision of the taxonomy of the species, listed eight subspecies, noting that their acceptance was provisional and that study of a more complete series of specimens would lead to a reduction in the number of recognized subspecies. Of the eight listed by Ansell (1972), only one was listed for East Africa, namely R. c. neumanni, one (R. c. kelleni) for Angola and Zambia, and the remaining six for southern Africa; Meester et al. (1986) reduced this list to five. Pending a much-needed taxonomic revision, the most reasonable approach appears that of Kingdon (1997), who lists only two subspecies: the nominate form from the southern part of the range, and R. c. neumanni for East Africa. Synonyms: acuticornis, bourquii, capensis, capricornis, cunenensis, fulvorubescens, grayi, hoamibensis, horstockii, ibex, kelleni, natalensis, neumanni, pallida, pediotragus, rufescens, rupestris, steinhardtii, stenbock, stigmatus, subulata, tragulus, ugabensis, zukowskyi, zuluensis. Chromosome number: 2n = 30 (Wallace & Fairall 1967a). All autosomes are metacentric; the X-chromosome is a large acrocentric and the Y is a small metacentric. Description A small, gracile antelope (about 0.5 m at the shoulder), with long, slender legs, a glossy light brown to reddish-brown pelage on the flanks and back and white underparts. Ear pinnae large with white inside, light grey on the back and a thin black line around the pinnae rims. Throat and chin white, nose and dorsal ridge of rostrum black, and there is a distinctly pale or white eyebrow line, giving the animal a neatly groomed appearance. The brown eyes are rimmed by black skin that enlarges their visual impact. Hooves narrow and sharp; false

Lateral, palatal and dorsal views of skull of Steenbok Raphicerus campestris.

hooves absent.Tail short, brown above and white underneath. Hairs on the tail and the posterior margins of the hindquarters are longer than on the rest of the body. Steenbok have preorbital glands, pedal glands in all feet, but no inguinal glands. Build and colouration similar in " and !, except that " has short, sharp, straight, smooth and upwardpointing horns (usually 100–150 mm long) that are relatively widely spaced on the head; as in some other bovids, horned !! are known. Juveniles are darker than adults, especially on the rump where there is a greyish tinge, and have longer hair all over the body. Juveniles also have two distinctive black spots on the crown of the head where horns would be expected, and these fade with maturity. Geographic Variation R. c. campestris: southern Africa, including Angola and Zambia. R. c. neumanni: East Africa. Similar Species Raphicerus melanotis. Sympatric in SW Africa. Slightly smaller (ca. 10 kg),

with slightly hunched or arched appearance; grizzled hairs on back, sides and neck; lateral hooves present; " has a preputial gland. R. sharpei. Marginally sympatric especially in Zimbabwe and S Mozambique. Slightly smaller (ca. 10 kg), with shorter legs and more hunched profile; horns shorter (max. 104 mm); pelage grizzled with white flecks. 311

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thorn bushes in areas where vegetation is disturbed. For example, they are often seen feeding along road verges. In southern Africa, they show a particular preference for heavily grazed areas, where the herb layer has a high forb : grass ratio and the woody layer is dominated by encroaching thorn scrub, typically comprising Acacia tortilis and Dichrostachys cinerea. Such conditions often occur around watering points although the availability of drinking water is not a habitat requirement for Steenboks. The key habitat requirement is the availability of high-quality food items (green browse, geophytes, berries, flowers or pods) throughout the year. In the lower Kuiseb River Canyon, Namibia, there was a strong relationship between the distribution of Steenboks and the occurrence of Acacia albida (staple food) and Salvadora persica (shelter) (Cloete & Kok 1986b).

Raphicerus campestris

Sylvicapra grimmia. Broadly sympatric. Larger, with sagittal hair tuft. Ourebia ourebi. Sympatric in parts of range. Larger, with longer neck, and black patch above the tail; inguinal glands present; bare glandular patch below the ear. Distribution Endemic to Africa. Steenboks have a disjunct distribution, with one population in East Africa (S Kenya, N and C Tanzania) and a larger one in southern Africa, the isolating barrier being the tall miombo Brachystegia woodlands of C Zambia, Malawi (from which there are no records) and N Mozambique. Historical Distribution In East Africa, their distribution covers most of S Kenya and N and C Tanzania.They formerly occurred in E Uganda but are now believed extinct. Not recorded from Somalia (Funaioli & Simonetta 1966, Simonetta 1988). In southern Africa, their range extends southwards from S Angola and W and SW Zambia, into most of Namibia (except the arid coastal parts), throughout Botswana and W, C and S Zimbabwe (though naturally absent from the Zambezi Valley below Victoria Falls), S Mozambique (being very common south of the Save R.), to South Africa. In South Africa, they occur almost throughout, being absent from S and SE KwaZulu–Natal and much of the neighbouring Eastern Cape (Ansell 1978, Kingdon 1982, East 1999, Skinner & Chimimba 2005). They are not recorded from Lesotho (Lynch 1994). Current Distribution Their distribution is largely unchanged in southern Africa, but in East Africa Steenboks no longer occur in Uganda, where most of the suitable habitat is now cultivated (East 1999). Habitat Steenboks are found in a variety of habitat types, from Kalahari semi-desert to alpine moorland zones up to 3500 m on Mt Kenya (Young & Evans 1993). However, within any particular ecosystem, Steenboks use a relatively narrow range of habitats with a common factor being the presence of pioneer plants and encroaching

Abundance Steenboks are common throughout their range. Aerial surveys underestimate population numbers, but ground surveys, in areas where the species is common, give density estimates of 0.3–1.0/km2 (East 1999). Based on these estimates, East (1999) estimated a total population size in excess of 600,000, but this clearly is an underestimate. In general, there are no reliable estimates of Steenbok population density, as census methods are too unreliable for this cryptic species. Adaptations The ability to survive without access to drinking water in arid regions might suggest that Steenbok physiology is specifically adapted for advanced water economy, but in fact this is not the case. Steenboks in the Namib Desert were found to have a water turnover rate of 135 ml/day and their rate of faecal water loss was 145 ml/day, while their dietary water intake rate was 343 ml/ day (Cloete & Kok 1986a). The surplus of 63 ml/day would be lost in breathing and evaporative cooling. From kidney structure and function there is no evidence that Steenbok renal efficiency is any higher than average for ruminants, indicating that water balance is maintained largely by behavioural means. These include feeding very selectively on food items of high water content, avoiding heat stress by lying in shade in the heat of the day, restricting diurnal activity mostly to morning and late afternoon, and being active nocturnally. Being small-bodied, Steenboks gain and lose body temperature relatively quickly. They have a low metabolic rate and a high overall thermal conductance (Haim & Skinner 1991). When ambient temperatures exceed about 38 °C, typical heat-shedding behaviour is to stand panting in the shade, with legs spread to dissipate heat from thermal ‘windows’ in the armpits, groin and belly. Their requirement for highly nutritious food items precludes sociality, since such food items are rare and widely dispersed in the environment. If Steenboks foraged in groups there would never be enough food to share at each feeding stop. An implication is that living alone, or sometimes in mixed-sex pairs, precludes the sharing of vigilance costs among herd members. Steenboks do not, in fact, invest more in vigilance, but rely almost entirely on crypsis for predation avoidance (du Toit & Yetman 2005). Steenbok are adapted for vigilance by having relatively large ears and apparently good eyesight. When an approaching predator is detected, the Steenbok will lie motionless in the grass with its ears laid flat and usually the predator will pass by. It is only when a predator approaches within a few metres that the Steenbok will bolt suddenly and at high speed, employing a bouncing hare-like gait and jinking in sharp zig-zags,

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then stopping suddenly to stand stock-still or dropping into cover to lie motionless again. Besides Sharpe’s Grysbok, Steenboks are the only antelope species known to take refuge in burrows, especially those of Aardvarks Orycteropus afer, for predation avoidance and possibly thermoregulation (Smithers 1971, J. T. du Toit pers. obs.). Foraging and Food Although they are frequently referred to in the scientific literature as mixed feeders, largely because of Hofmann & Stewart’s (1972) classification based on stomach structure, Steenboks are predominantly browsers throughout the year. In Kruger N. P., South Africa, both Cohen (1987) and du Toit (1993) found that grass leaves are ingested when they are green and tender after rains but, overall, grass contributes an insignificant amount to the mean monthly diet. These field studies are supported by a review of dietary preferences in African bovids (Gagnon & Chew 2000) and dietary studies involving analysis of stable carbon isotypes that show that Steenboks eat far less grass than is believed (Cerling et al. 2003, Sponheimer et al. 2003b). However, in Zimbabwe, a sample of 91 stomachs from shot animals comprised 30% grass on average (Smithers 1971), and the same author notes that in a sample of 25 stomachs from Botswana, there was an average of 50% browse and 50% grass. This is surprising but indicates that grazing can be important at times. Indeed, in Kenya, Hofmann (1973) noted from stomachs that grass predominated in the diet after rain had stimulated new growth. Steenboks in Kruger N. P. allocate most of their feeding time to forbs during the wet season (>80% feeding time) with the balance being made up mainly of leaves of woody plants, of which Flueggea, Acacia and Ziziphus species are staples (du Toit 1993). The monthly proportional allocation of feeding time to forbs varies closely with the three-month running mean of rainfall, which is an indicator of soil moisture content. During the late dry season in Kruger N. P. (Jul–Oct) forbs make up only about 30% of the diet, with the rest being mainly leaves of woody plants, although the fallen pods of Acacia tortilis are avidly sought out. It appears that the availability of these highly nutritious pods is a key factor enabling Steenboks to remain resident within their territories during the dry season and not to move down-slope to feed along river lines, as most of the other ungulates do during this ‘lean’ period. Also, Steenboks benefit from the micro-climate created within the recumbent canopies of trees felled by elephants, where forbs remain green well into the dry season. Bigger browsers are excluded from this resource by the cage of branches, through which Steenboks are able to creep.

Steenbok Raphicerus campestris.

The mean height above ground at which Steenboks feed varies from 140 mm in the wet season when forbs are the staple, to 390 mm in the late dry season when woody browse predominates in the diet. At full neck stretch, Steenboks cannot browse any higher than about 1 m above ground (du Toit 1990a) and they do not rise up on their hindlegs to gain additional height. Being one of the smallest ruminant species, Steenboks have a narrow dietary tolerance range and thus have to be efficient at maximizing net energetic gains from foraging. Steenboks spend about one-third of their day foraging (feeding and walking in search of food items) in both wet and dry seasons, and the same proportion of the night in the wet season, but in the dry season they spend almost 60% of their night foraging. When not foraging, Steenboks generally lie ruminating or resting to conserve energy (and heat, in winter), and throughout the year they spend about half of their time lying down. Rumination is important for processing ingested food as quickly and thoroughly as possible, and in the wet season this accounts for an additional 26 min for every hour spent foraging, rising to 33 min per foraging hour in the dry season. Steenboks seek out flowers and fruits when they are available, although frugivory appears to be regulated by plant toxins. For example, a radio-collared Steenbok in Kruger N. P. would eat only one Solanum panduraeforme fruit per day, even when they were abundantly available. Geophagia is common. When a Steenbok is seen standing head down near the water’s edge of a pan it may mistakenly be assumed to be drinking, but on close inspection it will invariably be licking the salt crust off dried mud. Steenboks in Kruger N. P. are sometimes seen digging with their front hooves and apparently consuming bulbs or tubers. The only documented observation of this behaviour is by Smithers (1983) in Botswana. Social and Reproductive Behaviour Steenboks are solitary, unless a " is consorting with a !, or a ! has a juvenile with her. Both sexes maintain territories and a defender will drive a conspecific intruder of the same sex out of its territory in an energetic chase, although physical combat is rare. It is unclear what cues are used among !! to assess dominance and submission, since without horns the options for physical combat seem limited. However, in "", opponents kneel on their forelegs and present their horns to one another, with the loser breaking away and running after only a few seconds. This behaviour is most frequently seen when a " is consorting with an oestrous ! and other "" attempt to gain access to her. Scent-marking entails depositing secretions from the preorbital glands on the browsed stumps of forbs and shrubs. Burger et al. (1999a) examined the organic constituents of the preorbital secretion of the Steenbok, and identified 109 different compounds. However, these authors noted that although the secretion is similar to that of the Cape Grysbok Raphicerus melanotis, it is far more complex, with more than 260 different compounds present in the secretion. Steenboks also scent-mark by means of urinating and defecating in latrines. A shallow scrape is prepared with the front hooves, the animal then squats to deposit urine and/or faeces in the scrape and then fresh soil is kicked over the latrine with the front hooves. This behaviour presumably is to keep the deposit moist and maintain the odour for longer than if it was uncovered. The homerange (and presumably also territory) of adult !! in Kruger N. P. 313

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a " : ! sex ratio of 1 : 1.04 in a sample of 4600 adults, and results from Namibia (Stuart 1975) and KwaZulu–Natal (Mentis 1972) based on large sample sizes yielded an even adult sex ratio.

Side and frontal views of male Steenbok Raphicerus campestris head.

covered about 0.6 km2 (du Toit 1993), although it is unknown if "" use more or less space than this. When a ! is in oestrus the attending " will kick and stroke her hindquarters with his forelegs and lick her genital region until she squats to provide a sample of urine, which he tests in typical flehmen posture. During this interaction the ! may be uncooperative and the " may make a loud cat-like growl. Mounting occurs suddenly, with the "’s body held almost vertical, forelegs folded back not clasping the !, and involving a few quick pelvic thrusts before dismounting. Mounting and penetration may be repeated two or three times in quick succession, after which the " and ! typically lie down together resting. The young lies hidden for its first three or four months of life while the mother moves around foraging, but she visits it regularly for suckling (Cohen 1987). When an oestrous ! has offspring with her the consorting " sometimes attacks the juvenile with his horns, which suggests that infanticide could occur, presumably when the attacking " has recently attained dominance, at which time the juvenile cannot be his offspring. If infanticide is a real risk, this probably explains the adaptive value of juveniles having black spots on the head where horns would be expected. An approaching adult " could see these ‘pseudo-horns’ and mistake the juvenile for an attending adult ", at least for long enough to allow the juvenile to move away. Reproduction and Population Structure Females attain reproductive age at about seven months. Steenboks are aseasonal breeders, although examination of 188 female specimens in Zimbabwe suggested a possible birth peak shortly after the onset of the rains (Nov/ Dec) (Wilson & Kerr 1969). A single young is born with a birthweight of 21 cm; Arsenault & OwenSmith 2008). Short grasses are often used, but taller swards are also well utilized. A wide variety of browse species is eaten, with Acacia leaves and twigs commonly found in the diet, along with Combretum spp., Dichrostachys cinerea, Grewia spp., Boscia spp., Maytenus spp. and Commiphora spp. Wilson (1975) listed 28 browse species eaten in Hwange N. P., and Monro (1979) 46 species eaten in Nyslvley. Fallen leaves of mopane may contribute the bulk of their diet during the dry season in the woodlands of southern Africa. Impalas also eat fruit, particularly pods, with those of Acacia tortilis and Acacia nilotica being actively sought for their high protein content. These feeding observations explain why most feeding is at ground level (Dunham 1982, du Toit 1990a). The high browsing pressure on Acacia spp. by Impalas is thought to be responsible for the low regeneration rate of these trees in some ecosystems (Prins & Van der Jeugd 1993, Moe et al. 2009). Joubert (1971) listed 21 browse plants and 12 grasses utilized by Black-faced Impalas in N Namibia. The ratio of dicotyledons to monocotyledons in the diet may also vary with sex and social status (Van Rooyen & Skinner 1989). Territorial !! eat less dicotyledons (31% of diet) than "" (48%) or bachelor !! (49%). This difference between the sexes was also noted at L. Mburo, Uganda (Wronski 1999, 2002) and in a study in Kruger N. P. using stable carbon isotopes (Sponheimer et al. 2003a). It may reflect the fact that dominant !! monopolize habitat where the grass layer is of prime quality, but probably reflects the fact that the time devoted to holding a territory prevents a ! seeking the dispersed, high-quality dicotyledons, as suggested by the greater proportion of dicotyledons in the diet of "" that share the homeranges of dominant !!. Social and Reproductive Behaviour Impalas are gregarious, forming herds of various sizes depending on the season and the

quality of habitat. In Hwange N. P., group size varies from 4 to 150 individuals with the largest herds found in open habitats (mean group size 15.4 in bush/grassland habitats, 12.2 in grassland and 7.3 in bushland; Bourgarel 2004). It is generally during the wet season, or the beginning of the dry season, that the biggest herds occur (Murray 1981). In N Namibia, Black-faced Impala herds range from 3 to 15 individuals, occasionally up to about 20, with aggregations of larger herds formed during or after the birth season (Joubert 1971). Impala social organization consists of territorial adult !! during the rut, bachelor groups and breeding herds. In the Sengwa area, "" seemed to be in separate, stable clans of 30–120 individuals, occupying discrete home-ranges of 80–180 ha depending on the season (Murray 1982a). Bachelor groups comprise juvenile, subadult and adult !! that are potentially territorial. An age-based hierarchy exists within bachelor groups. Highly mobile (in Hwange N. P., some marked subadults and adults moved 30 km from where they were born [M. Bourgarel pers. obs.]), bachelor groups tend to occur in lower-quality habitats where intra-specific competition and disturbance are less (especially during the rut). Cohesion within bachelor herds is poor, particularly when adult !! intending to become territorial become aggressive. After the rut, bachelor !! often mix with breeding herds. Breeding herds comprise adult, subadult and juvenile "", and subadult and juvenile !! that form cohesive herds (average spacing between individuals 285,000, mainly in Central African Republic and S Sudan. Little information is available on its current status in Sudan, although recent survey work conducted in the dry season estimated totals of 1070 and 115 animals for Southern N. P. and Boma N. P., respectively (Fay et al. 2007); the latter is a significant decline from the >50,000 animals estimated in the dry season in 1980 by Fryxell (1980).

Lateral, palatal and dorsal views of skull of Lelwel Hartebeest Alcelaphus buselaphus lelwel.

Lichtenstein’s Hartebeest in the southern clade totals about 82,000, with sizeable populations surviving only in Tanzania, particularly in the Selous ecosystem, and Zambia, in Kafue N. P. and the Luangwa Valley. The other member of the southern clade, the Red Hartebeest, totals about 130,000 and is increasing mainly due to reintroduction for consumptive use such as sport hunting. The overall total for the species at the time of East’s summary was about 360,000 animals. However, this total is strongly influenced by the contribution of the managed populations of Red Hartebeest in southern Africa. It is also a tiny proportion of historical numbers, and conceals the fact that Hartebeest are declining almost everywhere and that this rate of decline is accelerating with the increase in illegal hunting for the bushmeat trade. For example, Sournia & Dupuy (1990) gave the population of Western Hartebeest for the important Niokolo-Koba N. P. in Senegal as over 5000.When surveyed in 1990– 93, the population was ca. 2325, and by 1994–98 it was ca. 1175 (A. Galat-Luong, G. Galat & M. Mbaye pers. comm.). Likewise, in Comoé N. P. in Côte d’Ivoire numbers declined by 60% from 18,300 in 1984 to an estimated 5200 in 1998 (Fischer & Linsenmair 2001a). Densities of Hartebeest, like those of other African ungulates, must be considered with some reservations. Few populations are monitored sufficiently closely to know whether they are at carrying capacity. Many contemporary populations, including those in protected areas, are under severe hunting pressure or affected by fencing. While early explorer accounts were often exaggerated, few areas now seem even remotely close to the picture they gave of ungulate abundance. In addition, Hartebeest have specific habitat preferences and are never found at uniform densities across censused areas. With these caveats, Hartebeest density has been measured in a number of places: Coke’s Hartebeest averaged 2.6/km2 in the Athi/ Kapiti plains in Kenya before the severe 1961 drought and 0.8/km2 after it (Stewart & Zaphiro 1963);Western Hartebeest were recorded at 0.3/km2 in 1990–93 and 0.01/km2 in 1994–98 in Niokolo-Koba N. P. (Galat et al. 1998), 1.01/km2 in Ali N. P., Upper Volta (Green 1979), 0.45/km² in Comoé N. P. (Fischer & Linsenmair 2001a), and 1.66/km² in Bénoué N. P., Cameroon; densities of Lichtenstein’s range from 0.2 to 3.5/km2 (Booth 1985). Adaptations The skull of the Hartebeest is long and thin and the incisor row narrow (ca. 5 cm), probably as an adaptation to selective feeding. Hartebeest crop short green swards and green flushes that follow burning, when they are available, but they are most notably adapted to feeding selectively within medium and tall swards. The thin muzzle is pushed into the sward and is used to select grasses with a high proportion of leaf blade; Hartebeest can even nibble sheath from grasses that have had their leaves removed (Stanley Price 1978a, Murray & Brown 1993). This selectivity can be readily observed as Hartebeest lift their heads to look around while chewing (an anti-predator adaptation), often with the grass selected protruding from the mouth. Perhaps the most remarkable aspect of Hartebeest feeding is the ability to extract high-quality food from senescent swards (Murray & Brown 1993). Where Hartebeest and Roan Antelopes Hippotragus equinus overlap in West Africa, the two species graze on similar plants except in the driest time of year; at this point Roan Antelopes are forced to switch to browse while Hartebeest continue to feed on senescent grasses (Schuette et al. 1998). Hartebeest are thus particularly well adapted to survival in arid grasslands and in dry seasons and droughts. 517

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Lateral, palatal and dorsal views of skull of Lichtenstein’s Hartebeest Alcelaphus buselaphus lichtensteinii.

The digestibility of dry matter and organic matter in the Hartebeest is higher, and the intake lower, than other alcelaphines (Arman & Hopcraft 1975, Stanley Price 1978a, Murray 1993). The combination of an ability to select high-quality components from senescent swards, coupled with low appetite and high digestive efficiency, could be the key suite of adaptations that allowed the Hartebeest to outcompete and replace more generalist alcelaphine precursors. The horns and skulls of all male antelopes are adapted to fighting, but those of the Hartebeest show some unique features.They are also highly variable across Africa, probably reflecting different adaptations to fighting. In general, Hartebeest fight by pushing forward with the hindlegs into a horn ‘clash’, falling down onto their ‘knees’ as they do so. The horns meet with great violence and the ‘bang’ can be heard from hundreds of metres away. The horns then tend to interlock during ‘wrestling’ (see Social and Reproductive Behaviour), a phase of fighting in which animals push and twist their horns, trying to push the head of the opponent to one side. If they succeed, they disengage the horns and stab the face, neck and shoulder of the opponent with a backward hooking movement. Deep horn wounds are common in areas where competition is intense, and population densities high, and sometimes these wounds result in death (Gosling 1975). Horns consist of three main sections: a finely ridged short but thick basal section, a deeply ridged middle section and a smooth, sharply pointed tip. Observation of fighting and wear facets on horns shows that the middle section is used to catch the blow of an opponent, and the sharp tip is for stabbing. The parts are angled so that the horns interlock during ‘wrestling’. This angling has been achieved by rotational twisting and a complete reversal of the horns during a species-specific evolutionary development that is possibly reiterated during ontogenetic growth (Kingdon 1982); variations in this process are partly responsible for the characteristic variation in horn

shape across the different extant (and extinct) forms of Hartebeest. This angling reaches its most extreme in Lichtenstein’s Hartebeest, where the middle-section and tip forms an incurved hook that catches the horns of the opponent and prevents it disengaging. In Coke’s Hartebeest, the middle horn section is horizontal and catches the blow of an opponent above the head. In other Hartebeest, but at its most extreme in Lelwel’s, the middle section is oblique and guides the horn of an opponent into the narrowing ‘V’ shape between the base of the horns, again stopping the blow above the head. In Lichtenstein’s Hartebeest, the basal section is broad and spread over the top of the head to protect it. In all forms, except Lichtenstein’s Hartebeest, the basal sections are set on a bony pedicel that again probably functions to extend the leverage and stop an opponent’s horns well away from the brain. All of these structures are strong and heavy and designed to withstand heavy blows and violent ‘wrestling’. Differences in fighting intensity between different Hartebeest taxa have not been quantified. Hartebeest are renowned for their vigilance and for their speed and endurance. They can run at a fast canter for many kilometres and hunters report that they continue to run even when badly wounded. Presumably these traits were originally anti-predator adaptations in animals that live in open savanna environments (see Predators, Parasites and Diseases). However, their behaviour may also have been shaped by humans. The first Hartebeest appeared less than a million years BP in an area where stone tools made by successive Homo species have been abundant for 2.4 million years. The long dorsal processes probably function to support the powerful shoulder muscles and absorb the shocks of a fast prancing gait, but may also serve to increase the body surface in lateral displays to competitors and mates (see Social and Reproductive Behaviour). Hartebeest thermoregulate by panting rather than sweating. In experiments to measure this response under natural solar radiation, Coke’s Hartebeest started to pant at 32–34 °C; discounting the heat lost by re-radiation (80%) and convection, Hartebeest lose 61% of absorbed and metabolic heat by panting (Finch 1972). When individuals are hydrated, increased panting appeared to be a response to skin temperature but, when dehydrated (15% weight loss), the response was triggered at 39.5 °C by core (rectal) temperature (Finch & Robertshaw 1979). When dehydrated, Hartebeest thus accept an increase in body temperature in order to conserve water. Brain damage is presumably prevented, as in other antelopes, because blood that has been cooled by panting in the nasal mucosa removes heat from the blood entering the brain in the counter-current heat exchanger of the rete mirabile (Taylor 1969). These mechanisms allow a degree of water independence in the dry season. Under natural conditions, some Hartebeest seek shade to reduce exposure to solar radiation (Gosling 1975) and experiments (Finch 1972) confirm the physiological benefits. It is thus interesting that not all Hartebeest seek shade; some rest in the sun and some even stand on termitaria and other mounds in exposed positions. Perhaps this is because of the risks of predation from Lions Panthera leo and Cheetahs Acinonyx jubatus in areas where cover from trees and shrubs provides shade: Hartebeest might thus trade off the costs of thermoregulation against the risks of predation. Foraging and Food Hartebeest feed mainly on grass (Stewart & Stewart 1970) and studies involving stable carbon isotope analysis

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support field observation and faecal analysis in showing that they are almost exclusively grazers (Cerling et al. 2003, Sponheimer et al. 2003b). None the less, Hartebeest do browse, as evidenced by direct observations and the occurrence of browse in stomach contents (Van Zyl 1965, Wilson 1966c, Kok & Opperman 1975, Booth 1985; and see Gagnon & Chew 2000). Hartebeest feeding specializations may be the key to their explosive evolution that occurred at the expense of less-specialized grazing antelopes, notably the Hirola Beatragus hunteri and allies (Kingdon 1982). Hartebeest have long narrow noses and incisor rows that are adapted to feeding selectively within tall grass swards. They are particularly well adapted to taking bites with a high proportion of leaf blade from senescent swards and in feeding trials they rejected poor quality components of the diet at a higher frequency than wildebeest and Topis (Murray 1993). They can also scrape the leaf sheath off the stem in defoliated swards (Stanley Price 1974). Hartebeest cannot increase their bite size in taller swards and their intake rate is lower than that of Common Wildebeest and Topis (Murray 1993). Their low intake is probably because they have a lower metabolic rate than other extant alcelaphines and this, in combination with their ability to select a high quality diet and high digestibility (Arman & Hopcraft 1975, Murray 1993), gives them an important advantage in the dry season, a limiting time for many species. In the wet season they feed less selectively from short green flushes. Seasonal changes in their feeding can follow a classic catenary sequence. In Nairobi N. P. they feed from short green grasses on the catena apex in the rains, then progressively shift to longer, coarser grasses in or near sumps as the dry season advances (Gosling 1975). This may be the basis of their ecotone habit (Lamprey 1973): they are ecologically poised to take advantage of a shift from one preferred food to another. Significantly, when the resource-defence territories of !! are subdivided between two competing !!, the split generally occurs at right-angles to the grass zones within the original territory: each ! thus retains access to the full spectrum of seasonally optimum grass communities (Gosling 1975). A number of ecologists have investigated the basis of possible niche separation between Hartebeest and other grazing ungulates. Bell and others (Gwynne & Bell 1968, Bell 1970) showed that grazing ungulates in Serengeti N. P., including Hartebeest, selected different plant parts from grass swards, and more recent feeding trials (Murray 1993, Murray & Brown 1993) suggest that specialization on particular growth stages of grass may be the fundamental reason why a diversity of grazers can co-exist. These specializations have a spatial dimension as well as a temporal one since tall dry swards can persist for longer in some areas than rapidly growing and more nutritious growth stages. Thus, Hartebeest can remain as residents when other alcelaphines such as wildebeest that are dependent on more ephemeral growth stages are forced to migrate to seek food elsewhere. This specialization also gives Hartebeest the ability to live in relatively arid grasslands and explains the fact that they extended further north, into Ethiopia, Eritrea, North Africa and the Middle East, in contrast to the more restricted distributions of the wildebeest and D. lunatus. However, they do not penetrate truly arid areas such as the Sahara. Their ability to extract a high-quality diet from senescent swards also gives them considerable commercial potential since they can exploit dry swards on a year-round basis more efficiently than any competitors, both alcelaphine or cattle.

The grass species eaten by Hartebeest have been documented in a number of studies using direct observation in the field and analysis of stomach contents and faeces. In Coke’s Hartebeest in Tanzania, Lamprey (1973) found that grass occurred in the diet of Hartebeest at a frequency (96%) greater than in any other large herbivore in the area; 12 grass species were seen being eaten, the most common being Cynodon dactylon, C. plechtostachyu and Cenchrus ciliaris. In studies using faecal analysis in Kenya (Casebeer & Koss 1970, Stewart & Stewart 1971), the most common species detected were Themeda triandra and Ischaemum afrum. Pennisetum mezianum was avoided in the wet season although it was one of a number of grasses eaten at higher frequencies in the late dry season. Red Hartebeest also favour T. triandra, eating it throughout the year; other grasses eaten by this species in N South Africa include Eragrostis spp., Panicum stapfianum, Cynodon dactylon, C. hirsutus and Sporobolus spp. (Kok & Opperman 1975). Red Hartebeest are the only subspecies known to eat significant amounts of browse, but this only occurs in the dry season of particularly dry years (Skinner & Chimimba 2005). Swayne’s Hartebeest appear to select grasses not only on the basis of nutrients, but also on water content and this may explain why they can survive in areas where no apparent free water is available (Mattravers Messana 1993). Daily activity patterns are similar to those of many other species of plains ungulates. At dawn in Kenya, a few Hartebeest are still lying at the end of a resting period. However, most are already grazing in medium or long grass areas and this activity predominates until the temperature rises in mid-morning and animals move to short grass areas where they rest. Early morning grazing may be particularly important for this water-dependent species, because grass is often wet with dew at this time. At the start of resting, often between 10:00 and 11:00h, many animals stand, sometimes in the shade of trees or shrubs and ruminate; as time goes on, progressively more lie down, so that by 13:00–16:00h most are resting in this way. As it cools at around 16:00h animals start to move towards long grass areas, grazing as they do so (Gosling 1975). This basic pattern of activity is similar in Red Hartebeest (Ben Shahar & Fairall 1987). In Coke’s Hartebeest, grazing continues for one to two hours after sunset, when another resting period commences. There is at least one further grazing period before a resting phase just before dawn. However, activity at night needs further investigation. Movement to short grass areas during the day for resting seems likely to be an antipredator adaptation, but the relationship between such movements at night and activity cycles is poorly known. Activity also varies seasonally with more grazing during the middle of the day in the rains (Gosling 1975). Social and Reproductive Behaviour Like most of the plains antelopes, the primary determinant of the Hartebeest mating system appears to be the movements of "" in relation to food and water (Gosling 1986). Females invest most in offspring, particularly during pregnancy and lactation, and their ecology is thus shaped by selection to maximize nutrient intake for reproduction. There is no known post-natal care of their offspring by !!. As noted already, "" have horns, and there is a moderate level of agonistic behaviour between adult "". This sometimes results in fights with violent horn contact and, as a result, a few "" are seen with broken horns. The context of agonistic behaviour between "" has not been studied 519

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but it seems to occur when they come into close contact during drinking, earth-eating or feeding. One animal generally proves to be dominant in such encounters and the role of these dominance relations in mediating the access of "" to limiting resources is a priority for future study (for polygynous antelopes in general, as well as for Hartebeest). Sometimes "" seem to compete where no obvious resource is under dispute. For example, after rain, Red Hartebeest "" lie down and rub their face and neck in wet soil. Females engaged in this behaviour are sometimes approached and chased away by dominant animals (L. M. Gosling pers. obs.). Such behaviour seems linked to the establishment or reinforcement of dominance rather than to competition for a resource. The role of these dominance interactions in competition for access to mates is unknown (as is the existence of choice of mates, below). Females aggregate into groups that are more or less well defined. These groups split and reform as "" move about the landscape and the only consistent associations are between a " and her offspring. This association becomes looser as new calves are born. In Coke’s Hartebeest, up to four successive offspring can sometimes be seen clustered with their mother and typically the distance from the mother to each offspring increases with its age. The wide-scale movements of groups of "" are dictated by differences in the growth and persistence of different grass communities throughout the seasons and by movements during the day between resting and feeding locations. Hartebeest are waterdependent and, in the dry season, movements to permanent water sources become increasingly important. The savanna environment has a regular pattern of vegetation productivity in relation to rainfall and, in general, female movements are quite predictable. As a result, sedentary, ‘sit-and-wait’ mating strategies are a viable option for !! (Gosling 1986). Males compete to defend the areas that most "" visit and establish defended areas known as resource-defence territories; 38% of adult !! occupied territories in a three-year study of Coke’s Hartebeest (Gosling 1975), indicating that an average of up to three !! may potentially compete for each territory. As a result of this competition, territories that attract most "" are smaller and tenure is shorter for their owners; lower-quality territories have fewer "", but they are held for longer periods (Gosling 1974). Territories provide a spatial reference for dominance, their owners being dominant over all other !! while in residence. This becomes particularly important during mating and, as far as is known, only territorial !! (or dominant !! that follow groups of "", see below) mate. When !! leave their territories, for example to drink, they are subordinate to !! resident in territories that they pass through. Males temporarily outside their territories are generally dominant over non-territorial !!, although not always. The behavioural state of dominance in territorial !! thus appears to be under central nervous system control and can be turned on and off as they cross the boundary of their territory. While Hartebeest do use the preorbital glands to some extent for marking objects in their territories, territories are scent-marked mainly by dung piles and these are particularly large at boundaries where resident !! interact with neighbouring territorial !!; the interdigital secretion also probably plays a role in demarcation of territories (Reiter et al. 2003). Neighbours meet regularly in agonistic encounters, particularly where territories are relatively small. These encounters consist of much grooming (possibly to

avoid head-on orientations) and are marked by head-high postures with facing away. The focus of most encounters is ritualized scentmarking. Males sniff a dung pile, paw it, kneel, rub their head and horns, stand and defecate. The opponent then moves to the same position, noses the faecal pellets left by the preceding ! and repeats the actions. Repetitions of these sequences can go on for some time, sometimes with vigorous pawing that scatters the faeces and results in a depression at regularly used sites. In general, Hartebeest do not lie down in their dung piles during boundary encounters (in contrast to wildebeest) although they do lie on their own dung piles when resting (possibly a form of self-anointing with the odour used to mark their territory; Gosling 1982). Hartebeest break shrubs and dwarf trees such as Acacia drepanolobium with their horns then rub the surface of the preorbital gland on the end of broken branches; such marking sites are generally 1–3 cm in diameter (Gosling 1975). Males also ‘self-anoint’ by rubbing the secretion of the preorbital gland on their own sides leaving a distinct mark in the case of Lichtenstein’s Hartebeest (Dowsett 1966). This area is sniffed and sometimes nibbled by opponents during agonistic encounters and these animals may be assessing their opponent (as a territory owner) by matching the smell on the pelage with that of scent marks in the territory (Gosling 1982). In the case of Lichtenstein’s Hartebeest in Zimbabwe, self-marking increases in Sep just before the rut (Booth 1985); however, this behaviour occurs in all age and sex classes, which is curious and needs further investigation. Booth (1985) saw a territorial ! Lichtenstein’s Hartebeest marking a " on its rump just before mounting. Horn contact behaviour usually starts with ‘horn-tangling’ (light butting and twisting against the opponent’s horns). Serious fighting is generally absent during boundary encounters and also from most interactions with intruders. It occurs principally to determine ownership of a territory and at such times !! fight with great violence and are sometimes killed (see Adaptations); Dowsett (1966) reported one fight lasting more than an hour. Inter-individual body orientation is also very important in interactions between male Hartebeest. During encounters !! often orientate in the parallel or reverse parallel orientation. Encounters end with the !! gradually and reciprocally turning the head away after a period in the forward parallel orientation and eventually moving directly away from each other. They often graze at this point. Some interactions even consist almost entirely of two !! grazing next to each other (‘grazing encounters’) with subtle and reciprocated changes in body orientation. Agonistic behaviour with intruding !! varies according to the dominance status of the intruder. Encounters with high status !! can contain many of the elements that occur between neighbouring territorial !!, including escalation to horn contact. In rare cases, such interactions can escalate to full-blooded fights that become takeover attempts. However, the great majority of encounters with non-territorial !! end in the early flight of the intruder. Sometimes intruders simply turn and run as an owner approaches. Sometimes owners stand with the head erect and facing away. The intruder may then walk away with a lowered head. However, quite often the intruder approaches the owner and sniffs its cheek and neck, sometimes even nibbling down its neck. This behaviour (‘neck-sliding’) may function to smell the odour of the owner and test if it is the same animal that made the scent marks (dung piles) in the territory (the ‘scent-

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Hartebeest Alcelaphus buselaphus.

matching’ hypothesis; Gosling 1982). Sometimes owners overtake the intruder as it walks or runs away and, turning in front of the retreating animal, perform a characteristic bucking lateral display: the intruder responds by a low head toss and accelerated withdrawal. The bald fact that most breeding Hartebeest !! occupy resource-defence territories conceals a great deal of variation. In low and moderate population densities, territories are often large. For example, Dowsett (1966) recorded territories of 1.55–5.2 km2 for Lichtenstein’s Hartebeest in Zambia and Backhaus (1959) observed the behaviour of a Lelwel Hartebeest ! in a territory of at least 3 km2. In both of these cases, small groups of "" remained for long periods within one territory; in Dowsett’s study there were 1–9 "" in a sample of 11 such groups, while there were three "" in the group observed by Backhaus. In Uganda (Kidepo), several Lelwel "" were generally present in each male’s territory (4–10 km2), but this female distribution pattern was very variable according to season and year due to massive emigration and immigration (J. Kingdon pers. comm.). At higher population densities, territories are much smaller. In a high-density population in Kenya, Coke’s Hartebeest territories averaged 0.31 km2 in a sample of 73 territories observed

over three years; territories in preferred ecotone habitats that attracted most "" were smaller than those in scrubland territories (Gosling 1974). In Nairobi N. P., at the time of these observations, "" had home-ranges of ca. 5 km2 and these included 20–30 male territories. Female ranges may thus include from one (or a small number of territories) to over 30, a fact that must affect the potential for mate choice. Some !! occur in quite dense clusters, particularly around short grass clearings (Gosling 1974: Coke’s Hartebeest; Mattravers Messana 1993: Swayne’s Hartebeest). In such cases !! rest on the short grass areas and also graze them intensively when green flushes occur after rainfall. However, Hartebeest are classic edge species and they graze into long grass areas in the evenings, and increasingly as the dry season progresses. These clusters of resource territories are sometimes so dense that it has been suspected that they might have similarities to a ‘lek’, an arena where clustered !! display and "" choose mates. However, the quality of the food supply is always critically important in attracting "" to such territories, and thus in the mating success of the male owners. Thus, while small, such territories should probably be regarded as resource territories; they 521

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are not lek territories in the sense of the tiny territories on Topi leks. However, the clusters of territories found in Coke’s and Swayne’s Hartebeest would make it easier for "" to choose between territorial !!; but variation in the degree of mating skew has not been measured. Perhaps the best evidence for the role of sexual selection on male characteristics is sexual dimorphism in fighting structures (horns, pedicel and skull robustness). Not only do the sexes invest to a different extent in these traits, but the degree of difference between !! and "" varies across subspecies. This variation appears to be mainly a response to differences in the length of the breeding season. Species with shorter breeding seasons and thus a higher polgyny potential are relatively more dimorphic in horn dimensions, pedicel height and skull weight (Capellini & Gosling 2006). Sexual selection might also have favoured the occurrence of deep colouration together with other conspicuous patterns in the pelage of Red and Swayne’s Hartebeest. There are indications that competition might be intense in these two subspecies with breeding restricted to a short annual breeding season in Red Hartebeest and restricted spatially in Swayne’s Hartebeest in the territory clusters described by Mattravers Messana (1993). There may be enhanced male intra-sexual competition in both cases and, possibly, enhanced opportunities for female choice. However, more work is needed to fully explain variation in coat colouration, including the absence of conspicuous colouration in the two woodland subspecies, the Lichtenstein’s Hartebeest and Western Hartebeest. However, Hartebeest do show flexibility in male mating tactics: when they are at very low densities breeding !! follow groups of "" around their entire home-range. A well-authenticated case occurred in Amboseli N. P. where an individually known ! accompanied a group of "" throughout an 11 km2 area (D.Western pers. comm.). Similar behaviour appeared to occur in Masai Mara National Reserve, where L. M. Gosling (pers. obs.) observed another low-density population in the late 1980s. ‘Following’ strategies may arise when resources are very unpredictable so that a ! is forced to follow "" in order to ensure that he is with them when they become receptive. However, neither Amboseli nor the Mara were particularly unpredictable habitats and it is more likely that a low density of "" and reduction in the level of competition between !! was critical in the appearance of following tactics. Males in resource-defence territories should generally win against following !! because of the benefits of owner advantage in a small scentmarked territory. Thus, ‘following’ can probably only succeed when there is a very low density of male competitors. When "" enter a territory, !! stand at the boundary, circle behind them as they pass by and nose the vulva. Sometimes "" respond by urinating and !! sniff the urine as it falls and on the ground. However, remarkably (in view of its widespread occurrence amongst ungulates), Hartebeest do not show the oestrus testing display known as flehmen (a trait shared with Damaliscus). If "" are in oestrus, mating behaviour becomes prolonged. Again, body orientation is important: !! often stand in front of "" or even walk away from them to look intently at non-existent objects away from the ". These behaviours appear to be a subtle attempt to threaten the " into stopping as she walks away but not to alarm her so that she starts to run. When near to the " the ! employs the ‘ear-down’ posture with nose raised, ears down and the tail stiffly curved (Gosling

1974). In general, a " can escape a male’s attentions if she runs away quickly but if she is in oestrus, the ! will gallop hard to intercept her and bring her back, even if the " has run into the neighbouring territory. When mating is in progress, neighbouring !! sometimes run into the territory and gallop at full speed through the group, trying to scatter it. This tactic presumably increases the chance that the oestrous " will be displaced into its own territory. Copulation occurs repeatedly with the male’s forelegs just in front of the female’s haunches and the head dropped vertically over the female’s shoulders. Most copulation in Coke’s Hartebeest appears to be around mid-day, perhaps as an anti-predator adaptation since mating animals must be very vulnerable; Hartebeest are generally resting in short grass areas at mid-day and predators are usually inactive. Parturient "" usually isolate themselves and retire to scrubland to give birth. Female Coke’s Hartebeest eat the afterbirth and the calf stands about 30 min after birth (Gosling 1969a), considerably slower than wildebeest, which are more vulnerable in an open habitat. Young hide in long vegetation for about two weeks after birth (lying-out), emerging only to be suckled and for their mother to consume their urine and faeces; presumably this removes odours that might attract mammalian predators. Such hiding behaviour also occurs in Lichtenstein’s Hartebeest (Mitchell 1965, Ansell 1970) and Red Hartebeest (Kok 1975). Variation in development of the young and lying out behaviour require more detailed study. It might be predicted that Hartebeest with a more synchronized calving period, such as Red Hartebeest, would stand more quickly after birth, if, as is believed, such calving distributions are shaped by predation pressure. Males that fail to occupy resource territories live in groups. Males join male groups from about ten months of age up to two or three years of age. This timing depends partly on when they are separated from their mothers and thus, to some extent, on when the mother has further offspring. The mother keeps previous offspring further away from a new calf and, in the case of !!, this makes the offspring more exposed to aggression from the territorial !. Further, when a young ! is chased away by a territorial !, the mother is more likely to intervene and to flee with it if the ! is her only offspring. Once young !! are separated from their mothers, territorial !! chase them with exceptional severity and try to horn them as they flee. As a result young !! are sometimes found isolated and are sometimes injured. When young !! join male groups they may also be chased by adult members of the group; as a result they sometimes cluster with other young !! in subgroups at the edge of groups of older !!. Such ‘male groups’ (sometimes called bachelor herds) comprise about 62% of the adult male population in Coke’s Hartebeest (Gosling 1975). They occupy areas outside territories or, more usually, littleused parts of territories. It is not economically possible for territorial !! to keep all parts of their large territories free of non-territorial !!. Rather, they tend to chase any intruder that comes too close to them, wherever they might be in their territory. Active defence is also more likely during cooler times of the day. References in the literature to male groups defending territories are certainly an error: single high-status non-territorial !! may sometimes establish a small temporary territory, but groups never do so. Groups of nonterritorial !! vary in size from one to over a hundred individuals. Single non-territorial !! are common (particularly when high status !! isolate to try to find a vacant territory) and so it is impossible

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to tell if a lone adult ! is territorial or not without observing its behaviour, preferably for a number of days. By far the most common social behaviour in male groups is agonistic behaviour and !! form dominance relationships. When they reach high dominance status they leave the groups and wander about looking for vacant territories. If they have previously held a territory they return to the same area and sometimes succeed in taking it back, or in taking over an adjacent territory. Sometimes they occupy a territory when the owner is temporarily away, for example to drink, and an ownership contest occurs when the owner returns. Fights for territory ownership are by far the most severe type of contest. They consist of repeated violent clashes and often result in severe injury and sometimes in death. When a ! is defeated it may be pursued for several kilometres. There are few vocalizations.The most distinctive is the ‘quack’-like sound made by young calves. Significantly, the same call is produced by half-grown or young adult !! when they show the subordinate ‘head-in’ posture as they move away or flee from approaches or aggression by a territorial ! (Gosling 1974). The alarm snort is used in anti-predator contexts (below). Reproduction and Population Structure The seasonal distribution of reproduction varies between different parts of Africa and appears to be adapted to seasonal patterns of rainfall/primary production. In Nairobi N. P., where there are two seasons of rains, there are two peaks of calving by Coke’s Hartebeest. These peaks occur before the two seasons of rain so that the late phases of lactation, when "" need most food, coincide with the production of fresh green grass. This grass is also available as the young start to wean and it is not clear which of these two events is most important for this timing. Where there is only one season of rain there is only one breeding season.Thus, Lichtenstein’s Hartebeest calve from Jul– Sep in Zambia, Mozambique and Zimbabwe (Ansell 1960a, Mitchell 1965, Wilson 1966c, Booth 1985), although Ansell (1960a) gives a season of Oct–Nov for the low-lying Luangwa Valley. In Namibia, there is a corresponding unimodal pattern of calving for Red Hartebeest with a peak in Nov and Dec, a month or two before the start of the rains (L. M. Gosling pers. obs.). In Coke’s Hartebeest, the seasons of calving are quite widely spread, reflecting the distribution of rainfall (Gosling 1969a). Thus, it seems likely that selection for the timing of mating reflects the timing of water-dependent primary production. However, in the case of Red Hartebeest in Namibia the calving peak is considerably sharper than that of rainfall. The general reason for sharp calving peaks is probably that young born outside the peak are more likely to be killed by predators so that natural selection favours calving within the peak (Estes 1966, Gosling 1969a). Coke’s Hartebeest may be free of this constraint because young hide alone in long grass for the first two weeks of life (Gosling 1969a) and thus rely on crypsis rather than on saturating predation pressure. Under such circumstances, there could even be advantages in producing young when there are few others about to avoid the formation of ‘prey images’ by predators such as baboons and hyaenas. It is not entirely clear why Red and Lichtenstein’s Hartebeest have such sharp calving peaks, but perhaps young rely to a lesser extent on crypsis; like Coke’s Hartebeest, Lichtenstein’s Hartebeest calves hide after birth (Ansell 1970), but duration and variation have not been quantified.

1 month

6 months

6 months

9 months

1 year

9 months

2 years

1 year

young adult

2 years

prime adult male

very old

TOP: Coke’s Hartebeest Alcelaphus buselaphus cokii horn development (from Gosling 1975) ABOVE: Lelwel Hartebeest A. b. lelwel horn development (courtesy of J. Bindernagel pers. comm.).

Gestation is about eight months (Skinner et al. 1973). A single calf is born weighing about 15 kg (Wilson 1966c). Calves are weaned at about 7–8 months of age (Kok 1975). Most Hartebeest "" probably conceive for the first time between 15 months and two years old, although this may depend on condition due to variation in grassland production. Lichtenstein’s Hartebeest "" are sexually mature at 16–18 months (Mitchell 1965, Wilson 1966c) and Red Hartebeest at 28 months (Skinner et al. 1973). Physiological maturity in !! may also be reached at about 18 months; !! of around this age have been seen crouching and ejaculating after sniffing or attempting to mount "" (Gosling 1975). However, !! do not mate until they have acquired a territory, probably not before three years of age, at least in moderate- to high-density populations. It is possible that territories are acquired earlier in low-density populations where male intra-sexual competition is lower, but there are no data. Like most of the plains antelopes, Hartebeest generally have an adult sex ratio biased towards "". In Coke’s Hartebeest, while the ratio at birth does not differ from unity !! comprise only 42% of adults (over 20 months old). The main cause of this difference is mortality indirectly caused by male competition. In particular, young !! chased away from their mothers by territorial !!, and old !!, excluded from high-quality territories, are often injured and isolated in scrubland. Such animals are vulnerable to predation, particularly by Lions (Gosling 1974). A life-table, based on a sample of aged skulls from natural mortality in a population of Coke’s Hartebeest, shows a typical mammalian pattern with high mortality over the first two years of life, relatively low annual mortality up to six years, and then accelerating mortality up to 12 years; "" outnumber !! in the oldest year classes and some probably live up to about 15 years (Gosling 1974, 1975).Weigl (2005) gives longevity record as 22–23 years in captivity, and Flower (1931) reports 19 years for a Bubal Hartebeest. Predators, Parasites and Diseases Lions are the main predators of adult Hartebeest (Mitchell et al. 1965, Schaller 1972, Gosling 523

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1975). Adults are occasionally killed by Cheetahs, although the only case confirmed in a long-term study of Coke’s Hartebeest was an instance of four Cheetah !! hunting together (L. M. Gosling pers. obs.); Cheetahs do, however, kill large numbers of calves. Leopards Panthera pardus are known to kill calves in Serengeti N. P. (Bertram 1979) and probably kill small numbers throughout the species’ range. The alert response of adult Hartebeest "" to jackals Canis spp. and baboons Papio spp. suggest that they kill very young calves. Adult and juvenile Hartebeest are also hunted and killed by Spotted Hyaenas Crocuta crocuta (Kruuk 1972, Mills 1990, Di Silvestre et al. 2000) and African Wild Dogs Lycaon pictus (Creel & Creel 1995). Predation on Hartebeest may be affected by the abundance of alternative prey. In Serengeti N. P., when migratory ungulates, such as Common Wildebeest and Plains Zebras Equus quagga, move into an area, Lions feed mainly on the migrants and pressure on resident species, including Hartebeest, is temporarily reduced (Bertram 1979). Hartebeest are extremely vigilant and respond to predators, particularly Lions, at great distances. When alarmed, Hartebeest face the predator directly with the head at maximum elevation, the nose drawn in and the ears directed forwards. They seek any rise in the ground to obtain a better view. At intervals related to the apparent level of danger, they make characteristic loud snorts by rapidly forcing air through the nostrils. They may approach and even follow a moving predator such as a Lion or Cheetah to keep it in sight and thus under sensory control. Hartebeest respond sensitively to the alarm behaviour of other nearby animals. Gosling (1975) observed an isolated territorial ! in scrubland respond to its neighbour’s alarm snorting by snorting itself and adopting the same body orientation, even though it could not see the predator (a Lion) that the first animal had detected. The main function of belonging to groups is probably due to the benefits of selfish-herd membership (detection and dilution effects). When disturbed by predators, lone territorial !! in scrubland move closer to each other. Members of other species, for example wildebeest, sometimes take advantage of the extreme vigilance of Hartebeest by seeking refuge within Hartebeest groups when predators are detected. Flight distances appear to be related to vulnerability. In a group watching a predator, "" with young calves keep to the rear of the group and are usually the first to flee, running straight away from the back of the group. Active defence against predators is rare although young Hartebeest !! have been seen attacking Black-backed Jackals Canis mesomales after the jackals had killed a young gazelle (L. M. Gosling pers. obs.). Hartebeest carry a large number of disease organisms, but there are few records of them showing any clinical symptoms of disease. Large-scale mortality may instead be generally due to starvation when food resources are exhausted within range of permanent water; examples include the die-off of Coke’s Hartebeest in 1973 in Nairobi N. P. in Kenya (Hillman & Hillman 1977) and of Red Hartebeest in the Kalahari in 1985 (Knight 1995a). Hartebeest generally have low susceptibility to rinderpest virus infection (Plowright 1982). For example, in the great pandemic of 1889–97, while Damaliscus spp. amongst other antelopes died in large numbers, Red Hartebeest were relatively unaffected (Scott 1970). However, sometimes Hartebeest were severely affected by rinderpest, as in the case of Swayne’s Hartebeest in Somalia in 1897 (Simon 1962) and the Western Hartebeest in 1913–17 (Pecaud 1924, in Plowright 1982). Hartebeest typically live at lower

density than the most susceptible species (African Buffalo Syncerus caffer and wildebeest) and it would be interesting to know how far density explains this variation: Swayne’s Hartebeest were known to reach high densities on open grassland before their numbers were reduced by overhunting. Antibodies have also been detected against bluetongue virus in Western Hartebeest (Formenty et al. 1994) and Red Hartebeest (Simpson 1978), against bovine ephemeral fever in Coke’s Hartebeest (Devies et al. 1975) and bovine virus diarrhoea in Lichtenstein’s Hartebeest (Anderson & Rowe 1998). Alcelaphine herpes virus-2 has been identified in a number of subspecies; it is closely related to alcelaphine herpes virus-1, which is carried by wildebeest and causes malignant catarrhal fever (Reid et al. 1975, Seal et al. 1989) but has not been linked to clinical MCF in Hartebeest. Hartebeest have relatively low susceptibility to the anthrax bacillus, but they do die in moderate numbers when infected; examples are the 1999 and 2000 outbreaks in Mago N. P., Ethiopia, involving Lelwel Hartebeest (Shiferaw et al. 2002). The haemoprotozoan Theileria sp. has been isolated from the serum and from ticks carried by Red Hartebeest (Spitalska et al. 2005), but there were no clinical signs of theileriosis. Trypanosoma brucei, the protozoan parasite that causes African sleeping sickness, has been detected in Western (Jamonneau et al. 2003) and Coke’s (Geigy & Kaufmann 1973) Hartebeest. However, Hartebeest are not favoured by tsetse flies (Glossina spp.) and were not detected as hosts in a sample of 13,145 blood meals collected throughout Africa (Clausen et al. 1998). Most, perhaps all, adult Hartebeest carry a moderate number of ticks, including Amblyomma spp., Boophilus spp., Haemaphysalis aciculifer, Hyalomma truncatum and Rhipicephalus spp. (Hoogstraal 1956, Matthysse & Colbo 1987, Walker et al. 2000, Ntiamoa-Baidu et al. 2005). These are vectors of a number of viral diseases (see below and an overview in Walker et al. 2003).Ticks sometimes occur in clusters, notably around the edges of the ears. Records of the infestation of various wild bovid hosts suggest that Hartebeest are remarkably free of ticks compared with such species as Impala Aepyceros melampus, wildebeest and African Buffalo (Walker et al. 2000). This finding is supported by studies of the numbers of ticks per individual in relation to body size, which show that Hartebeest carry less ticks and have a smaller proportion of engorged ticks than would be expected (Olubayo et al. 1993). This may be linked to effective grooming with the incisors and hindfeet and also to their relatively open habitat. Oxpeckers Buphagus spp. rarely feed on Hartebeest. In a study of Coke’s Hartebeest, oxpeckers sometimes attempted to land, but were nearly always immediately driven away (L. M. Gosling pers. obs.); in another study that included Coke’s Hartebeest, none was seen feeding (Koenig 1997). The low incidence of ticks means that Hartebeest are probably little affected by tick parasitosis (direct damage due to wounds and blood loss). Hartebeest are sometimes infected by Sarcoptes scabiei, the mite causing sarcoptic mange (Pence & Uekermann 2002). Gastrointestinal parasites of Hartebeest include helminths and nematodes (e.g. Bindernagel 1972, Boomker et al. 2000). Nematode faecal egg counts of Hartebeest in N Kenya were higher in drought years than non-drought years (Ezenwa 2004b), possibly due to nutritional stress. Most adult Hartebeest appear to be infested with nasal botfly larvae, including those of Oestrus ovis (Howard 1977, Wetzel 1984, Mbassa 1986). These flies lay eggs in the nose and the larvae migrate upwards into the nasal cavity. L. M. Gosling (pers. obs.) once saw

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a Coke’s Hartebeest ! in Nairobi N. P. sneeze out a larvae and on inspection it proved to be O. ovis. Some individuals contain large numbers of these larvae and in necropsy of a Red Hartebeest ! in Namibia, L. M. Gosling (pers. obs.) was able to confirm that these penetrate even into the cavity of the pedicel. It is sometimes said that the larvae penetrate the brain, but evidence is lacking. Hartebeest sometimes sneeze unexpectedly and this could be a response to botfly larvae; the alarm snort is similar but is accompanied by alert behaviour, focused on a predator. Conservation IUCN Category: Least Concern (A. b. buselaphus – Extinct; A. b. cokii – Least Concern; A. b. lelwel – Endangered A2acd; A. b. swaynei – Endangered C2a(i); A. b. tora – Critically Endangered C2a(i); A. b. major – Near Threatened; A. b. lichtensteinii – Least Concern; A. b. caama – Least Concern). CITES: Not listed. The feeding habits of Hartebeest bring them into direct conflict with grass-eating livestock and, as numbers of livestock have increased, Hartebeest numbers have declined everywhere except in a few protected areas. Hartebeest also are valued for their highquality meat and as the bushmeat trade escalates out of control, partly fuelled by the increase in modern guns, many Hartebeest populations are being hunted to extinction. Hartebeest are thus declining in most parts of their range and are now rare outside protected areas, many of which are too small to support viable populations. Protected areas holding important populations of Hartebeest include: Niokolo-Koba N. P. (Senegal), although this population declined by half in the 1990s alone (Galat et al. 1998); Comoé N. P. (Côte d’Ivoire), Mole and Digya National Parks (Ghana) and Pendjari N. P. (Benin) for Western Hartebeest; Zakouma N. P. (Chad), Manovo–Gounda–St Floris N. P. (Central African Republic), Southern N. P. (Sudan), Mago N. P. and surrounds (Ethiopia) and Murchison Falls N. P. (Uganda) for Lelwel Hartebeest; Mazie N. P. and Senkelle Wildlife Sanctuary (Ethiopia) for Swayne’s Hartebeest; Tsavo N. P. and Masai Mara National Reserve (Kenya) and Serengeti and Tarangire National Parks (Tanzania) for Coke’s Hartebeest; the Selous ecosystem and Ruaha–Rungwa–Kisigo complex (Tanzania), Kafue N. P. and Luangwa Valley (Zambia), and Niassa G. R. (Mozambique) for Lichtenstein’s Hartebeest; and, for Red Hartebeest, Kgalagadi Transfrontier Park (South Africa/ Botswana), Etosha N. P. (Namibia) (East 1999) as well as various nature reserves and conservancies, such as the expanding population of Red Hartebeest of about 18,000 animals in the 400,000 ha Seeis Conservancy in E Namibia (H. Förster & B. Förster pers. comm.). The failure to protect Hartebeest adequately is particularly tragic because there is poor recognition of local variation among Hartebeest populations and even less recognition of their evolutionary importance as the outstanding model of a mammalian adaptive radiation in the savanna environment. Many local variants have disappeared (buselaphus, nakurae). Hartebeest were kept in captivity by the ancient Egyptians, illustrated in the tombs of the pharaohs, and used for ceremonial purposes. However, the wild populations from which these animals were taken are now extinct, perhaps partly because of habitat destruction in the Mediterranean region, but mainly due to hunting. Bubal Hartebeest in North Africa were similarly reduced and probably died out around the 1950s (see Distribution); groups in zoos (including London and Paris) that could have ensured their survival were allowed to die out. The intergrade populations of the Kenya Rift

Valley were displaced by intensive farming and shot by settlers and troops; the last surviving Nakuru Hartebeest (a remnant of a once abundant intergrade population between A. b. cokii and A. b. lelwel in the Kenya Rift Valley), a !, was seen and photographed in 1967 (Gosling 1969b). Other subspecies and intergrades are severely threatened or may already have disappeared (swaynei, jacksoni, keniae, tora); there are now less than 800 Swayne’s Hartebeest (Antonínová et al. 2008) and Tora Hartebeest have not been seen for some 10–15 years in any part of their range. The Hartebeest are the most important example of an antelope adaptive radiation in which most members still just survive; they illustrate better than any other group an adaptive radiation into varied savanna habitat and the evolutionary consequences of long-term climatic change. They thus represent an opportunity to conserve the spectacular manifestation of an evolutionary process rather than a museum collection of isolated taxa in protected areas. There is also a case for conservation at a local level. For example, the declining Kenya Hartebeest is a unique local race and should be a conservation priority in a country that values its wildlife and depends on it economically. An important exception to the pattern of decline is the Red Hartebeest in southern Africa, which is important for various forms of sustained use such as trophy hunting. The latter case is a remarkable example of the success of the southern African approach to wildlife conservation, which involves the transfer of ownership of wildlife and thus any financial profit from its use to the landowner. So long as such practices continue on a rational basis, Red Hartebeest populations will remain secure; indeed its numbers are expanding. Lichtenstein’s Hartebeest also seems to be relatively secure, partly because there are several very large reserves within their range (which is tsetse country) and perhaps because this species is exploited to a lesser extent. The lesson from population changes between Hartebeest taxa across Africa is that populations with consumptive use are stable or expanding while the others are plunging to shortor medium-term extinction. Future conservation measures should attempt to expand and improve the operation and sustainability of sustainable use schemes. However, they should also be combined with non-consumptive exploitation in properly guarded protected areas. A further obstacle to be overcome is that unless income from consumptive and non-consumptive use is shared by local communities, Hartebeest will not survive in the medium term.Their meat is highly regarded and they will be poached to extinction except where tangible economic benefits for entire communities outweigh the individual’s short-term need or taste for food. The expanding Hartebeest populations in southern Africa are descended from the small number of survivors of the slaughter of the seventeenth to nineteenth centuries by European colonists. Hartebeest were eradicated from most of their range (Skead 1980, 1987) and much genetic variation was undoubtedly lost at this time. For example, even though its taxonomic status is uncertain, the Hartebeest known as the Cape Red Hartebeest (and believed at the time to be a subspecies) was completely exterminated by hunting and only those known as Northern Red Hartebeest survived around the Kalahari and in what is now known as Namibia (Harper 1945). However, the survivors are numerous again and expanding as the considerable economic potential of the species is appreciated and exploited. The main conservation problem in southern Africa is that animals are sold and translocated to new areas without regard to natural spatial patterns of genetic variation. Local legislation needs to be revised, if only because populations are 525

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most likely to thrive if introduced into areas to which they are locally adapted, that is, their natural range. There is an urgent need for an Africa-wide conservation strategy for the Hartebeest, one that transcends local custom and makes use of best-practice across the continent. Hopefully, before it is too late, conservationists at least will become aware of the impending loss of the finest existing example of a pan-African large mammal radiation. Measurements Alcelaphus buselaphus A. b. caama TL (!!): 2144 (2073–2200) mm, n = 8 TL (""): 2096 (2070–2110) mm, n = 3 T (!!): 470 (404–504) mm, n = 8 T (""): 472 (430–500) mm, n = 3 HF c.u. (!!): 557 (534–572) mm, n = 8 HF c.u. (""): 524 (503–545) mm, n = 3 E (!!): 195 (192–201) mm, n = 8 E (""): 185 (175–192) mm, n = 3 WT (!!): 152.0 (137.0–156.0) kg, n = 8 WT (""): 120.0 (105.0–136.0) kg, n = 3 Botswana (Smithers 1971)

caama22 caama1 caama2

southern lineage

licht1 licht2 cokii1 swaynei10 lelwel2 cokii46 lelwel3

eastern lineage

lelwel4

northern lineage

swaynei1 swaynei4 tora5 major6 major7

western lineage

Tentative phylogenetic tree for Alcelaphus as estimated from cytochrome b data (after Flagstad et al. 2001).

A. b. lichtensteinii TL (!!): 2380 (2090–2540) mm, n = 6 TL (""): 2360 (2010–2420) mm, n = 5 Sh. ht (!!): 1230 (1220–1360) mm, n = 6 Sh. ht (""): 1250 (1190–1300) mm, n = 5 WT (!!): 177.1 (156.7–203.9) kg, n = 10 WT (""): 166.3 (160.4–181.2) kg, n = 10 Zambia (Wilson 1966c) A. b. cokii TL (!!): 2327 (2210–2460) mm, n = 5 TL (""): 2205 (2105–2275) mm, n = 5 T (!!): 515 (490–545) mm, n = 5 T (""): 497 (460–515) mm, n = 5 HF c.u. (!!): 505 (500–520) mm, n = 5 HF c.u. (""): 492 (480–500) mm, n = 5 E (!!): 201 (190–213) mm, n = 5 E (""): 186 (180–195) mm, n = 5 Sh. ht (!!): 1170 (1130–1180) mm, n = 5 Sh. ht (""): 1120 (1110–1160) mm, n = 5 WT (!!): 142.5 (129.0–159.8) kg, n = 5 WT (""): 126.2 (116.0–135.0) kg, n = 5 Serengeti N. P., Tanzania (Sachs 1967) Maximum recorded horn lengths for the subspecies are: 74.9 cm for A. b. caama for a pair of horns from Windhoek, Namibia; 61.9 cm length Lelwel Hartebeest Alcelaphus buselaphus lelwel female (top) and male (bottom). for A. b. lichtensteinii for a pair of horns from Mumbwa, Zambia; 61.0 cm for A. b. cokii for a pair of horns from Kenya; 73.0 cm for A. b. major Key References Dowsett 1966; Gosling 1969a, 1975, 1986; for a pair of horns from Nigeria; 70.1 cm for A. b. lelwel from the Aouk Kok 1975; Mattravers Messana 1993; Wilson 1966c. R., Chad; 58.1 cm for A. b. tora from the Sudan; and 51.4 cm for A. b. swaynei from Somalia L. Morris Gosling & Isabella Capellini 526

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GENUS Connochaetes Wildebeest Connochaetes Lichtenstein, 1812. Mag. Ges. Naturf. Fr. Berlin 6: 152.

Common Wildebeest Connochaetes taurinus myology.

Common Wildebeest Connochaetes taurinus skeleton.

There are two recognized species, the CommonWildebeest Connochaetes taurinus and the Black Wildebeest or White-tailed Gnu C. gnou.While the latter is monotypic, there are five distinctive subspecies of C. taurinus, at least one of which (C. t. mearnsi) may be different enough to be considered a separate species (Georgiadis 1995). Black Wildebeest ranged the temperate Highveld and Karoo of South Africa, from where C. gnou antiquus was described. Connochaetes taurinus is known from fossil remains in North and East Africa, and from lower, middle, and upper Pleistocene beds in South Africa. According to mtDNA analyses (Arctander et al. 1999), the Common Wildebeest shows a pattern of colonization going from southern Africa toward East Africa, probably following the expansion of savanna habitat during the past 2.5 million years. While the related Alcelaphus and Damaliscus species continued their pan-African distribution, western and northern Connochaetes populations disappeared and the species now only survives in the southern refugium, from mid-Kenya southwards. The adaptations of the two wildebeest species to their respective savanna ecosystems made them the keystone species among guilds of grazing ungulates. Although their size and conformation are very different (the mass of an adult CommonWildebeest is 250 kg compared with a 157 kg Black Wildebeest), they share traits that help explain their success. The distributions of the two species overlapped periodically after the speciation of the Black Wildebeest ca. 1 mya (Corbet & Robinson 1991), but habitat preferences and behavioural differences have contributed to reproductive isolation (Brink 2005). However, the two species are also genetically close enough to produce viable hybrids (Fabricius et al. 1988), and indeed hybridization and introgression now pose significant risks to the Black Wildebeest due to injudicious translocations that have brought the species into contact with each other at numerous localities in South Africa (Grobler et al. 2011). Wildebeest are adapted to exploit extensive grasslands that can support great concentrations of grazing antelopes, producing highly nutritious green pastures during growing seasons (Estes 1991a). Productivity is maintained by recycling of the manure distributed by the massed animals. Both species are equipped with broad incisor

arcades to take large bites of short grasses (in both species, the dental formula is I 0/3, C 0/1, P 3/2, M 3/3 = 30, with the second lower premolar absent, as in Beatragus).They are bulk rather than selective feeders. The Common Wildebeest is virtually a pure grazer, whereas the Black Wildebeest also browses to some extent on karroid shrubs. Both species are territorial, which represents the original sedentarydispersed social organization shared by all but a few antelopes in the subfamily Antilopinae. In the resident phase, the Black Wildebeest defends a far larger territory than its congener. In the migratory phase, the Common Wildebeest defends temporary, very small territories. How close migrating Black Wildebeest territorial !! would tolerate one another is unknown: there has been no migratory population since the species was nearly exterminated in the late 1800s. Though efforts to restore the species have been outstandingly successful, the thousands of Black Wildebeest now live in isolated herds on farms and ranches in the Highveld and also outside their natural range as far afield as Namibia. What sets wildebeest apart from all other antelopes – and must contribute to their dominance of competing migrants – is their reproductive system.They have abandoned the ancestral system wherein newborn calves go through a hiding stage (for example, Hartebeest Alcelaphus buselaphus) that lasts days and even weeks, during which mothers must remain in attendance. In highly mobile aggregations, selection would favour reducing or even eliminating the hiding stage. Wildebeest abandoned it altogether and evolved an entirely new follower-calf system. The main features are extreme precocity of the newborn, which gain mobility within minutes of birth, aggregations of pregnant "" on calving grounds, a short peak during which some 80% of the calf crop is born, and associations of mothers and calves in maternal herds (Estes 1976, Estes & Estes 1979). In this way, not only are predation and predators limited, but herds with older calves provide essential cover for neonates (substituting for high grass) during the first day or two when they are most vulnerable. Richard D. Estes 527

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Connochaetes gnou BLACK WILDEBEEST (WHITE-TAILED GNU) Fr. Gnou à queue blanche; Ger. Weissschwanzgnu Connochaetes gnou (Zimmermann, 1780). Geogr. Gesch. Mensch. Vierf. Thiere 2: 102. South Africa, ‘Die lander der Caffern, ziemlich tief ins land vom Cap gerechnet in grossen Waldern ohnmeit der Uchtermanns Brenjes hogde und Camdebo’; since selected as Eastern Cape Prov., Somerset East Dist., Agterbruintjieshoogte (Grubb 1999).

ABOVE: Lateral and palatal views of skull of Black Wildebeest Connochaetes gnou. LEFT: Black Wildebeest Connochaetes gnou.

The moniker Gnu derives from the Khoikhoi (aka Hottentot) descriptive term for the bellowing snort the animal gives when alarmed (Skinner & Chimimba 2005). Taxonomy Monotypic, with no subspecies recognized. Synonyms: capensis, connochaetes, gnou, gnu, operculatus. Chromosome number: 2n = 58 (Wurster & Benirschke 1968, Corbet & Robinson 1991). Hybridization with Common Wildebeest has been recorded, and hybrids are fertile (Fabricius et al. 1988). Description Similar in appearance to its close relative the Common Wildebeest Connochaetes taurinus, but smaller in size, with strongly built forequarters sloping to more slender hindquarters. Rich, dark-brown colour; mature !! have a black face and a darker, almost black appearance; newborn calves are fawn-coloured. Head and face is less elongated than in Common Wildebeest, with a broad muzzle, erect facial tuft, preorbital glands and distinct tuft of hair under the chin. Further tuft of hair found on the chest, between the forelegs. Stiff, upright, trim mane, creamy-white with dark tips. Horse-like tail is creamy white but dark at base, almost reaching the ground. Pedal glands are present on front hooves and excrete a sticky substance; inguinal glands are absent. Males larger than "". Horns, present in both sexes, are smooth with expanded bases, directed forward and downwards before curving up sharply. Adult !! have heavy horns with prominent bosses, while horns of ""

are slighter. Calves have long spike-like horns that start to curve at approximately nine months; !! attain full horn shape at 4–5 years (von Richter 1971b). In addition to the differences in horn shape and horn base architecture, the skull of the Black Wildebeest can be distinguished from that of the Common Wildebeest in being generally smaller, shorter and dorsoventrally flattened. The angle of the braincase to the face is greater in the Black Wildebeest than the Common Wildebeest, the orbits are absolutely and proportionally enlarged, while the frontal suture is fused (Brink 2005). These features appeared progressively over the last million years, and can be observed in rudimentary form in the earliest fossil populations of ancestral Black Wildebeest (Brink 1993, 2005). All molar teeth are erupted by the third year of age, with the upper third and fourth premolars erupting at 28–30 months; patterns of tooth eruption are discussed in detail by von Richter (1971b). Geographic Variation

None recorded.

Similar Species Connochaetes taurinus. Larger, greyish in colour with dark brindle stripes on the neck and shoulders; head and face elongated; chin with long beard and limp hair, and shaggy mane of long black hair under the throat; black tail almost reaching the ground; horns smooth, arising from swollen bosses and directed outwards and slightly downwards before curving up; horn tips pointed

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Habitat A species characteristic of the open plains Highveld grasslands (the Highveld BZ) and Karoo shrublands (the Karoo subzone) of southern Africa. The high central plateau grasslands are characterized by flat to rolling hills, and mountainous areas with altitudes ranging from 1350 to 2150 m. These areas are dominated by a single layer of grasses, with cover dependent on rainfall and the degree of grazing. Abundance Several thousand are all that remain of the millions that once roamed South Africa’s Highveld and Karoo regions. The total population is estimated at more than 18,000 animals (including the introduced population in Namibia), 80% of which occur on private land and 20% in protected areas. However, population size is increasing, especially on private land, with a large extralimital population now established in Namibia, where importations from South Africa led to a dramatic rise in the estimated total numbers, from 150 in 1982 to more than 7000 in 1992 (East 1999).

Connochaetes gnou

inwards and often slightly backwards. Although the ranges of the two species formerly barely overlapped (with Black Wildebeest confined to the temperate, treeless Highveld and Karoo regions of South Africa, Swaziland and Lesotho), stocking of both species outside their natural range has led to fertile hybrids where the two species are kept on the same property (Fabricius et al. 1988). Distribution Black Wildebeest are endemic to southern Africa. According to von Richter (1974) they formerly migrated in large numbers in both east–west and north–south directions, probably following the onset of rains and changing vegetation cycles. Historical Distribution Formerly confined to the central inland plateau of South Africa, specifically the Highveld regions of Free State, Gauteng and North West Provinces, parts of the Eastern Cape and Northern Cape provinces, and marginally in the grassveld regions of western KwaZulu–Natal, in the foothills of the Drakensberg Range (von Richter 1974). It has been suggested that their occurrence in the western parts of KwaZulu–Natal may have been due to a local movement from the Free State during the winter months in search of better feeding grounds. Black Wildebeest were also recorded seasonally in grassland areas of W Swaziland and the western parts of Lesotho (where they were exterminated before 1900). A detailed discussion of the historical distribution of the species is given by von Richter (1971a). Current Distribution Black Wildebeest have been widely reintroduced to areas within their former distribution range, as well as to farmland and reserves well outside their natural range. For example, although they do not occur naturally in Namibia, they have been introduced and now occur widely throughout Namibia’s farming districts (East 1999). They have also been reintroduced in Lesotho (in Sehlabathebhe N. P.) and in Swaziland in Malolotja N. R. (East 1999).

Adaptations Black Wildebeest actively condition their own preferred habitat by persistent grazing and trampling (von Richter 1974, Kok & Vrahimis 1995), especially applicable to sub-optimal areas, where territorial !! open up fairly dense vegetation. These !! are closely attached to their territories throughout the year; here, vegetation horning during demonstrative-threat displays plus sustained grazing in the immediate vicinity of the stamping ground keeps the grass in the favoured short state, thereby enhancing visibility (Kok & Vrahimis 1995). Although scattered trees may be found in concentration areas, animals rarely use trees for shade and shelter. Generally these darkly coloured ungulates are exposed to direct solar radiation and other, often extreme, climatic conditions. During early morning and late afternoon, when temperatures are lower, animals present the long axis of their bodies to the sun to facilitate heat uptake (Vrahimis & Kok 1992). Conversely, during the heat of day, animals orientate themselves to expose the smallest surface to direct sunlight, thereby compensating for excessive heat load. In summer, when the hottest time of the day usually coincides with the highest windspeed, it is more likely that animals orientate with respect to wind direction, thus maximizing amount of airflow over the entire length of their bodies (Vrahimis & Kok 1992). Additionally, animals prefer lying down to standing during the heat of the day, thereby reducing the intake of reflected radiation from the ground. Although they are subjected to high ambient temperatures during the day, the brain temperature in Black Wildebeest is usually within 0.2 °C of arterial blood temperature. Selective brain cooling is absent, even though brain temperature may rise to 42.0 °C (Jessen et al. 1994). Jessen et al. (1994) established that heat storage as a thermoregulatory tactic is relatively unimportant for Black Wildebeest. The Black Wildebeest may be considered to have extraordinary homeothermic capacity, the key being the insulation provided by its fur. Furthermore, due to enlarged evaporative surfaces in the long snout of the genus Connochaetes, heat is also dissipated by panting. Even in a thermal environment characterized by large contrasts between day and night, the circadian variation of body temperature in the Black Wildebeest appears to reflect an endogenous rhythm rather than a reaction to cyclic thermal loads. In addition, it was recorded that increased metabolic rate, as during 529

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episodes of chasing, increases blood temperature (1.3 °C within 4 min). Without free access to water, heterothermy may be obligatory (Jessen et al. 1994). However, regarding selective brain cooling, it has been established that the mammals with the greatest capacity for such brain cooling possess carotid retes, bilateral networks of arterioles in, or just outside the cranial cavity, in the main arterial blood supply to the brain (Gillilan 1974 and Simoens et al. 1987, in Mitchell et al. 2002). In the artiodactyls, the retes have a thermoregulatory function and are heat exchangers in which the arterial blood destined for the brain is cooled by venous blood returning from the evaporating surfaces of the nasal cavity (Baker 1982 and Mitchell et al. 1987, in Mitchell et al. 2002). Studies conducted in a free-living Black Wildebeest revealed that, as one would normally expect for artiodactyls, selective brain cooling could develop, but, unexpectedly, seldom actually did in their natural habitats (Jessen et al. 1994). Thus, Jessen et al. (1994) found that at a brain temperature that exceeded 42 °C, established under induced intense exertional hypothermia, selective brain cooling was abandoned and brain and blood temperatures were similar. Foraging and Food The Black Wildebeest is predominantly a grazer and prefers short grassveld (von Richter 1974). A feeding study listed 41 plant species utilized, and showed that 63% of the Black Wildebeest’s main diet consisted of grass and 37% of karroid shrubs (Van Zyl 1965). Grasses such as Sporobolus spp., Themeda triandra and Cynodon dactylon formed the bulk of their diet. Karroid shrubs were browsed during the colder months of the year, probably correlating with the decline in the nutritional value of the grass. Roberts (1963) found that, in the central Free State, 86.8% of the diet consisted of grasses, especially Eragrostis lehmanniana (31.3%), Themeda triandra (25.3%), Panicum coloratum (20.3%) and Cynodon dactylon (9.9%). Research conducted on feed utilization and digestion in Golden Gate Highlands N. P. in the Free State indicates that the diet of Black Wildebeest consisted of 80% grass and karroid shrubs and that the relatively high fermentation rate is due to suitable substrate for microbial activity found in that area (Van Hoven & Boomker 1981). These observations of grass predominating in the diet are borne out also by studies involving stable carbon isotopes (Sponheimer et al. 2003b). The feeding preferences of a tame Black Wildebeest cow were investigated over a two-year period (S. Vrahimis pers. obs.). This animal was hand-reared, but subsequently integrated into a freeranging herd. Results obtained during the free-ranging stage showed that this animal utilized grass (93.7%), karroid shrubs (3%) and herbs (3.3%). While mainly a non-selective grazer there was evidence of some preference for certain grass species. This was evident due to the fact that the animal would feed on a specific grass species for quite some time (approx. 20 min.), before moving on and feeding on another grass species, once again for a period of time. All social groups exhibit a typical bimodal feeding pattern with high-intensity grazing peaks during the early morning and late afternoon (Vrahimis & Kok 1993). Generally, !! spend more time grazing than "". For territorial !! this is perhaps explicable on the basis of their intensive involvement in territorial behaviour and, therefore, greater energy costs, while in the case of bachelor herds, which usually occupy marginal areas with poor grazing (often to avoid harassment by territorial !!), the animals have to feed longer. In addition, the

two categories of grass plants found in grasslands described by Low & Rebelo (1996), namely, sweet grasses having a lower fibre content which maintain nutrients in winter (palatable), and sour grasses having a higher fibre content which tend to extract nutrients during winter (unpalatable), will also determine the time spent feeding during winter in different grassland areas. Black Wildebeest are dependent on water and drink regularly, mainly in the late afternoon. Social and Reproductive Behaviour Black Wildebeest are gregarious, their social groups comprising female herds, bachelor herds and territorial !!. Average size of a female herd is 28 animals (range 14–49; n = 144) and comprises adult and subadult "" and calves, and in most cases an attending territorial bull. A number of yearling !! are usually also found in these herds. Bachelor herds contain !! of various ages, including yearlings, with an average of 21 individuals (range 11–32; n = 52). Tolerance of the !! towards one another is notable in these groups (von Richter 1971b). Two types of territorial !! are recognized: isolated, solitary bulls, usually older animals, and those found in a territorial network. Definite dominance hierarchies are evident in such a territorial system where most matings take place. Home-range and size of territories is largely dependent on the density of animals and the size of the available area and available food source. Von Richter (1971c) described territorial !! displaying vigorously, defending their territories and challenging neighbouring bulls or intruders throughout the year in the Free State. Territoriality is a prerequisite for reproduction, as non-territorial !! are excluded from the rut. Encounters between territorial !! are highly ritualized and resemble those of Common Wildebeest (von Richter 1974). Estes (1991a) records standing in erect posture, a rocking canter, calling, defecation preceded by pawing, kneeling, ground horning, rolling on stamping ground, herding and chasing as the means of advertising territories. The territories of neighbouring !! are separated by distances of 180–450 m (von Richter 1972). Agonistic behaviour between territorial !! is comprised of dominance-threat displays, defensive-submissive displays and fighting (ramming horns). In several instances, carcasses of !! with interlocked horns have been found. Black Wildebeest are among the most vociferous of all antelope species. Territorial !! have a very specific call, best described as a loud ‘woink’, which carries over a long distance and is heard day and night. Calling is more intense at night, and the fact that this is the main signal available after dark is well demonstrated during full moon (when covering of the moon by a cloud leads to more intensive calling). When this call is made, the head jerks back and the mouth opens wide. Males respond to each other’s calls. Observations conducted during the mating season include the recording of interactions between territorial !! and "". Interactions consist of herding, chasing, inspecting for receptiveness and mating. On a daily basis, a build-up of interactions was evident from 13:00 to 18:00h; 39–66% of all interactions consisted of inspections of the receptiveness of "", while chasing comprised a third of all activities. Most matings observed were between 14:00 and 18:00h, but it is suspected that mating often occurs at night. Female herds move from one territory holder to another and the time spent with each can vary from a few hours to several days. Herd movements occur mainly at night. When approaching a territorial

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!, the herd is usually met by the ! at the periphery of the territory, performing a typical, stiff-legged gait, which Estes (1991a) referred to as a ‘rocking canter’. Once the herd has entered his territory, the ! moves about in the herd inspecting "" at random. When approached within a herd, "" lift and swish their tail. Heads are often held at an angle-horn position (agonistic behaviour – readiness for combat) and circling of the ! and " then follows. Occasionally "" drop their heads and interlock horns with the !. Urination on demand and urine testing (flehmen) continuously take place (Estes 1991a). When about to mate, the ! approaches in the typical lowstretch posture (Estes 1991a), resting his chin on the female’s rump, stands bipedally, and mates. Most matings take place on the periphery of the herd where the ! normally stands separately; a receptive " usually approaches the ! here with her tail lifted and swishing. Parturition takes place within the herd and all births witnessed were between 08:00h and 12:00h (S. Vrahimis pers. obs.). Females about to calve become very restless, lying down and standing up continuously. During the final stage the " lies flat on the ground. Whether the " eats the afterbirth, as sometimes occurs in Common Wildebeest, is unknown. Von Richter (1974) recorded an average of 9 min elapsing before a calf can stand on its feet. Throughout the day, lying down dominates (85%) all categories of activities for calves 1–3 months of age. During subsequent months this activity stabilizes to a much lower level (55–56%) of the diurnal time budget. The most noteworthy development in diurnal activities involves a progressive increase in the time spent feeding (4–33%). From the second month, the formation of nursery or crèche groups is evident, showing synchronized activity distinct from the main body of the herd, such as playing and exploring. At approximately three months, aggregations of calves are more pronounced, with the young spending the largest part of the day together, indicating the start of the weakening of the mother–calf bond (Vrahimis & Kok 1994). Regarding associations with other species, the distribution range of Black and Common Wildebeest (Aylward 1881, Sidney 1965, Skead 1987) was known to overlap slightly, especially in the Free State and regions of the Highveld. Associations of Black Wildebeest with the now extinct Quagga Equus quagga quagga were recorded (Bryden 1889, Sclater 1900/1901, Lydekker 1926). Ostriches Struthio camelus also co-existed with both wildebeest species. Early explorers and hunters passing through the central open plains of South Africa described huge aggregations of wildebeest, Quaggas, Plains Zebra Equus quagga, Blesbok Damaliscus pygargus phillipsi, Hartebeest Alcelaphus buselaphus and Springbok Antidorcas marsupialis (Cumming 1980 [1850], Bryden 1893). Reproduction and Population Structure Black Wildebeest are seasonal breeders, and breeding appears to be triggered by shortening daylength (Skinner et al. 1973). The majority of calves are born within a three-week period (von Richter 1971c), from midNov to end Dec. Generally, the peak mating season is from midMar to end Apr. Most "" conceive at the age of 16 months and calve when they are two years old (S. Vrahimis pers. obs.), after a gestation period of approximately 8.5 months (Skinner et al. 1973). Males can mate successfully at the age of 16 months, but first have to secure a territory before being allowed to mate. In one area all adult !! were removed, leaving only young !! (16–18 months)

present during the mating season. These !! mated successfully, but the calving percentage was not as high as expected (68%). Average age of territorial !! is four years, but in areas where excessive hunting occurs, two-year-old !! were territorial. A single calf is born (there being no records of twins; von Richter 1974, S. Vrahimis pers. obs.) and the average birth-weight recorded is 14 kg (range 12.5–15.5 kg; n = 4). Calves are weaned at 6–8 months; the composition of the mother’s milk is discussed by Van Zyl & Wehmeyer (1970). Birth rates are generally high, ranging from 72 to 97% (S. Vrahimis pers. obs.). Von Richter (1971c) recorded lower reproductive rates (47–68%) in some areas of marginal habitat suitability, and stated that the reproductive performance, in a specific area, can be variable and is linked to seasonal climatic conditions, as 86% and 100% of cows were recorded birthing in the same area on different occasions. A lack of predators in the enclosed areas where Black Wildebeest are held today lowers mortality. The current primary cause of mortality is separation of calves from their mothers during game capture operations. Von Richter (1971c) recorded 5–11% mortality in two different areas, also ascribed to separation of cows and calves. Wildebeest have a unique reproductive system, which includes a short, sharply defined calving season that generates a superabundance of newborn calves (Estes 1991a). Connected to the species’ former migratory habits, there is a tendency to concentrate in large numbers, a preference for short grass and calves follow their mothers shortly after birth, all of which are incompatible with a concealment strategy. Although Black Wildebeest are no longer able to migrate and no longer permitted to occur in large concentrations, this could be a vestige of behaviour important for the survival of wild animals in the past. Today, Black Wildebeest are extensively managed in protected areas that do not allow the formation of natural population structures. In most areas animals are captured and removed annually, depending on the animal numbers, climatic conditions and grazing. Attempts are made to maintain a balance, with sex ratios kept close to parity and equal representation of all age classes. Pre-natal sex ratio of 91 foetuses examined showed 1.17 !! : 1 ". Von Richter

Black Wildebeest Connochaetes gnou.

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Conservation IUCN Category: Least Concern. CITES: Not listed. By 1900 the Black Wildebeest was nearly exterminated by hunting and the reduction of available habitat due to human settlement (von Richter 1971a, 1974), as well as the periodic outbreak of diseases. Fisher et al. (1969) stated that Black Wildebeest were subjected to constant unregulated persecution for more than a century, culminating in slaughtering of immense numbers for their skins in Predators, Parasites and Diseases Presently, in South Africa, the 1870s. Millais (1895) expressed concern, fearing the extinction these animals are confined to nature reserves and private farmland, of the species, as formerly, ‘tens of thousands of these wildebeest where major predators no longer occur. In historical times, explorers had been scattered in troops of from twenty to fifty over the face and hunters described instances where large numbers of animals of the Southern Transvaal and Free State Highveld, and after careful died as the result of the outbreak of diseases, apparently scabies inquiries, there were hardly more than 550 in existence’. These or mange (Harris 1840, Cumming 1980 [1850], Bryden 1889). animals were exterminated throughout the greater part of their Von Richter (1974) suspected that rinderpest and foot and mouth former distribution range and recorded as being extinct in the wild disease, which seriously affected Common Wildebeest, could have (Stevenson-Hamilton 1917, Fitzsimons 1920, Shortridge 1934). had an influence on Black Wildebeest. Fatal, sporadic outbreaks of Fortunately, the species was saved from extinction by conservationanthrax in Black Wildebeest were recorded up to 1943 (Neitz 1965). minded farmers, especially in the Free State, by protecting herds on Black Wildebeest are also asymptomatic carriers of the virus that their farms. causes malignant catarrhal fever, a disease deadly to cattle. A survey Following the turn of the nineteenth century, Black Wildebeest conducted in 1988 to establish the extent of malignant catarrhal numbers started to increase rapidly (Fitzsimons 1920). However, fever in the Free State showed that there were 13 confirmed cases the drought in 1933 had a severe impact on this species, and von recorded from 1977 to 1987. The survey involved 205 landowners Richter (1971a) provides information from one population where that had either Black or Common Wildebeest, or both species on only 15–20 animals, mainly !!, survived out of a population of their properties (Vrahimis & Prinsloo 1988). As a result of malignant approximately 400 animals. According to Fisher et al. (1969), until catarrhal fever outbreaks in South Africa restrictions were placed on 1936 the only Black Wildebeest surviving were all on private land, the relocation of both Black and Common Wildebeest in 1982 and and later that same year animals were released into the then Free no new introductions were allowed. Malignant catarrhal fever was State G. R. (Somerville). When the Free State G. R. was abolished, then also listed as a notifiable disease and all cases had to be reported. the herd was transferred to Willem Pretorius G. R. in 1956, where Due to the fact that it was later found that malignant catarrhal fever numbers had increased to 370 animals by 1966. Gradually, over the was not a significant problem, these restrictions were lifted in April years, reintroductions took place on a large scale. Translocations 1993, once again allowing for free movement of both wildebeest were mainly carried out from the small number of original herds in species in South Africa. the Free State and to a lesser extent from a herd in the North West The endo- and ectoparasite burdens of Black Wildebeest have been Province, as well as from the De Beers estate in the Northern Cape investigated at two localities in South Africa, which revealed a small (von Richter 1971a). number of species and relatively small number of helminths and ticks Although it is now supposedly safe from extinction, Black at both localities (Horak et al. 1983b). This was ascribed to fairly Wildebeest numbers are still relatively low. Presently, the largest cold climates prevailing in the survey regions and the fact that Black threat to the species is hybridization with the Common Wildebeest. Wildebeest appear to be fairly resistant to parasitic infestation. The Sidney (1965) mentions the occurrence of hybrids between Black nematode burden of Black Wildebeest examined in another survey and Common Wildebeest in KwaZulu–Natal, which is probably was found to be extremely low when compared with that recorded one of the first recorded instances. Fabricius et al. (1988) provide by Horak et al. (1983b) and was also perceived to be a reflection evidence that hybrids are fertile. As most recorded instances of of the adverse climatic conditions, namely hot summers and cold hybridization have been Common Wildebeest !! cross-breeding winters with low rainfall during either season (Boomker et al. 2000). with Black Wildebeest "", Corbet (1991) speculates that this In Black Wildebeest, as in the Common Wildebeest, larvae of could be due to the larger size of the former species being able to oestrid botflies inhabit the sinuses and nasal septa, apparently without dominate the latter. The external appearance of hybrids can vary, causing obvious harm to the host. Horak et al. (1983b) observed with the most prominent and reliable feature being the shape of an erratic, but progressive, increase in the numbers of first stage the horns. Although first generation hybrids are easily identified, larvae of Gedoelstia spp. on the dura mater of Common Wildebeest hybrids interbred with pure stock Black Wildebeest are difficult to calves until they reach the age of approximately 13 months; likewise, recognize on appearance alone and advanced backcrosses are hard Horak (2005) noted that large numbers of first instar larvae of G. to detect (Grobler et al. 2011); Ackermann et al. (2010) discuss hassleri appeared to accumulate on the dura of Black Wildebeest in several anomalous cranial morphological characteristics of hybrids. the Eastern Cape from June to August. Flies of the genus deposit first It has always been believed that hybridization only occurs under instar larvae on the cornea or conjunctiva of the eyes of their hosts artificial conditions, but, recently, in a protected area (approx. from where they migrate either via the optic nerve tract or artery 6000 ha) in KwaZulu–Natal where large herds of both Black and to the subdural cavity and dura mater, and then via alterior routes to Common Wildebeest were housed together, all the wildebeest had the nasal passages where they moult to the second instar (see Horak to be destroyed because of hybridization. & Butt 1977). (1972) reported that game-capture operations changed sex ratios, whereas in one national park, from which no animals were removed, the sex ratio was near parity.Weigl (2005) gives a longevity record in captivity of 21.8 years, although in the wild longevity is estimated at approximately 14 years (based on the known age of a single animal; S. Vrahimis pers. obs.).

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For several years Tussen-die-Riviere N. R. in the Free State had large populations of both Black and Common Wildebeest together, with no apparent problem of hybridization. It was decided in the early 1990s that these animals should be separated and that preference should be given to Black Wildebeest (being endemic to central South Africa) and all Common Wildebeest were removed from the reserve. At one stage, this was one of the largest populations of Black Wildebeest in South Africa, but annual hunting on the reserve resulted in almost the entire Black Wildebeest population being concentrated in a narrow area south of the Orange R., causing serious habitat degradation. After all the Common Wildebeest on the reserve were culled, Black Wildebeest from other protected areas in the Free State were subsequently translocated back to Tussen-dieRiviere N. R.

Measurements Connochaetes gnou HB (!!): 1800 (1700–1880) mm, n = 92 HB (""): 1660 (1530–1750) mm, n = 53 T (!!): 540 (460–610) mm, n = 92 T (""): 500 (420–560) mm, n = 53 HF c.u. (!!): 470 (440–500) mm, n = 92 HF c.u. (""): 450 (430–480) mm, n = 53 E (!!): 170 (150–190) mm, n = 92 E (""): 160 (140–180) mm, n = 53 WT (!!): 160.0 (134.0–200.0) kg, n = 253 WT (""): 131.0 (117.0–149.0) kg, n = 64 Free State, South Africa (S. Vrahimis pers. obs.) Maximum recorded horn length is 74.6 cm for a pair of horns from the Free State, South Africa (Rowland Ward) Key References

Estes 1991a; von Richter 1971b, c, 1974. Savvas Vrahimis

Connochaetes taurinus COMMON WILDEBEEST Fr. Gnou bleu; Ger. Streifengnu Connochaetes taurinus (Burchell, 1824). Travels in Interior of Southern Africa 2: 278 (footnote) [1824]. Apparently ‘Kosi Fountain’, but lectotype came from South Africa, North West Prov., Vryburg Dist., ‘Chue Spring, Maadji Mtn [Klein Heuningvlei]’; see Grubb (1999).

corniculatus, fasciatus, gorgon, hecki, henrici, johnstoni, lorenzi, mattosi, mearnsi, reichei, rufijianus, schulzi. Chromosome number: 2n = 58 (Wallace 1978b, Corbet & Robinson 1991).

Common Wildebeest Connochaetes taurinus albojubatus.

According to Skinner & Chimimba (2005), the now uncommon name Gnu applies to the Black Wildebeest C. gnou and derives from the Khoikhoi (aka Hottentot) descriptive term for ‘the bellowing snort they give when alarmed’. But as an onomatopoeic term, it sounds more like the territorial advertising call ‘GA-nou!’. ‘Wildebeest’ is the misnomer bestowed on both species – first the Black – by the Dutch colonists because of a fancied resemblance to the ‘wild ox’ of Eurasia. Taxonomy Five subspecies are usually recognized (Ansell 1972, Grubb 2005). Synonyms: albojubatus, babaulti, borlei, cooksoni,

Description A large, long-faced antelope with cow-like horns, heavy-set body but thin legs, and forequarters higher than hindquarters. Head large, muzzle broad, with wide incisor row and flexible lips; nostrils covered with skin flap. Eyes obscured by long lashes, iris yellow, tapetum reflection greenish. Ears narrow, ca. 200 mm; frontal mask black (but subject to individual and subspecific variation). Neck short and thick, with mane upstanding or lax, and beard from chin to forelegs. Coat short, glossy with vertical ‘stripes’ of longer, dark hair on neck, shoulders and chest. Colouration of torso blue-grey, tan, or brown with vertical stripes of longer, dark hair on neck and thorax (subject to partial moult, most developed in breeding season, making individuals look darker). Tail, mane, backs of ears and facial blaze black; beard black, tan, or off-white. Lower legs lighter coloured. Calves fawn to tan, mask, beard, dorsum and tail black (but up to one in ten has blond crown). Change to adult colouration complete in third month. Hooves with little taper; false hooves well-developed. Tail hock-length with long hair reaching to heels (widely used as a fly whisk). Preorbital glands are developed in both sexes, but larger in !, and covered with hair, the central duct absent or vestigial, and the secretion appears as a clear oil. Pedal glands are present on the forefeet, but rudimentary on hindfeet (Pocock 1910, Ansell 1969). The secretion of the pedal glands is black, pungent and detectable to the human nose (resembling fresh tar); the constituents of the secretion have been examined by Wood (1998). Inguinal glands are absent. Horns cowlike, present in both sexes, unridged and extend sideways with tips pointing inward; horns of ! are wider and thicker, 533

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ABOVE: Lateral view of skull of Common Wildebeest Connochaetes taurinus. LEFT: Common Wildebeest Connochaetes taurinus taurinus.

with well-developed bosses beginning in third year. Attwell (1980) considered the loss of the second lower premolar in adults (which retain only the third and fourth premolars) as representative of an advanced stage of evolution in ungulate dentition. Adult dentition is complete by just over three years; Attwell (1980) discussed age determination based on tooth eruption, attrition and cementum layers. Horn development to 24 months is illustrated in Kingdon (1982: 530) and herein. Geographic Variation C. t. taurinus (Blue Wildebeest or Brindled Gnu): Namibia and South Africa, north of the Orange R., to Mozambique and from Mozambique to Zambia south of the Zambezi R., and from SW Zambia, west of the Kafue R., to SE and S Angola. Bluish-grey coat, black beard and upstanding black mane. Also commonly called the Brindled Gnu because of the stripes of darker hair on the neck and shoulders extending back to about the middle of the body. C. t. cooksoni (Cookson’s Wildebeest): restricted to the Luangwa Valley, Zambia; believed to have ranged, but only as vagrants, onto the adjacent plateau into C Malawi (Ansell & Dowsett 1988). Browner than other races. C. t. johnstoni (Nyassa, Johnston’s or White-banded Wildebeest): north of Zambezi R. in Mozambique to east-central Tanzania, and formerly in S Malawi, but now extinct there (Ansell & Dowsett 1988). Sometimes referred to as the ‘White-banded Wildebeest’ for the pale chevron between its eyes (often absent in the Tanzania population). Distribution largely confined to Acacia savanna of

major river valleys within the miombo (Brachystegia) woodland zone. Former northern distribution limit Wami R., Tanzania. C. t. albojubatus (Eastern White-bearded Wildebeest): N Tanzania to C Kenya just south of the Equator, west to the Gregorian Rift Valley; southernmost point in recent past (1950s) at least to the Handeni–Kondoa road (5° 30' S) (J. Kingdon pers. obs.). Lighter grey than C. t. mearnsi, larger and about 50 kg heavier, with wider horns but less-developed boss; beard white to tan. C. t. mearnsi (Western White-bearded Wildebeest): N Tanzania and S Kenya west of the Gregorian Rift Valley, reaching L. Victoria at Speke Bay.This is the smallest, darkest and most numerous race – the CommonWildebeest of the Serengeti Plains. Horns shorter but boss most developed; beard white to tan. Male territorial advertising call distinctively different from other races: resonant croak with mouth closed repeated many times, compared with more metallic series of 6–12 calls with mouth open of other races. Female mimicry of male secondary characters includes a penile tuft of adipose tissue, rarely if ever present in other subspecies (Estes 1991b). Often unrecognized as a subspecies distinct from C. t. albojubatus (Heller 1913b, Allen 1939, Kingdon 1982), yet DNA studies support the phenotypical differences and indicate that mearnsi is the longest isolated, and genetically most unlike the other races (Arctander et al. 1999). A small resident population of C. t. mearnsi existed in the Gregorian Rift Valley near L. Naivasha, but was eliminated by fencing and shooting by the end of World War I (Roosevelt & Heller 1915). The only known place of potential contact now is via the Rift Wall opposite L. Natron, which is sometimes traversed by albojubatus (D. Peterson pers. comm.).

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Similar Species Connochaetes gnou. Fossil evidence indicates the two species separated ca. 3 mya (Gentry 1978, Vrba 1979, Flagstad et al. 2001); although the ranges of the two species formerly barely overlapped (with Black Wildebeest adapted to the temperate, treeless Highveld and Karoo), stocking far outside its natural range (Botswana, Namibia, Zimbabwe) has led to fertile hybrids where the two species are kept on the same property (Fabricius et al. 1988). These are much smaller, darker antelopes with more evenly developed limbs, dangerous horns that project forward like meathooks, and a white tail. Distribution Endemic to Africa, historically ranging in short grasslands and open bushland and woodland from N and E Namibia, Botswana, N South Africa (generally, north of the Orange R., in the Northern Cape, North West, Limpopo, Mpumalanga, Gauteng, Free State and NE KwaZulu–Natal Provinces) and Mozambique to ca. 38° E (Chuyulu Hills) in S Kenya, west to L. Victoria at ca. 01° S. Palaeontological evidence indicates that C. taurinus lived in the Saharan region in the late Pleistocene (Gentry 1978), and more recently in West Africa, possibly in a ‘bovid’ western refugium (Arctander et al. 1999). In pre-colonial times, Common Wildebeest dominated the teeming assemblages of plains game in most of the Acacia savanna ecosystems of eastern Africa. Migratory populations rivalling those of the Serengeti Plains existed in Kenya’s Athi-Kapiti Plains; in Tanzania, stretching from the Gregorian Rift Valley to Mt Meru, mainly comprising S Masailand; and in much of Botswana. Populations in the order of 25,000–50,000 inhabited the broad river valleys incised in the miombo (Brachystegia) woodlands of SE Tanzania and N Mozambique (C. t. johnstoni), the Liuwa Plains of Barotseland, SW Zambia, extending into SE Angola; the Limpopo Valley between Zimbabwe and Mozambique; South Africa from the lowveld of the Limpopo and Mpumalanga provinces west into the foothills of the Drakensberg; and in NE Namibia, Common Wildebeest migrated between Etosha G. R. into

Connochaetes taurinus

adjacent Botswana and Angola, before a game-proof fence was erected along the Namibia–Botswana border, followed by fencing of the eastern border of Etosha N. P. Loss of range outside of protected areas has resulted in the replacement of migratory populations with smaller more sedentary populations within protected areas, as in the case of Etosha N. P. (Berry 1997, East 1999). The only range state in which the Common Wildebeest has been exterminated is Malawi. According to Sidney (1965), the Nyassa Wildebeest’s range in Malawi used to be bounded by L. Malawi on the north, the Shire R. on the west and the Zambesi on the south. The alluvial plains of the river and Lakes Chilwa and Chuita must once have supported a sizeable population. But as one of the best agricultural regions of the country, it was capable of supporting a dense human population; the few animals that persisted at the south end of L. Chilwa were finally shot out in 1925 (Sweeney 1959; see also Ansell 1982, Ansell & Dowsett 1988). Common Wildebeest have also been introduced to regions outside their former distribution range, including the Eastern Highlands of Zimbabwe, parts of KwaZulu–Natal and private farmland in Namibia. Habitat A quintessential plains antelope preferring open short grassland, and closely associated with Acacia savanna. Rainfall averages between 400 and 800 mm across its range. Migratory populations disperse over the arid part of the range during the wet season, and concentrate in higher-rainfall areas with permanent water during the dry season. Prefers well-drained soils with firm footing, avoiding wetlands and waterlogged soil, although their dry season range typically includes extensive areas of black-cotton soil associated with Acacia drepanolobium, A. seyal and Balanites aegyptica. They are rarely found above 1800–2100 m (e.g. floor of Ngorongoro Crater in Tanzania), but may be transient in montane grassland and hilly terrain (e.g. movement between seasonal pastures; Estes & Small 1981, Estes 2002a, Musiega & Kazadi 2004). Abundance During the late 1990s, the global population was estimated at around 1,298,000 (correcting for undercounting biases in aerial surveys; East 1999), with the migratory Serengeti–Mara population representing about 70% of global species numbers (942,000, having dropped below 1 million following the severe 1993 drought). Other population estimates were: Blue Wildebeest, 150,000 (with about half in protected areas, and one-quarter on private land and conservancies); Cookson’sWildebeest, 16,000 (about 60% in protected areas); Nyassa Wildebeest, 96,000 (about two-thirds in protected areas, particularly Selous G. R.); and Eastern White-bearded Wildebeest, 94,000 (with about two-thirds in and around protected areas). The most recent estimate of the total population size of Common Wildebeest is around 1,550,000 (Estes & East 2009), largely due to the rebounding of the Serengeti population of Western White-bearded Wildebeest to about 1,300,000 (Thirgood et al. 2004); other subspecies populations are estimated at 130,000 Blue Wildebeest, 5000–10,000 Cookson’s, and 50,000–75,000 Nyassa. However, the latest estimates of Eastern White-bearded Wildebeest indicate a steep decline in the subspecies’ populations to a current level of perhaps 6000–8000 animals.While the Common Wildebeest remains one of Africa’s most abundant game species at present, these figures highlight both the significance of the migratory Serengeti population and the vulnerability of the species to further adverse developments, such as those that 535

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have affected the Eastern White-bearded subspecies during the last decade (Estes & East 2009). Maximum densities of 35/km2 have been recorded in the Serengeti and Ngorongoro Crater populations (Runyoro et al. 1995), although the Ngorongoro population density was closer to 55/km2 up until 1980 (mean of 14,000/250 km2; Estes 2002a). Elsewhere, population densities estimated by aerial surveys are lower: less than 0.15/km2 in areas such as Kafue N. P. (Zambia), Hwange N. P. (Zimbabwe) and Etosha N. P. (Namibia) and the southern Kalahari, to 0.6–1.3/km2 in Kruger N. P. (South Africa) and North Luangwa N. P. (Zambia), Selous G. R. and Kajiado (see East 1999). Adaptations The size, proportions and shape of the Common Wildebeest are all adapted to the species’ migratory habits, including the ability to sustain a canter with minimal energetic cost, facilitated by high shoulders and slender legs (Alexander 1977, Pennycuick 1979, Christiansen 2002). Common Wildebeest have a number of physiological adaptations including the ability to minimize water loss by allowing body temperature to rise in hot weather and by seeking shade (Taylor 1970a, Ben-Shahar & Fairall 1987). When body temperature rises above ca. 40 °C, animals dissipate heat by nasal panting. However, the Black Wildebeest is more heat- and cold-tolerant than Common Wildebeest, and Hofmeyr (1981) found the latter to be at a disadvantage compared with Hartebeest Alcelaphus buselaphus, as its thin pelage and dark skin fail to provide significant protection against solar heat gain. In areas

Common Wildebeest Connochaetes taurinus albojubatus head and myology.

devoid of shade, Common Wildebeest orientate their bodies to minimize insolation; they also face into the wind (Berry et al. 1984). Common Wildebeest are considered water-dependent, able to go without drinking for several days on green pasture, but no more than two days while subsisting on dry grass. Daily requirement has been estimated at 10 litres under arid conditions (Taylor 1968b). However, in Botswana’s Kalahari Desert, Common Wildebeest go for months without drinking in sandveld areas where local rains have produced abundant plant growth such as tsama melons Citrullus vulgaris and tubers (Estes & East 2009). This suggests greater adaptability to environmental conditions than commonly supposed. The rumino-reticulum of the Common Wildebeest is comparable with that of African Buffalo Syncerus caffer or cattle in proportion to size, with average capacity of 40 litres. Its wide muzzle, wide incisor arcade and flexible lips are adapted for non-selective bulk grazing, preferably on grasses in an early growth stage, between 3 and 10 cm, and green with more leaf than stem (Talbot & Talbot 1963, Bell 1970, Hofmann 1973, Gordon & Illius 1988). Foraging and Food Classed as a variable grazer by Gagnon & Chew (2000) in their review of the dietary preferences of African bovids, and studies using stable carbon isotopes suggest that Common Wildebeest have such a high component of grass in their diet that they are best considered as hypergrazers (Cerling et al. 2003, Sponheimer et al. 2003b); Hofmann & Stewart (1972) classify them as a fresh-grass grazers dependent on water. During the wet season in the Serengeti ecosystem, Common Wildebeest concentrate on the highly nutritious, productive shortgrass plains, which sustain very high herbivore densities. In the dry season they pursue a rotational-passage grazing system as they move rapidly through medium and tall grassland in search of green pastures (McNaughton & Banyikwa 1995, Wilmshurt et al. 1999b). Grasses with the most leaf and least stem are selected. A comparative study of selectivity and feeding rate of Common Wildebeest, Hartebeest and Topis Damaliscus lunatus jimela in Serengeti N. P. showed that the Common Wildebeest has the fastest bite rate on swards with low biomass and high protein content of green leaf (Murray & Brown 1993). Preferences for species and growth stages vary depending on season and availability and in different populations and ecosystems (Berry 1980, Ben-Shahar 1991, Ben-Shahar & Coe 1992, Murray 1993, Bodenstein et al. 2000, Ego et al. 2003). Shallow-rooted colonial species that carpet the ground are preferred (Bell 1970). Favourites include Cynodon dactylon (probably most important overall; Andere 1981), Brachiaria brizantha, Sporobolus marginatus, Cenchrus ciliaris, Eragrostis superba, Digitaria milanjiana, D. macroblephara and small sedges in short grassland (e.g. Kyllinga). Many other species are eaten in early growth stages or when preferred decreaser species are rare, such as Themeda triandra, Chloris gayana, Pennisetum mesianum and Panicum maximum. Food availability is often limited in the late dry season following extensive wildfires. Forty years’ worth of data collection prove conclusively that the Serengeti population is limited by food supply (Mduma et al. 1999). Nomadic/migratory movements are discussed further under Social and Reproductive Behaviour. Social and Reproductive Behaviour The following account is based mainly on the author’s observations of C. t. mearnsi in

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Ngorongoro Crater and Serengeti N. P. between 1963 and 2006, supplemented with information from other populations as indicated. The Common Wildebeest is a highly social species, with a grouping and mating system based on male territoriality. Migratory populations move in vast herds and aggregate in dense concentrations on green pastures. Actively territorial !! segregate yearling and older non-territorial !! from "" and young, making three distinct social classes: female or nursery herds of "" and young; all-male (bachelor) herds; and territorial !! – the latter the only class normally ever found alone. The Common Wildebeest has two related but different dispersion patterns: (a) sedentary/dispersed and (b) migratory/aggregated. The former is representative of most social, territorial antelopes; the latter, derived state is adapted to migratory habits. Depending on environmental variables, one form can change into the other, either way. In the sedentary/dispersed state, nursery herds of "", calves and attached (mainly female) yearlings occupy preferred grazing grounds, which competing bulls divide into a network of individual territories. The intolerance of breeding (= territorial) !! keeps bachelors in less favourable, lightly or undefended habitat. Female herds, comprised of 2 to ca. 25 cows and young (with a mean number of ten in Ngorongoro Crater), represent semi-exclusive associations derived from the tendency of "" with young calves to band together, and also due to durable mother–daughter bonds (which persisted two years in the case of a marked barren mother; Estes 1966). Strangers attempting to join resident herds in the Ngorongoro Crater were generally rejected, chased out by the attendant bull if treated with hostility by the cows. Each herd acted largely independently and remained within a restricted home-range, often less than 100 ha during the wetter half of the year (Nov–May), encompassing the territories of four or five resident territorial !!. Captive migrating "" kept in a 25 ha enclosure behaved in the manner of resident herds, including establishment of a dominance hierarchy (A. Moss pers. comm.). In Ngorongoro Crater, as pastures stopped growing after the rains ended, known female herds began to aggregate on the remaining green pastures. At the end of the day, each herd returned to its accustomed home-range (Estes 1966). However, as the dry season intensified, the small herds stayed in aggregations on the remaining greenbelts and within a few weeks apparently lost their separate identity. Nevertheless, when the short rains rejuvenated the Crater pastures, some of the known animals returned to the same homeranges (Estes 1966, 1969, 1976). Bachelor herds include !! of all ages from yearlings to old bulls, with herd sizes ranging from two to thousands, depending on the size of the population and openness of the habitat: the more open the habitat, the larger the herds – as usual in social ungulates (Estes 1974, 1991a). Competition between bachelors is much less spirited than in male herds of Topis, Hartebeest and most other African antelopes (Estes 1969). This relatively unaggressive behaviour contrasts with the behaviour of territorial bulls. Males mature and become territorial at 5–6 years of age (Watson 1969). Rarely, a bull of three years can participate (briefly) in the rut (Estes 1966, 1969). In resident populations, bulls may hold the same territories for years, either continuously or discontinuously, depending on variable grazing conditions and proximity of "" (Talbot & Talbot 1963, Estes 1969). In Ngorongoro, average spacing between bulls is roughly 60 m in areas frequented by small herds, but even there bulls were alone 80% of the

time (Estes 1969). Except for the rut, territorial competition is most intense early in the rains (Nov/Dec) when rangeland abandoned in the dry season is reoccupied by "". The fact that mating opportunities are virtually non-existent so near the calving season suggests that the psychological advantage of established ownership promotes yearround occupancy where competition is strong. Serious fights mostly involve attempts to establish a territory, which territorial neighbours fiercely resist. Rather than contest the territory of an established bull, a newcomer’s best strategy is to win rights to a small plot between territories by wearing down the resistance of the neighbours, and then gradually enlarging it. Creating a ‘stamping ground’ is one of the first acts in laying claim to a space. It is created by pawing, kneeling and horning the ground, defecating, lying and rolling. These bare patches, centrally located and strewn with dung, are the olfactory foci of the territory; there may be several on a property, but usually one where the bull spends most of its inactive time. Female herds when present also frequently rest on the stamping ground. Fights between established bulls are uncommon, but neighbours engage in elaborate daily challenge rituals that serve to reaffirm the fitness of each bull to hold property. The ritual includes mutual urine testing with the vomeronasal organ. Male–male urine testing is highly unusual, as the primary function of this organ in ungulates is to detect female reproductive status through assaying hormone breakdown products in their urine (Estes 1969, 1972, Krieger et al. 1999).When a territorial bull leaves his property, he reverts to the status of a bachelor male as soon as he has passed beyond the territories of his immediate neighbours (Estes 1969). The mobile/aggregated state is adapted to migratory and nomadic movements in extensive savanna ecosystems with a rainfall gradient from ca. 200 to 800 mm. Wilmshurst et al. (1999b) found that movements of Common Wildebeest are broadly similar to those of other large herbivores that migrate in response to resource gradients. In Serengeti N. P., the short, colonial grasses found in the lowerrainfall areas are kept growing by frequent rain and fertilized by herbivore concentrations, whose manure is buried overnight by hordes of dung beetles. Equally important, colonial grasses only 2–4 cm high virtually carpet the ground (Bell 1970, McNaughton 1984), unlike the medium and longer grasses in the higher-rainfall areas where wildebeest circulate in the dry season seeking swards of intermediate-height and greenness that offer the optimal combination of bulk and digestibility (Fryxell 1991, Morrison & Bolger 2012). Wolanski et al. (1999) suggest that excessive salinity triggers the migration of Common Wildebeest and Plains Zebras off the Serengeti short-grass plains. Eight Common Wildebeest collared and monitored for 1086 wildebeest days in the Serengeti ecosystem moved an average 4.47 km/ day (S.D. 5.26 km, maximum in one day 57.5 km). Daily movements increased from Mar–Jun with a peak in May (Thirgood et al. 2004, Hopcraft 2010). The structure and composition of aggregations vary according to habitat conditions, season, time of day, prevailing activity and level of territorial behaviour. The tendency of Common Wildebeest to associate in peer subgroups (see later) means that distribution is not random – which makes sampling to determine sex ratio and recruitment difficult, even though immature age classes are distinct (!! to 3–4, "" to 2.5 years). Most samples of the Serengeti population suggest an equal adult sex ratio (Watson 1967, 1969, 1970, Sinclair 1979, Mduma 1996), whereas samples of resident populations generally indicate a female bias in the order of 1 : 1.5 or 1 : 2 (R. D. 537

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Jun–Jul

Oct–Nov

Dec–May

The annual migratory route of eight female Common Wildebeest Connochaetes taurinus in the Serengeti-Mara ecosystem (each series of coloured dots represents the movement of one animal tracked using a GPS collar). Common Wildebeest retreat to the Mara River in the northwest during the dry season, often detouring via the (unprotected) Western Corridor on their way. Green corresponds to Serengeti National Park (Tanzania) and Masai Mara National Reserve (Kenya); yellow corresponds to surrounding buffer zones (map courtesy J. Grant, C. Hopcraft, Frankfurt Zoological Society and the University of St Andrews).

Estes pers. obs.). Common Wildebeest aggregations on the Serengeti short-grass plains are typically dispersed with wide individual spacing. But when feeding in tall swards, aggregations are compact with minimal individual spacing, possibly from fear of Lions Panthera leo. Except during several months of the dry season when territorial activity is attenuated, mature bulls accompanying the migration space out and become active as soon as an aggregation stops moving. In short order they round up "" and weed out the bachelors, enforcing the same kind of segregation as in resident populations: nursery herds contained in a territorial network with bachelor !! on the periphery. Such groups have been called ‘pseudo-herds’, as the association of "" is merely temporary (Watson 1969). Only cows with young of the year keep together. They begin to separate after nine months, but female and some male yearlings are frequently associated with nursery herds, at least some of which continue to trail their mothers. When on the move in files or columns, both sexes and all ages join together. However, when the movement is halted by an obstacle (river crossing, tall vegetation, defile), bulls come to the forefront and are the first to risk going forward (R. D. Estes pers. obs.).When CommonWildebeest are in tree savanna they cluster in the shade during the hottest hours. At night, aggregations gather on the shortest available grassland and rest/ruminate in ‘bedding formations’, which may contain hundreds of animals but being linear, with enough separation to permit individuals to move in or out, can disperse immediately when alarmed, as when tested by hunting hyaenas (Estes & Estes 1979). Territorial !! are capable of reproducing at any time and for a good six months of the year have continuing but declining mating

opportunities, as some 20% of "" fail to breed during the rutting peak; a few breed as much as five months later. The onset of the rut is marked by a peak of noise and activity. No other African wildlife event can match the spectacle of the Serengeti rut, which occurs during the migration from the short-grass plains into the woodland zone. An idea of the intensity of sexual competition can be gained from the number of territorial bulls in dense concentrations: up to 250/km2 (R. D. Estes pers. obs.). This remarkably close packing of territorial bulls facilitates synchronous breeding of cows in migrating hordes: no " in oestrus is likely to go undetected for long. The rutting uproar continues day and night, reaching a crescendo when aggregations are on the move. As long as "" are present, territory holders neither rest nor feed but exert every effort to detain "", prevent raids by their nearest rivals, and seize opportunities to break up a neighbour’s herd. Herding bulls do not engage in challenge rituals, but frequently butt heads. Tightly clustered herds within a network of single bulls is a hallmark of the rut clearly visible from an airplane or hilltop. Considering that up to half-a-million cows are mated during the rut, it is surprising how seldom copulations are witnessed. By far the most effort exerted by territorial bulls goes into herding, chasing and fighting. Their efforts may look, but are not, futile, as having a herd improves a bull’s chances of gaining a cow in heat, because "" suffer less sexual harassment by going into a group rather than stopping with a lone ! (R. D. Estes pers. obs.). The more cows a bull encounters, the more mating opportunities. Bulls that establish territories in the main stream of a migrating horde obviously are fitter than bulls with peripheral territories where most passers-by are bachelor !!. Good locations also vary with time of day or night. Bulls under a broad shade tree may have up to 100 "" and young during the hottest hours while unshaded neighbours have a few or none. Bulls that stay put on territories in a close-cropped, abandoned pasture may be rewarded come evening, when an aggregation returns to spend the night on short grassland. In the hurly-burly of the rut, it is not unusual for a bull to overlook the presence of an oestrous " in his herd, but having detected one, a bull copulates with her dozens of times, up to three or four mounts with ejaculation in a minute when not distracted by the need to repel invading neighbours. A preference to continue mating with the same bull is shown by "" in heat, which will follow and even solicit copulation. But if forced to leave, oestrous "" have been seen to copulate with a succession of bulls until no longer receptive (Estes 1991a). Oestrus lasts at most one day. While the energetic costs of rutting are rigorous for actively competing bulls, interludes when no "" are in the immediate vicinity enable them to rest and feed. Bulls that have been active over a sustained period may take a day or two off in a bachelor herd (R. D. Estes pers. obs.). Accordingly, the Common Wildebeest rut leaves breeding !! less spent than ungulates of the Northern Temperate Zone that persevere to exhaustion. However, serious injuries – to hooves, legs, horns, eyes – incapacitate possibly 0.05% of the competing bulls. Those with broken horns can hold their own as long as they have the bosses to absorb the impact of butting. But a bull that has lost a whole horn can no longer stand up to an intact rival. The end of the rut is marked by reduced intensity and volume of calling and by the presence in resting herds of adults of both sexes. The fat deposits bulls accumulate in the months prior to the rut are depleted by the end of it (Sinclair 1977b).

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The fact that 80% of the annual calf crop is born within three weeks is linked to the evolution of ‘follower young’, a key adaptation to the mobile/aggregated state of migratory populations. Apart from the two wildebeest species and the Blesbok/Bontebok Damaliscus pygargus, the young of all other antelopes go through a hiding or lying-out stage of varying length. Selection for follower young born in a short time stems from the Common Wildebeest habit of massing on short, green grass. Tan calves are conspicuous among dark-coloured adults. As aggregations rapidly deplete pastures and move on, mothers guarding ‘hider calves’ adapted for concealment in tall, tan grass would be left behind. Changing to follower young led Common Wildebeest to lose all aspects of the ancestral hider strategy. Calves have lost the hiding instinct and accompany their mothers as soon as they can stand. This takes as little as 3 min (mean of 7 min), and the feeble stage when neonates cannot keep up with a herd is over within about two days (Estes 1976, Estes & Estes 1979). A drastic speed-up in the development of locomotory skills is key to this precocity. Common Wildebeest calves have almost adult muscle/bone length ratios at birth, whereas hider calves have only two-thirds the adult muscle/bone ratio at birth (Grand 1991). Common Wildebeest calves may be the most precocious of all ungulates. Actual parturition is described in detail in Estes & Estes (1979). Mothers only lick their calves in the first hour postpartum; neonates neither receive nor need the stimulation of anogenital licking to void wastes. Mothers recognize their newborns by scent, and aggressively reject any but their own (Estes & Estes 1979). Calves may be seen to eat dung within a few days, presumably acquiring rumen bacteria. They start sampling grass within a week. Within a day or two mother and calf are mutually imprinted. Calves rest in crèches or lie alone while their mothers feed. Some mothers leave sleeping calves to join wildebeest filing to water.This practice came to light while following Ngorongoro cows returning from drinking, which were acting as though they had lost calves – running and bawling, sometimes joining forces and inspecting every herd with calves. After going some 3 km in this manner, several distraught mothers found their calves – just where they had left them. Up until then, a lone wildebeest calf was assumed to be orphaned and doomed (R. D. Estes pers. obs.) Associative behaviour is manifested within days of birth, in crèche formation and group play (chasing, cavorting, head-butting). Beginning as yearlings, peer subgroups of the same sex and age are discernible even in huge Serengeti aggregations. The annual calving season promotes further subdivisons into groupings of "" in the same reproductive state. In the months before and after calving the following subgroupings can be observed, in addition to the aforementioned divisions: pregnant "", maternity groups of cows with calves, non-pregnant "", and cows that have lost calves (both these often found grouped in male aggregations) (Estes & Estes 1979). While resting, inter-individual spaces of approximately one metre keep herd members beyond horn reach. Only mother and calf lie in contact. Star-formations can be seen in small herds that enable members to monitor a full 360 degrees. Such circles form through settling back-to-back facing away rather than head-to-head. Bachelor herds can be discerned at a distance by their more regular individual spacing. Rubbing the muzzle or head on the rump or shoulder of another wildebeest, commonest between "", is also performed during the territorial challenge ritual. It appears sociable, although

these contacts apparently often include depositing secretions of the preorbital glands, which might be an assertion of dominance (Estes 1969, 1991a). Comfort behaviours such as lying down and rolling (rare or unknown in other ruminants) are most commonly seen in territorial bulls (though Common Wildebeest cannot roll over completely like zebras). Common Wildebeest also have the habit of rubbing their preorbital glands on branches and trunks of trees, usually as the prelude to horning bouts. Olfactory communication is involved, as marking and horning are directed especially at sites that have already been marked and horned; and bouts begin with sniffing of the site. The behaviour is also infectious as Common Wildebeest will line up and take turns rubbing and thrashing the same tree. However, vegetation horning is primarily an activity of adult !!; other animals horn much less often and vigorously. Bouts may last anywhere from 15 seconds or less to as long as five minutes. It is clearly aggressive in character and may either be directed, as in encounters between territorial bulls, or an undirected display of aggressive mood. What is extraordinary and generally overlooked is the environmental impact of Common Wildebeest horning in large populations. Data collected on the incidence and effects of horning beginning in 1979, in 1980–81, 1986, 1998 and 2003 show that on the Serengeti Plains and in tree savanna of the woodland zone two out of three young trees of suitable size and accessibility have been horned, many repeatedly over a period of years. In numerous cases the stems and branches of trees (especially Acacia spp. and Balanites aegyptica) have been destroyed every time they produced stems of thrashable size; the resulting growth in the case of A. tortilis, a dominant tree, is a supine hedge, within which a main stem of up to 10 cm diameter may be found. The evidence suggests that destruction of woody vegetation by Common Wildebeest was second only to fire and elephants in transforming Serengeti savanna woodlands into tree grassland during the 1970s and 1980s (Estes et al. 2008). In undisturbed herds or aggregations of Common Wildebeest, the advertising calls of territorial bulls are the source of virtually all vocalizations. Calling by other classes is common in moving aggregations and consists mainly of exchanges between mothers and their offspring, which reach a crescendo when they become separated, as when massed at waterholes, crossing terrain that funnels many animals close together, or when stampeded by predators. Whereas differences in the advertising calls of C. t. mearnsi are discernible from other subspecies, the higher-pitched calls of other classes sound the same.Yet, mothers and dependent offspring recognize one another’s calls. It enables nearly all separated pairs to reunite, facilitated by remaining at the separation site and running back and forth bawling (R. D. Estes pers. obs.). With a dozen or more ungulate species in savanna herbivore communities, the Common Wildebeest inevitably is often found in the same place and time with some of them, especially with other grazers. The Plains Zebra Equus quagga, which eats many of the same grasses (Bell 1970, Casebeer & Koss 1970, Ben-Shahar & Coe 1992) is popularly believed to be closely associated, and indeed it is common to find aggregations that include both species. Research on the incidence and basis of association between Common Wildebeest and eight other grazing ungulates (Plains Zebra, Grant’s Gazelle Nanger (granti) granti and Thomson’s Gazelle Eudorcas thomsonii, Topi, Hartebeest,Waterbuck Kobus ellipsiprymnus, Impala Aeypceros melampus and Common Warthog Phacochoerus africanus) of the Serengeti 539

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Common Wildebeest Connochaetes taurinus albojubatus.

ecosystem found a large overlap in distribution with the Common Wildebeest and that most species were eating the same plants in the same vegetation types (Sinclair 1985). Most species were found closer to Common Wildebeest than expected if distribution was random. In the wet season, 71% of the Plains Zebra population could be found near the main Common Wildebeest concentrations, but on the leading edge where they could graze before the sward was eaten down by the latter. But in the dry season, after migrating to the Masai Mara part of their range, the Plains Zebras avoided inter-specific competition by staying further away from the Common Wildebeest and grazing in taller grassland. The results of this study strongly suggest that the basis of the association is mutual protection against predators. As the Common Wildebeest is the preferred prey of all the large carnivores, the other species are safer in proximity to this most abundant herbivore. The conclusion: ‘In general, predation appears to play as important a role as inter-specific competition in structuring this community’ (Sinclair 1985: 916; and see Hopcraft 2010). Despite proximity, behavioural interaction and communication with associated ungulates are surprisingly limited. They get out of one another’s way and respond to each others’ alarm signals, but largely ignore their species-specific displays. Most ungulates react to the alarm signals of other species, including monkeys and baboons, jackals Canis spp., oxpeckers Buphagus spp. and other birds. Reproduction and Population Structure The timing of calving and rutting is geared to the climate so that both occur under favourable conditions (Sinclair et al. 2000). Onset of the annual rut varies from year to year by as much as three weeks, under the influence of variable climatic conditions. For example, following the great East African drought of 1960, when most cows lost their calves, C. t. mearnsi and C. t. albojubatus calved two months earlier than normal; Estes 1966). In East Africa, the rut comes after the rains (Jun) when the animals are in top condition, thus ensuring an adult conception rate normally better than 90%. Some "" breed as yearlings (ca. 20% in the Serengeti ecosystem), but most conceive a year later (ca. 28 months). In the 1960s and 1970s, while the

Ngorongoro and Serengeti populations were increasing at the rate of 10% a year, the yearling conception rate was closer to 80% (Estes 1966, Watson 1969, 1970, Estes & Estes 1979, Sinclair et al. 2000). Research conducted in the Grumeti Reserves bordering Serengeti N. P. in 2002–2004 on 17 captive "" (Monfort et al. 2001) yielded new information about the reproductive physiology of the species (Clay 2007, Clay et al. 2010). Regular assays of faecal hormones excreted by each " revealed their reproductive-endocrine rhythms and determined that CommonWildebeest are seasonal, polyoestrous, spontaneous ovulators. The duration of oestrous cycles and luteal phases were quantified, and a gestation period of eight months was confirmed (Clay et al. 2010). The average duration of postpartum anoestrus was 100 days. No significant correlation was found between lunar phase and the timing of luteal phases as proposed by Sinclair (1977b). How the oestrous cycles of "" are synchronized has long been a subject of speculation (Rutberg 1987, Sinclair 1977b, Sinclair et al. 2000). A major objective of this study was to test the author’s hypothesis that chorusing of bulls, which reaches a peak during the rut, serves this function (R. D. Estes pers. obs.). A month prior to the 2003 Serengeti rut, recordings of rutting calls were played to two groups of captive cows continuously for three weeks, one of which included a bull, while a third control group was kept isolated. Faecal progestin assays showed that the oestrous cycles of the two exposed groups were synchronized more closely than in the control group. This research provides the first evidence of an auditory cue that modulates reproductive timing in the Bovidae (Clay 2007). Calving occurs typically ca. two months before the period of most rain and maximum grass production (Talbot & Talbot 1963). Common Wildebeest calving is more tightly synchronized than that of any other African ungulate, comparable to the birth peak of Caribou Rangifer tarandus (Dauphiné & McClure 1974). The Serengeti migratory population usually calves in Feb, between short and long rains, but the resident population in the Western Corridor is subject to the wetter climate associated with L. Victoria and calves 1–2 months earlier. In southern Africa, the calving season is generally Nov/Dec (though sometimes later; Attwell 1977), but in

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Liuwa Plains, SW Zambia, calving occurs in late Oct to early Nov just before the rains start (Howard & Conant 1983). Calves weigh between 14 and 25 kg at birth, with calves in southern Africa being larger (Talbot & Talbot 1963, Attwell 1977). Maternal stress is considered to be most severe during early to midlactation (Eltringham 1979, Altmann 1980, Murray 1982b). During the abbreviated feeble stage, calf survival depends on the presence of harderto-catch calves a few days older, which serve as cover for newborns.The more calves in a maternity herd, the higher the calf survival rate. Survival in the first month averages ca. 70% in migratory aggregations vs. 50% or less in resident small herds (Estes 1976). A calf’s only defence is its own mother, which is quite effective against a single Spotted Hyaena Crocuta crocuta (Estes & Estes 1979).Young of the year begin to form separate groups at about nine months, following weaning at around eight months (Watson 1967). However, some continue to follow their mothers until the next calf crop is born. The potential payoff is that cows that lose calves allow their yearlings to nurse. Estimated calf survival to yearling stage in the Serengeti population in nine years between 1965 and 1994 averaged 22.9% (12.3–35.1) (Mduma 1996, Appendix 2). Weigl (2005) gives a longevity record for the species in captivity of 24.3 years, which is about five years longer than maximum longevity recorded by Attwell (1977) in KwaZulu–Natal (1982). But the oldest animals recorded in Etosha N. P. were 13–14 years (Berry 1980) and individuals of over 12 years in Serengeti N. P. were considered very old (Mduma 1996). For information on sex ratios see Social and Reproductive Behaviour. Predators, Parasites and Diseases Common Wildebeest are typically the preferred prey of Lions and Spotted Hyaenas wherever they occur (Kruuk 1972, Schaller 1972, Elliott et al. 1977, Mills & Shenk 1992, Scheel 1993b).The fact that they tend to be the dominant large herbivore in savanna ecosystems may largely explain this preference. But there is also quantitative evidence that wildebeest are less vigilant than most associated prey species, including Plains Zebras, the second most-preferred prey (Scheel 1993b). Aggregations are easier to approach than singles and small herds, probably because each individual feels and is safer in a large group. Observations of scanning rates showed that rates declined with increasing herd size, including herds containing individuals of different species (e.g. zebras) (Scheel 1993a). Scanning rates were also influenced by light and cover. Common Wildebeest scanned more at twilight and on moonlit nights than during the day. Scanning rate also correlated with nearby cover large enough to conceal hunting Lions. Greater vigilance, as indicated by increased scanning rates, would be expected in bushy habitat and tall grass, but oddly enough, wildebeest scanned less in such places than in more open habitat. Likewise animals in the interior of a group tend to scan less than those on the periphery. This is usually explained by the fact that animals on the outside are more vulnerable (Elgar 1989, FitzGibbon 1990a), but having their view blocked by other animals could explain reduced scanning rates of individuals inside a herd. Similarly, thick cover that renders the ability to detect approaching predators problematic could explain reduced scanning rates in such habitats (Scheel 1993a). Of eight species in the study, Common Wildebeest and Common Warthogs were the least vigilant, and both were preferred by Lions. Hence these species were at greatest predation risk (Scheel 1993b). However, Common Wildebeest and Plains

3 months

4 months

10 months

16 months

24 months

Common Wildebeest Connochaetes taurinus horn development.

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Family BOVIDAE

Zebras were considered equally vulnerable to Lions in Kruger N. P., and no selection by season or sex was noted (Mills & Shenk 1992). Although the hunting technique of the Spotted Hyaena is adapted to selecting the most vulnerable individuals, a single adult hyaena is capable of running down and killing a healthy full-grown Common Wildebeest (Wasson 1990). The hyaena’s specialization on newborn calves during the calving season is a major factor in maintaining the short peak, as mortality of calves born before the peak is virtually total (Estes 1976). Cheetahs Acinonyx jubatus and African Wild Dogs Lycaon pictus as well as Lions partake of the birth bonanza. Jackals very rarely prey on wildebeest calves. The birth of thousands of calves during the short peak completely gluts the predator market (Rutberg 1987, Ims 1990, Sinclair et al. 2000). The infamous pan-African rinderpest epizootic of the 1890s, a virus transmitted from cattle brought into Somalia around 1887 during the Italian campaign, caused the mortality of up to 90% of the continent’s cattle, African Buffalo, tragelaphines Tragelaphus spp. and wildebeest. No other livestock or wildlife disease has had such widespread mortality, and the survivors of the rinderpest pandemic, with at least partial immunity, mostly recuperated within one to two decades (Simon 1962). Called the ‘yearling disease’ because older Common Wildebeest calves were vulnerable to rinderpest in lean years (Talbot & Talbot 1963), rinderpest disappeared from the Serengeti population in 1963 (Dobson 1995). The population then increased five-fold from 250,000 to 1,250,000 by the mid-seventies (Sinclair 1979). Apart from such rare epidemics, studies of Common Wildebeest and African Buffalo suggest that disease rarely kills animals in good body condition. But disease, like predation, can cull animals in poor condition (Sinclair 1977a, Prins 1996). Common Wildebeest are carriers of malignant catarrhal fever (sometimes referred to as ‘snotsiekte’), a viral disease fatal to cattle. The Masai have to keep their herds away from the short-grass plains during and after the Common Wildebeest calving season, to avoid pastures contaminated by afterbirths (Homewood et al. 1987). Anthrax, an endemic disease of African wildlife (Prins 1996), most prevalent in arid ecosystems, has been implicated as a major cause of the decline of the Common Wildebeest population in Etosha N. P. (Berry 1993). Protecting cattle from foot and mouth disease has been the justification for the veterinary cordon fences criss-crossing Botswana in total disregard of their effects on the wildlife, so that the ruling oligarchy of cattle barons can continue to profit from the European Union (EU) subsidized export of beef to Europe (Williamson & Williamson 1985a, Williamson 1994b; and see Conservation). Cleaveland et al. (2005) isolated bovine tuberculosis, caused by Mycobacterium bovis, from two of 18 migratory Common Wildebeest samples in the Serengeti system in 2000. Similarly, healthy animals are hardly affected by external and internal parasites, but can be further weakened when in poor health. Ticks are the most serious external parasites, and botflies (oestrid flies) are common parasites of Common Wildebeest throughout their range (Howard & Conant 1983). Larvae of Gedoelstia spp. are typically deposited on the cornea of the eyes, and then make their way to the nasal/sinus cavities via the ocular-cranial route or ocular-vascularpulmonary route. Larvae of Oestrus spp. are deposited in the nostrils. The larvae develop to mature larvae in the sinus cavities and then crawl out to pupate. Botflies do not seem to cause Common Wildebeest any harm, although Gedoelstia larvae have been found in the braincase and on the dura mater (Talbot & Talbot 1963, Horak et al. 1983b). Horak

et al. (1983b) recovered parasites from 55 Blue Wildebeest in Kruger N. P. and recorded 13 nematodes (particularly Haemonchus bedfordi), four cestodes, one trematode, the larvae of five oestrid flies, three lice, seven ixodid ticks, one mite and the nymphae of a pentastomid species (Linguatula nuttalli, indicative of the large number of Lions in Kruger N. P., the final hosts of the pentastomid). Remarkably, they recovered very few adult ticks of any species. Only one species, the lungworm Dictyocaulus viviparous, was attributed to any ill-effects observed in Common Wildebeest, causing fairly extensive pulmonary lesions, though seemingly not severe enough to cause death. Conservation IUCN Category: Least Concern. CITES: Not listed. Reductions in historical geographic range have been relatively minor, with only the elimination of C. t. johnstoni from Malawi being perhaps most notable. However, all the major Common Wildebeest populations except the Tanzania populations of C. t. mearnsi and johnstoni have undergone declines, some by as much as 90%. Fences that blocked migration between wet- and dry-season ranges have had the most obvious impact. Denying access to water and to higher-rainfall refuges during severe droughts have caused mass die-offs. But many other anthropogenic factors have taken their toll: ever-expanding settlement, mechanized agriculture, overstocking with cattle and habitat degradation, elimination of water sources through watershed deforestation and expropriation for irrigation, poaching, bushmeat trade, armed conflicts – and last but not least, game eradication programmes in failed efforts to eliminate the wild hosts of sleeping sickness (nagana) and other diseases of domestic livestock. The decline in numbers and episodes of mass mortality of Common Wildebeest in Botswana caused by veterinary cordon fences that blocked drought-induced migrations received considerable notoriety, particularly after thousands died at L. Xau in the north-east of the Kalahari Desert in 1980 (Owens & Owens 1980), followed by other large die-offs in subsequent years (Williamson & Williamson 1985a, Parry 1987, Estes & East 2009, Gadd 2012). But Spinage (1992) documents historical evidence that the decline actually began much earlier. Archive records show that there was at one time a significant migration every winter east and south-east from the southern Kalahari to the Molopo R. into South Africa through the unfenced boundary between Tsabong and Khuis (with Common Wildebeest moving as much as 40 km per day). Wildebeest here were once so numerous that they were regarded as a menace by local farmers because they competed with local cattle for grazing and transmitted malignant catarrhal fever. Large-scale killing of Common Wildebeest followed, at least until 1961, when the species was classified as a game animal that could only be hunted with a licence. Fencing of national park boundaries caused Common Wildebeest populations to crash in Kruger N. P. (Whyte & Joubert 1988) and Etosha N. P. (Berry 1997). Mechanized farming, cattle ranching and fencing of the Loita Plains wet-season range of the Masai-Mara population of C. t. mearnsi led to a decline of 75% between 1977 and 1997 (Homewood et al. 2001, Ottichilo et al. 2001, Serneels & Lambin 2001, Ogutu et al. 2011). Progressive fencing of the Athi-Kapiti Plains keeps reducing the range available for wildlife; the Kitengela corridor between the plains and Nairobi N. P. is now so tortuous that Common Wildebeest rarely come into the park, which was formerly an important dry-season concentration area. Completing the fencing of the park by closing the connection to the Athi-Kapiti Plains is now advocated by many (Cowie 2004).

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The migratory population of theWesternWhite-beardedWildebeest defines, and is largely conserved in, the Serengeti ecosystem, including Ngorongoro Conservation Area and the Masai Mara National Reserve. Thirgood et al. (2004) radio-tracked eight Common Wildebeest during 1999–2000 in relation to protected area status in different parts of the ecosystem, and found that the collared animals spent 90% of their time within well-protected core areas. However, two sections of the migration route – the Ikoma Open Area and the Mara Group Ranches – currently receive limited protection and are threatened by poaching or agriculture. Comparing current Common Wildebeest migration routes with those recorded during 1971–73 indicates that the western buffer zones appear to be used more extensively than in the past.Thirgood et al. (2004) suggest that the current development of community-run Wildlife Management Areas as additional buffer zones around the Serengeti represents an important step in the conservation of this UNESCO World Heritage Site. The Serengeti migration was seriously threatened in 2010 by the proposed construction of a paved highway from Musoma on L. Victoria to Tanga on the Indian Ocean coast across the northern part of Serengeti N. P. Intended to transport oil, minerals and produce to the coast from landlocked countries to the west and north, this road would have carried thousands of vehicles a day and permitted roadside development along the route, threatening sooner or later to truncate the migration. International outcries by scientists (e.g. Dobson et al. 2010; and see Holdo et al. [2011], who predicted a decline in the wildebeest population by one-third), conservationists, tour operators, UNESCO and IUCN finally persuaded President Jakaya Kikwete to accept an alternative route south of the Park and to construct only a gravel road across the northern Serengeti, as proposed in the original 2005 plans. However, in Sep 2011, the Minister of Transport announced plans for a railway across the Serengeti along the same route, pledging that, despite opposition, the project will be completed or reach advanced development within the next five years. Priorities for research and conservation in the future include: (1) identifying, restoring and maintaining access to water and drought refuges for Common Wildebeest, zebras and other water-dependent herbivores; (2) maintaining corridors and dry season refuges, and promoting Transfrontier Conservation Areas (TFCA) that enlarge and protect viable ecosystems; (3) applying pressure from national and international conservation organizations to stop the EU from subsidizing beef exports from Botswana to Europe at above-market prices (Williamson 1994b); and (4) enforcing separation of Black and Blue Wildebeest introduced on ranches in South Africa, Botswana and Namibia to prevent hybridization. Existing TFCAs of potentially great importance for restoring Common Wildebeest populations to something approaching their former abundance include: the Kavango–Zambezi TFCA, spanning an area of approximately 287,000 km² at the confluence of Namibia, Angola, Zambia, Zimbabwe and Botswana and including the Caprivi Strip, Chobe N. P., the Okavango Delta and the Victoria Falls; the Great Limpopo N. P. connecting Zimbabwe, Mozambique and South Africa, facilitating the doubling of the size of Kruger N. P. by adding an adjoining section of Mozambique and the Gonarezhou N. P. in Zimbabwe; and the Kgalagadi Transfrontier Park, which combines South Africa’s Kalahari Gemsbok N. P. with Botswana’s Gemsbok N. P. Serengeti wildlife would also benefit from the proposal to incorporate the Mara range in a TFCA. Efforts are also under way to

link Selous G. R. in S Tanzania with Niassa G. R. in N Mozambique via Selous–Niassa Wildlife Corridor. Another promising conservation initiative is establishment of private wildlife conservancies adjoining and extending protected areas, as exemplified by Sabi Sand G. R., and the Timbavati and Klaserie Private Nature Reserves on the western boundary of Kruger N. P. Measurements Connochaetes taurinus C. t. mearnsi TL (!!): 2495 (2330–2760) mm, n = 40 TL (""): 2346 (2270–2415) mm, n = 11 T (!!): 623 (565–730) mm, n = 40 T (""): 587 (550–635) mm, n = 11 HF c.u. (!!): 501 (470–540) mm, n = 40 HF c.u. (""): 489 (462–510) mm, n = 11 E (!!): 202 (183–230) mm, n = 40 E (""): 195 (187–207) mm, n = 11 Sh. ht (!!): 1226 (1110–1340) mm, n = 40 Sh. ht (""): 1171 (1070–1230) mm, n = 11 WT (!!): 201.1 (171.0–242.0) kg, n = 40 WT (""): 163.0 (140.8–242.0) kg, n = 11 Serengeti N. P., Tanzania (Sachs 1967) WT (!!): 210.0 kg, n = 40 WT (""): 165.0 kg, n = 43 Western Masailand (Talbot & Talbot 1963) C. t. cooksoni WT (!!): 235, 241 kg, n = 2 WT (""): 219, 224 kg, n = 2 E Zambia (Wilson 1968) C. t. albojubatus WT (!!): 243.0 (222.0–271.0) kg, n = 10 WT (""): 192.0 (179.0–208.0) kg, n = 10 S Kenya (Ledger 1964) C. t. taurinus Sh. ht (!!): 1472 (1410–1565) mm, n = 17 Sh. ht (""): 1353 (1290–1410) mm, n = 17 WT (!!): 237.2 kg, n = 98 WT (""): 190.4 kg, n = 95 KwaZulu–Natal (Sh. ht: Attwell 1977; WT: Hitchins 1968) WT (!!): 251.7 kg, n = 97 WT (""): 214.8 kg, n = 106 Kruger N.P. (Braack 1973) Maximum recorded horn length in the species is 86.0 cm for a pair of horns from Messina, Limpopo Province, South Africa (Rowland Ward) Key References Attwell 1977; East 1999; Estes 1966, 1969, 1976, 1991a; Estes & East 2009; Hopcraft 2010; Mduma et al. 1999; Sinclair 1979; Sinclair et al. 2000; Talbot & Talbot 1963; Watson 1967, 1969. Richard D. Estes 543

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Tribe HIPPOTRAGINI Horse-like Antelopes Hippotragini Sundevall, 1845. Öfversigt. Kongl.-Vetensk. Akad. Förhand. for 1845, parts 2 and 3, p. 31.

Beisa Oryx Oryx beisa myology.

Beisa Oryx Oryx beisa skeleton.

Hippotragini are large, barrel-bodied antelopes with long, slender and well-annulated horns, long ears and broad, heavy hooves (with well-developed pedal glands on all four feet).The tribe is represented by six extant African species in three genera, Hippotragus, Oryx and Addax, and a seventh species (the Arabian Oryx Oryx leucroyx) in Arabia. Four of these are desert or near-desert species, and the morphology of both fossil and living forms confirms the common ancestry of all hippotragines and the likelihood of an early accommodation to heat, desiccation and a marked trend towards grass-eating (Gentry 1978). False hooves are particularly well developed in the oryxes and the Addax Addax nasomaculatus. The coat is sleek, in various shades of tan, white and black and the face is striped rather like a goat or gazelle. The thick, tapering necks and mane earned these antelopes their tribal name Hippotragini, which means ‘horse-goat’. Sexes are similar, but males are heavier and have thicker horns. Preorbital glands are absent, although they are vestigial in some species (for example, in Hippotragus and Scimitar-horned Oryx Oryx dammah); inguinal glands are absent. Pedal glands are present on all four feet. Females of all species have two pairs of inguinal nipples. The earliest hippotragine fossils are difficult to separate from the earliest caprines and the earliest alcelaphines (Gentry 1978), which is consistent with the three tribes emerging from a common stock during the early Miocene (Hassanin et al. 2012). Fossils from India and Europe indicate that hippotragines were once more widespread and might even have penetrated Africa from a Eurasian source (Simpson 1953, Gentry 1978, both authors who allied this tribe with the preeminently Eurasian Bovinae, whereas

Kingdon [1982] associated hippotragines with caprines). Molecular studies have confirmed a relationship with Caprini, but suggest an even closer affinity with the Alcelaphini (e.g. Gatesy et al. 1997, Matthee & Davis 2001, Hassanin & Douzery 2003, Hassanin et al. 2012). As for geographic and ecological origins, incipient deserts in mid-Miocene North Africa or even outside Africa are possible centres of endemism or evolutionary origin. In as much as caprines almost certainly originated in Asia, there could also be a continental dimension for Hippotragini, fossils of which were first found in Eurasia and might signify a non-African emergence, possibly as the larger-bodied, lowland branch of a proto-Caprine lineage (Kingdon 1982). An early accommodation by ancestral hippotragines to cooler, drier climates (and more grass in their diets) might have begun during a marked trend towards global cooling and increasing aridity in Africa at about 14 mya (Cerling et al. 1997a, Retallack 2001, Zachos et al. 2001). Because deserts make considerable demands on the physiology of any animal it is certain that such adaptations were acquired incrementally by the ancestors of Hippotragini. Lineages that have a head-start in any set of favourable adaptive traits tend to retain their advantages in suitable habitats. In the case of other Antilopinae lineages, adaptations to drought were first made by small-bodied animals at around the Oligocene/Miocene boundary, but there is no certainty that the immediate ancestors of Hippotragini were as well suited. Indeed, Kingston & Harrison (2007) have shown that Pliocene Hippotragini were still less than wholly committed to a grass (C4) diet than the contemporary species. None the less, it seems likely

544

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Family BOVIDAE 10

5

0 mya

Addax

Oryx beisa

O. gazella

O. leucoryx

O. dammah

Hippotragus niger

H. equinus

H. leucophaeus

Tentative phylogenetic tree for Hippotragini (modified after Hassanin & Douzery 2003 and Hassanin et al. 2012).

that the hippotragine trend towards large-bodied grazers coping with desiccation might have begun in the mid-Miocene, either in North Africa or neighbouring areas of western Asia. This period well preceded the divergence between Hippotragus and Oryx, which has long been known to have been early (Kingdon 1982), but molecular studies have now narrowed this divergence down to the mid-Miocene (approximately 10.5 mya) followed by a surprisingly early divergence (8.5 mya) between Hippotragus niger and Hippotragus equinus (Hernández Fernández & Vrba 2005). By 10.5 mya, hippotragines must have been well established in Africa and a predominantly northern African focus for proto-Oryx/Addax and a southern focus for proto-Hippotragus would seem likely. This was a particularly benign period (also marked by a major exchange of fauna with Eurasia), but a dry south-west African focus would have already existed. It is therefore very interesting that a recently extinct hippotragine, the Bluebuck Hippotragus leucophaeus, shared characteristics of both the Roan and Sable Antelopes, but was smaller than either of them. This relic species survived in the Cape of Good Hope, a dry but cool region, until 1779. Its affinities were closer to H. equinus than to H. niger but it seems to have retained more generalized features than either. It may also be significant that Sable Antelope populations have been found to have the greatest intra-specific genetic diversity known for any mammal (more than 18% of divergent maternal ancestry, Pitra et al. 2002), with almost imperceptible differences in general morphology. Perhaps this level of genetic divergence has been influenced by a lineage history of more than 8 million years since the H. niger lineage diverged from the morphologically very similar equinus/leucophaeus lineage. It is significant for our broader understanding of hippotragine evolution that the 8+ mya split between H. niger and H. equinus (if the molecular clock calibration is correct) took place a mere 2 million years after the probable north/south split between proto-Hippotragus

Facial myology of Sable Antelope Hippotragus niger. Upper-right toothrows of Scimitar-horned Oryx Oryx dammah (above) and Roan Antelope Hippotragus equinus (below). TOP:

ABOVE:

and the proto-Oryx/Addax lineage. This is consistent with benign conditions having continued over much of tropical and southern Africa during the late Miocene, but this ended with the closing of the Straits of Gibraltar and the beginning of the Messinian (6.5–5.3 mya), when the Sahara Desert first spread over the whole of northern Africa. Before the Messinian was over, at least two hippotragine lineages of true desert antelopes had evolved, but the ancestors of Addax were clearly the earliest and best equipped large herbivore to have adapted to the very driest conditions of the Sahara. This left Oryx to find a slightly less extreme niche across a wider range of deserts and desert fringes. Oryx species differentiated during other periods of intense aridification in the late Pliocene and Pleistocene. Ancestral Gemsboks took advantage of one of these arid periods in the later Pleistocene to colonize south-west Africa by means of an arid corridor that connected Somalia with the Namib and Kalahari deserts. Today, Hippotragini are broad-spectrum grazers with broad, heavily crenellated teeth, well adapted to cope with tough foods (see Figure above). Their selection of food plants also shows few signs of being particularly specialized. All hippotragines, but most subtly, Hippotragus species, have specialized in exploiting zones with an impoverished fauna and flora. Their narrowed choice, even within 545

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0

5

10 km

Area of high herbivore density Lions Roan Antelope

Map of Kidepo Valley N. P. showing area of high herbivore density with records of Lions Panthera leo and Roan Antelopes Hippotragus equinus over 23 months (Kingdon 1982 from data collected by I. Ross).

the tropics, is marked by an attachment to particular localities in which they build up intimate knowledge of large home-ranges (which includes systematic avoidance of areas with numerous predators and competitors). Rigours of the environment are often sufficient to explain why Oryx occupy areas with few other herbivores, but this in insufficient to explain why Roan and Sable Antelopes also shun apparently suitable areas in spite of rainfall being relatively high and plant productivity also being good, even if seasonal. Powerful, longhorned hippotragines are certainly not excluded from pastures by aggression from competing species, so their apparently voluntary spatial restriction must be guided by environmental clues that are not readily apparent to a human observer. It has been demonstrated, in several localities, that Roan, Sable and Beisa Oryx Oryx beisa avoid areas with high densities of other herbivores (Kingdon 1982). Why should hippotragines be limited to small localities or discontinuous vegetation belts in which the density of other herbivores is markedly lower than in neighbouring areas? Early studies of predation revealed that Roan and Sable Antelopes were disproportionately susceptible to predation by Lions Panthera leo, even when their numbers were already low (Pienaar 1969a, Harrington et al. 1999, McLoughlin & Owen-Smith 2003). Likewise, Beisa Oryx time their visits to water

Dry season visits to artificial waterholes in Tsavo N. P. showing temporal separations between Beisa Oryx Oryx beisa and carnivores. Arrows indicate peaks of visits (from Ayeni 1975).

to coincide with least likelihood of predator activity (Ayeni 1975). In Kidepo Valley N. P., Uganda, Roan Antelopes very seldom intruded into areas with a high density of grazers and, concomitantly, such areas supported numerous Lions. Thus, over a two-year period, Roan consistently preferred drier, better-drained ground where both predators and potential competitors were fewer (see Kingdon 1982). This suggests that predation can be the decisive force in shaping habitat-choice for vulnerable species and that food resources, as well as direct competition for those resources, may be less influential in determining local dispersal over the landscape. The role of predation in selecting for detailed and species-specific patterns of habitatchoice may be operating at both the direct, ontogenetic level as well as at a genetic level. In the first instance, direct attrition as a result of sustained predation could bring about such patterns. In the second instance, survivors could have been selected for their ability to choose habitat in response to a wide variety of environmental clues that, over many generations, effectively reduced their chances, or their offspring’s chances, of being taken by a predator. Behaviour, modes of communication, social structures, ecological dispersal patterns and anatomy have probably all been influenced by an ancestral exposure to dry impoverished habitats. Young animals, once emerged from their ‘hider’ phase, are exceptionally social and active.They spend much of their time playing or rushing around with stylized gaits, horning objects or mock-fighting. Hippotragines are unusual in that the !! have horns that are as long as those of the "". Female social units tend to have closed membership and horns provide them with the means of excluding outsiders from scarce resources and resisting any attempts by "" to limit their movement or threaten their offspring. Hippotragines pose special challenges for conservation, not least because of their extensive ranges. As agricultural and ‘development’ interests continue to excise, diminish, constrict or obliterate conservation areas, many local populations will be extirpated, as has already happened in many areas. Jonathan Kingdon

Sable Antelope Hippotragus niger kirkii adult male.

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GENUS Hippotragus Roan and Sable Antelopes Hippotragus Sundevall, 1845. Öfversigt. Kongl.-Vetensk. Akad. Förhand. for 1845, parts 2 and 3, p. 31.

Polytypic genus with two extant species: the Roan Antelope Hippotragus equinus, in both savanna woodlands and grasslands of sub-Saharan Africa, and the Sable Antelope Hippotragus niger, confined to the southern savanna regions of Africa and, more particularly, to the miombo (Brachystegia dominated) woodlands. Judging from their relatively abundant representation in Pliocene fossil beds, Hippotragus, or its close allies, were more common then than they are today and, supposedly, more ecologically diverse. For example, a very large type of Roan Antelope, H. gigas, is known from Sterkfontein and other late Pliocene deposits and a closely related genus, Praedamalis daturi, is known from upper Pliocene beds at Laetoli (Kingston & Harrison 2007).These authors have attributed the apparent decline of Hippotragus and hippotragines to competition from more recently evolved grazers, notably alcelaphines, but the susceptibility of Hippotragus to predation can only have accelerated the decline of this genus and could be the single most significant factor defining the ecological niche of Hippotragus within the African antelope spectrum (Kingdon 1982). Roan and Sable Antelopes are subject to less predation in the miombo woodlands they occupy, while also minimizing competition with the diverse array of other large grazers that live in surrounding savannas (see discussion in Sable profile under Adaptations). Likewise, other members of the tribe escape competition by occupying the most arid habitats. When the carbon profiles of teeth from eight hippotragine specimens from Pliocene beds at Laetoli were compared with those of contemporary Hippotragus (and Oryx), the living species were confirmed to be unambiguously grass-eaters, with predominantly C4 diets, whereas the Pliocene species were all much less exclusively C4 dominated, indicating that they were mixed feeders, relying more on C3 browse than contemporary species (Kingston & Harrison 2007). That carbon profiles may reflect the gross diet but fail to show up some subtleties of actual foraging patterns should, however, be borne in mind. For example, in his studies of Sable Antelopes in several localities, R. D. Estes (pers. obs.) has found that during the dry season animals eat quite a lot of foliage and forbs. Even so, the findings of Kingston & Harrison (2007) clearly reveal differences that could be interpreted in more than one way: (1) the fact that the fossil samples come from Pliocene populations could be indicative of a wider spectrum of dietary types within Hippotragus; (2) a preponderance of C3 browsers in the Pliocene samples could be taken as evidence for less grass-dominated habitats; and (3) the fact that several tribes of the Antilopinae, notably Hippotragini, Alcelaphini and Reduncini, have all become progressively more and more specialized for grass-eating strongly suggests a phylogenetic dimension. It seems likely that the three explanations may all be correct and are probably not easily disentangled. Although Roan and Sable Antelopes are the sole survivors of this genus, the now extinct Bluebuck Hippotragus leucophaeus was a relict species, which lived along the south-western coast of the Cape Province from about Caledon to Plettenberg Bay (Ansell 1972, Meester et al. 1986). Considered by Haltenorth (1963) to be a dwarfed subspecies of Roan Antelope, molecular studies have placed the divergence

Roan Antelope Hippotragus equinus skeleton.

between H. leucophaeus and H. equinus in the region of 3 mya (Hernández Fernández & Vrba 2005). This molecular clock dating suggests that gigantism in the H. equinus lineage arose after this divergence. The delicate build, smaller size and peculiar colouring of H. leucophaeus could represent morphological responses to the impoverished or chemically peculiar soils of the Cape or they could signify conservative retentions in a formerly more widespread precursor within the ancestral lineage of H. equinus.The last individual Bluebuck was shot around 1800, the first African antelope to be hunted to extinction by European settlers (Klein 1974). Detailed reviews of the species are provided by Mohr (1967) and Klein (1974), and see Smithers (1983) for a synopsis. Of special interest are recent revelations of hybridizations between Roan and Sable Antelopes in Kruger N. P., South Africa (Estes & Whyte 2006) and the discovery that several members of the Giant Sable herd in Cangandala N. P., Angola, are clearly hybrids sired by a Roan Antelope bull (Vaz Pinto 2006). A major question, and one that is of crucial importance for the conservation of Hippotragus species, is whether Roan–Sable hybrids are viable. While Robinson & Harley (1995) have described such hybrids as viable it is the genetic details that matter. The lineages of H. niger and H. equinus are estimated to have remained separate in spite of having maintained partially sympatric ranges for at least a portion of their estimated 8.5 million year existence as separate lineages (as determined by molecular clock, Hernández Fernández & Vrba 2005). This reinforces the urgent need to study the genetics of currently known hybrids to determine whether any danger exists that they might contaminate the H. niger variani genotype, a potential catastrophe for this very distinctive population that is already on the brink of extinction. Richard D. Estes & Jonathan Kingdon 547

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Hippotragus equinus ROAN ANTELOPE Fr. Hippotrague rouan; Ger. Pferdeantilope Hippotragus equinus (É. Geoffroy Saint-Hilaire, 1803). Cat. Mamm. Mus. Nation. Hist. Nat. p. 259. ‘Inconnue’; now thought to be South Africa, Western Cape Prov., Plettenberg Bay (Grubb 1999). For dating the name to É. Geoffroy Saint-Hilaire, 1803 see Grubb (2001b, 2004, 2005).

Lateral and palatal views of skull of Roan Antelope Hippotragus equinus.

Roan Antelope Hippotragus equinus.

Taxonomy Ansell (1972) distinguished six subspecies, based primarily on morphology. Matthee & Robinson (1999b) analysed the mitochondrial DNA population structure of 13 samples representing four of Ansell’s subspecies (H. e. equinus, H. e. cottoni, H. e. langheli and H. e. koba) and confirmed the existence of at least four, well-delineated maternal clades (of course, mtDNA relates only to maternal lineages). Subsequently, Robinson & Alpers (2001) investigated both mtDNA and nuclear DNA patterns (via microsatellites), and greatly expanded the sample database from the earlier work. Their results distinguished several geographically defined genetic groupings. First, there is a clear genetic separation, both mitochondrial and nuclear, of the western African assemblage (based on samples from Benin, Ghana and Senegal). This group is referable to Ansell’s H. e. koba. Second, three other assemblages are recognized: the north-eastern assemblage with N Tanzania, Kenya and Uganda; the south-eastern assemblage with Zimbabwe, Zambia, Malawi and S Tanzania; and the south-western assemblage with Namibia, Botswana and South Africa (the ‘Kruger National Park lineages’ are not included in this assemblage because of probable previous genetic mixing with animals from elsewhere). Third, the authors reported that Ansell’s H. e. charicus (from Nigeria, Cameroon, Chad, Central African Republic) should not be considered part of an eastern and southern clade, nor of the western clade. It seems that there is still no definitive support for a central clade, which H. e. charicus and H. e. bakeri (from Sudan)

might belong to. With the exception of H. e. koba representing the western African assemblage, there is no exact correspondence between Ansell‘s subspecies and results based on DNA. Other Roan populations form a geographically diverse assemblage with no clear genetic correspondence between subspecies. Alpers et al. (2004) distinguish two evolutionarily significant units (per Ryder 1986) for the Roan: the western African group, and a ‘rest-of-Africa’ group (including cottoni, equinus and langhelidi subspecies), but noted that more work is needed to clarify the status of charicus and bakeri subspecies. Synonyms: aethiopica, aurita, bakeri, barbata, charicus, cottoni, docoi, dogetti, gambianus, jubata, koba, langheldi, rufopallidus, scharicus, truteri, typicus. Chromosome number: 2n = 60 (Robinson & Harley 1995). Despite very different phenotypes, and subtle differences in chromosomes (Fordyce-Boyer et al. 1995), the Sable Hippotragus niger and Roan Antelope are genetically close enough to produce viable hybrids (Robinson & Harley 1995; see Estes & Whyte 2006, Vaz Pinto 2006, and Hippotragus niger species profile for further discussion). Description Large-sized antelope, the second tallest after the Common Eland Tragelaphus oryx, and third heaviest after the Common Eland and the Bongo Tragelaphus eurycerus. Horse-like body-shape with short, stiffly erect, greyish-brown mane edged with black extending from neck to withers. Pelage uniform sandy-fawn, or light reddish-fawn to grey, or dark rufous. Conspicuous facial mask with striking colouration comprising white parts on cheeks, muzzle, chin and on two oblique and parallel stripes along each eye, constrasted against deep dark black background. Face pattern varies both individually and regionally, with black markings more extensive in

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north, and light markings in south. Black patches of facial mask become greyish with age, even turning whitish in very old individuals. Ears long and narrow and terminate in tufts 3–5 cm long. Shoulders higher than hindquarters. Legs long and hooves large; false hooves prominent. Tail reaches heels, with black tassel on lower half. Preorbital glands are vestigial and there are pedal glands on all feet; inguinal glands are absent. Both sexes have paired ringed horns that rise steeply and curve evenly backwards with diverging tips. Horns strongly ridged for most of the length, but tips are smooth. Sexual dimorphism is weak, !! being only slightly heavier than "", while horns, head and neck are slightly thicker in !! than "". Geographic Variation As noted above, six subspecies have been identified by Ansell (1972); however, the validity of some of these subspecies is still in doubt, and recent genetic studies have shown that only the western African subspecies (koba) constitutes a genetically separate group from those inhabiting the rest of Africa (Robinson & Alpers 2001, Alpers et al. 2004). Western populations: H. e. koba: Senegal to Benin. General colour pale tawny, although specimens from West Africa tend to be reddish; forehead chestnut in both sexes. Central populations: H. e. charicus: E Nigeria, Cameroon, Chad and Central African Republic. No discernible differences with H e. koba. H. e. bakeri: Sudan. Browner than the other races; forehead blackish in !!, reddish-brown in ""; chest pale, as are ears and front of pasterns. Eastern and southern populations: H. e. langheldi: S Sudan, Ethiopia, N DR Congo, Uganda, Kenya, Tanzania, Rwanda, Burundi. General colour pale reddish; forehead reddish-brown in both sexes. H. e. cottoni: S DR Congo, Angola, Zambia, C and N Malawi, N Botswana. No discernible differences with H. e. equinus (see below), except they are redder than other specimens. H. e. equinus: Zimbabwe, South Africa and Mozambique. General colour greyish, with forehead black in both sexes. Similar Species Hippotragus niger. Sympatric through most of the southern savannas. These two grazing species tend to remain in different vegetation types, the Sable being a species less of grassland/tree-savanna and more of open miombo woodland. The Sable is smaller on average, characterized by sexually dimorphic colouration – deep black pelage in adult !! (sorrel to chestnut in "" and young); horns are longer, commonly exceeding 1 m. Distribution Endemic to Africa. Formerly one of the most widely ranging of African antelopes, found throughout savanna woodlands and grasslands of sub-Saharan Africa, but now greatly reduced in range (East 1999), though remarkably still locally abundant in parts of West Africa (Poché 1974). Roan were probably present in Egypt until 2000 BC (Osborn & Osbornová 1998).

Hippotragus equinus

Historical Distribution In West Africa, Roan Antelopes ranged from C and S Senegal, Gambia, S and E Guinea-Bissau and S Mauritania east through SW Mali, N Guinea, N Côte d’Ivoire, Burkina Faso, Ghana, Togo, N and C Benin, SW Niger, and C and N Nigeria. From there, they ranged eastwards through N Cameroon (from L. Chad to the Adamaoua Plateau), S Chad, Central African Republic and N DR Congo and S Sudan to the western lowlands of Ethiopia in the Sudanian region between latitudes 15 and 19° N; they ranged as far north as the south-western savannas of Eritrea. Southwards they ranged through NE and SW Uganda, S Kenya, Tanzania, NE Rwanda and Burundi to Angola (except in Cabinda and the arid southwest), S DR Congo, NE Namibia, Botswana (where older records evidence that they formerly occurred much further south than they do today; Smithers 1971) and Mozambique (East 1999). In South Africa, an observation dating to 1778 suggests that a population of Roan Antelope formerly inhabited the Western Cape in the vicinity of Plettenberg Bay (consistent with the fossil record), but were likely extirpated from the region during the late 1700s (Faith 2012). In recent times, their range has extended only as far as the bushveld/ lowveld regions of the North West, Limpopo and Mpumalanga Provinces, and the Northern Cape. Current Distribution The range of the Roan Antelope has contracted greatly since the beginning of the twentieth century, and they have been eliminated from parts of their former range surviving mainly in and around protected areas as well as in non-protected areas with low densities of people and livestock (East 1999). A striking feature in the distribution of the Roan Antelope is a clear contrast between West and central Africa, where it remains one of the most common antelopes, and East and southern Africa where it is one of the rarest. In West and central Africa, its range has contracted in the face of hunting and habitat degradation although the species remains locally common. On one hand, Roan Antelopes have disappeared 549

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from areas lying on the fringe of the typical habitat such as S Mauritania, Gambia (where they now occur only as vagrants in the east), N Sierra Leone, N Liberia, C Mali and C Niger; on the other hand, they remain locally abundant in the heart of typical habitat such as SE Senegal, SE Burkina Faso, N Benin, SW Niger, N Cameroon, SE Chad and N and E Central African Republic (East 1999). Dowsett & Dowsett-Lemaire (1989) give Roan Antelopes as occurring in the savanna of Congo, based on records from two localities – Les Grandas and Lefini; however, the species is certainly no longer recorded there. In East and north-east Africa, the Roan Antelope is extinct in Eritrea and in Burundi, but still survives in W Ethiopia (e.g. in Gambella N. P.) and in SW and SE Sudan (J. O. Heckel pers. comm.). In Kenya, Roan Antelopes are present only in Ruma N. P., where they are entirely surrounded by cultivation (East 1999). In Uganda, East (1999) reported them surviving only in Pian-Upe G.R. They are still widely present, albeit patchily, in Tanzania (notably Biharamulo, Moyowosi, Kigosi, Katavi, Rukwa, Rungwa). In Rwanda, they are restricted to Akagera N. P., although they have been in perpetual decline since the park’s boundaries contracted (Williams & Ntayombya 1999, Apio & Wronski 2011). In southern and south-central Africa, Roan Antelopes are still widely distributed in Zambia, where they survive mainly within protected areas and game management areas, including the Luangwa Valley and Kafue N. P. In Angola, there is limited information, but they are reported to survive in some areas (P.Vaz Pinto pers. comm.). In Malawi, they are now confined mainly to four protected areas in the north and south, with the largest population in the small Nyika N. P. (East 1999). In Mozambique, there are two locations with very limited populations of Roan Antelopes: in the hunting areas of western Tete Province and in Great Limpopo Transfrontier Park (East 1999); there are also plans to reintroduce them to Gorongosa N. P. In Namibia, the present ‘natural range’ of the Roan Antelope includes the East and West Caprivi Strip, Khaudom G. R. and Nyae Nyae Communal Conservancy. They were introduced to Etosha N. P. in 1970 and to Waterberg Plateau Park in 1975. However, the Kaross area in the west of Etosha, where the main Roan population is located, falls below the 300 mm rainfall isohyet and the population has not thrived (Martin 1983). Roan Antelopes are today confined to the northern savannas of Botswana, while in Zimbabwe they have disappeared from the Highveld but are still present in limited numbers in the Hwange ecosystem, in the mid-Zambezi valley and in patches of the lowveld (East 1999). The Roan Antelope is among the most threatened antelopes in South Africa. The species still survives in the north of Kruger N. P. (but see later), and has been reintroduced elsewhere, including Marakele N. P. in Limpopo Province. A breeding population of Roan is maintained in Percy Fyfe N.R. in Limpopo Province. Although there is no evidence that Roan Antelopes formerly occurred in KwaZulu–Natal, they have been introduced to reserves such as Weenen and Karfloof Nature Reserves (Rowe-Rowe 1994). The indigenous population that formerly occurred in NE Swaziland is extinct, but Roan Antelopes have since been introduced to Mkhaya G. R. (Monadjem 1998). Habitat The Roan Antelope is usually present in dystrophic (nutrient-poor) savannas, such as miombo woodland, and inhabits

gently undulating terrain of open woodlands and savannas with long grass, low tree density and canopy cover (Pienaar 1963, Joubert 1976, Wilson & Hirst 1977, Dörgeloh 1998a, b, Perrin & Taolo 1999b). Cover of high grasses and woody plants plays an important role for both grazing and calving (Taolo 1995, Dörgeloh 1998a). Calving "" select habitats with long grass and woody plants, which provide shelter for the newly born calves (Allsopp 1979, Dörgeloh 1998b). The highest densities of Roan Antelopes are found in areas with an average rainfall of around 1000 mm, where soils are mainly infertile and support grazing of low quality. They tend to avoid areas of short grass and woodland where the trees form a closed canopy or where the bush forms thick, closed stands (Heitkönig & OwenSmith 1998). Permanent water is important. Shrub encroachment seems to have a negative effect on Roan Antelopes (Joubert 1976, Wilson & Hirst 1977), even though they tolerate low shrub growth (Ben-Shahar 1990). In Comoé N. P., Côte d’Ivoire, Roan Antelopes are notably present in woodlands during the wet season. They do not cover great distances, but range over different types of altitude and vegetation depending on the season. At the beginning of the wet season they leave the plains bordering the rivers and migrate towards higher areas, where they are frequently observed in the open woodlands (Steinhauer-Burkart 1987). In Bouba Ndjida N. P., N Cameroon, situated in the savanna woodland belt and receiving an average annual precipitation of 1200 mm (falling mainly from May to Oct), Roan Antelopes occur in Terminalia laxiflora wooded savanna. The trees of this habitat do not exceed 3 m in height and grass cover consists mainly of tall perennial Andropogoneae. These grasses grow up to 3 m high during the wet season and constitutes the principal forage for the majority of large herbivores. Roan occur to a lesser extent in the Isoberlinea doka savanna woodland, but seem to avoid fringing forest (Van Lavieren & Esser 1979). In Bénoué N. P., Cameroon, Roan Antelopes prefer Burkea–Detarium open savanna rather than Isoberlinia woodland and Anogeissus riparian forest during the dry season. During the wet season, Burkea–Detarium open savanna is still the preferred habitat, but they also occur in T. laxiflora and T. macroptera open savannas (Stark 1986b). In Zakouma N. P., Chad, Roan often occur in the Combretacea wooded savannas (52% of observations) or in the Acacia seyal savannas. They are less frequently observed in more open areas (Dejace et al. 2000). In Kenya’s Lambwe valley, Allsopp (1979) observed that Roan selected open Themeda/ Setaria grassland, and never observed them in Hyparrhenia grassland or in dense continuous thickets. In the Waterberg region in Limpopo Province, South Africa, part of the moist savanna biome dominated by woody plant species such as Terminalia sericea, Burkea africana and Combretum spp. and characterized by abundant and perennial water-courses and rivers, Roan Antelopes occupy an area of flats with sandy acidic soils and low rock cover. They also venture onto rocky slopes further away from water resources (Ben-Shahar 1990). In Kruger N. P., favoured habitat is on slightly undulating land consisting of heavy clay soils derived from basalt; hillier areas on clay–loam soils derived from dolerite are also used to a lesser extent (Joubert 1976). Abundance East (1999) estimated the total population of Roan Antelopes at about 40,000, although correcting for undercounting bias in aerial surveys gives a slightly higher estimate of 76,000 animals,

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of which 60% occur in and around protected areas. According to East (1999), the largest populations survive in Burkina Faso (>7370), Cameroon (>6070), Zambia (>5080) and Tanzania (>4310).Winter (1997) noted that more than 50,000 Roan Antelopes survived in S Sudan in the 1970s, including 4100 in the Jonglei region and 2000 in the Boma region. Spinney (1996) commented that Roan Antelopes in Sudan survived locally in relatively stable numbers. During recent surveys conducted during the dry season of S Sudan, only 21 individuals were counted in Southern N. P., with a small group sighted in the Jonglei and two other sightings in Boma N. P. (Fay et al. 2007). Historical accounts indicate that the Roan Antelope was extremely rare in certain localities but could reach considerable densities in areas where environmental factors were favourable, for example in parts of West Africa (Poché 1974). Densities vary greatly across their distribution, but the average density is quite low (Wilson & Hirst 1977). In natural conditions, the density does not exceed 4/ km2, even in areas where nutrients are not deficient (Wilson & Hirst 1977). But under intensive management, Roan Antelopes may be stocked at high densities of up to 20/km2 (Dörgeloh et al. 1996). For example, in Nylsvley N. R. (South Africa) density could reach about 8/km2 (Dörgeloh 1998b). Effective management could explain why, in the 18 years preceding 1992, the Roan population in South Africa grew at approximately 1% per year. The result of the national inventory in 2001 was 1237 animals. Aerial surveys provide estimated population densities of 0.007/ km² in Kruger N. P. (Kröger & Rogers 2005), 0.2–0.6/km² in Pendjari N. P. (Benin) (Chardonnet 1995, Rouamba & Hien 2002), and 0.8–1.5/km² in Pama–Arly–Singou (Burkina Faso) (Lungren et al. 2004). Aerial surveys of Arly N. P., where the species is common, have produced densities of 1.1/km² (Barry & Chardonnet 1998) and 0.34/km² (Bouché et al. 2004). Aerial surveys of Bouba Ndjida N. P. gave a density between 0.2 and 0.6 herds/km² with a mean herd size between 4.2 and 6.9, producing a total population size estimated at 4114 (Van Lavieren & Esser 1979). Ground surveys provide densities ranging from 0.03 to 0.10/km² in areas like Manda N. P. (Chad) (Chai 1996), Comoé N. P. (Fischer 1996, Fischer & Linsenmair 2001a) and Dinder N. P. (Sudan) (Hashim et al. 1998). In Waza N. P., where Roan densities have been monitored over several decades, they declined from 2.4/km² (1960s, following 20 years of above-average rainfall and effective protection) to 0.7/km² (mid-1990s, following droughts and transmission of disease by livestock) to 0.2/km² (early 2000s, following increased poaching) (Scholte et al. 2007). In Konkoumbouri Hunting Area (Burkina Faso), density was 3.9/km² (Bouché & Lungren 2004), which is considerably higher than the density of 0.27/km² recorded in the same area based on aerial surveys (Bouché et al. 2004). Adaptations Roan and Sable Antelopes share common aridadapted ancestors with other hippotragines. These ancestors came from North African or western Asian biomes (Kingdon 1982) and probably came to occupy both northern and southern savannas following a dry period in the mid-Miocene (Hernández Fernández & Vrba 2005). Since semi-arid savannas were already occupied by several ungulate species, it is possible that Hippotragus had to adapt themselves to broad-leafed deciduous woodlands and grasslands of the mesic savanna that were freer from large herbivore competitors.

Changes contingent on latitudinal shifts probably had consequences for their social organization: territoriality in !!, traditional homeranges for herds of "", spatial segregation of bachelor !!. Habitat changes were accompanied by morphological changes such as lengthening of the head and an increase in gape and the incisor– molar diastema, changes presumably well adapted for feeding on tufted perennial grasses. It has been suggested that the Roan Antelope’s occupation of large home-ranges, dependence on water and relatively slow gait might all be symptomatic of derivation from a still larger, giant-sized ancestral stock (Kingdon 1997). Roan kept the typical hippotragine ability to walk long distances daily, which allows them to reach remote feeding grounds beyond the reach of most competitors; furthermore, being less water-dependent than many potential competitors, such as Kobs Kobus kob, they have access to good quality forage further away from water and away from other herbivores (with the exception, of course, of the Savanna Elephant Loxodonta africana). Foraging and Food Roan Antelopes are considered to be predominantly grazers, taking only a small amount of browse. In their review of the dietary preferences of African bovids, Gagnon & Chew (2000) classified Roan as variable grazers, while studies involving stable carbon isotope analysis (from southern Africa only) are consistent with observations (e.g. Joubert 1976, Heitkönig 1993) that suggest that grass predominates in their diet (Sponheimer et al. 2003b). However, although Joubert (1976) found that browse did not contribute significantly to the diet (less than 5% of total intake), several other authors have mentioned that browse could play an important role during critical periods of the year, especially in the late dry season (Poché 1974, Wilson & Hirst 1977). In Zimbabwe, Child & Wilson (1964) showed that Roan Antelopes used to feed from browse plants such as Diplorhynchus mossambicensis (leaves), Capparis tomentosa (leaves), Piliostigma thoningii (fruit), Diospyros kirkii (fruit), D. mespiliformis (fruit) and bamboo (flowers). Schuette et al. (1998) studied the diet of Roan Antelopes in West Africa by means of microhistological analysis of faeces and found that Roan Antelopes switched from being predominantly grazers (>95% grass) in the wet season to mixed feeders (200 for all radio-collared "" followed longer than a few months. Although large aggregations occur in seasons of green grass growth, "" that calve in green conditions usually still persist in isolation and form consortships in the immediate postpartum period. At calving, experienced adult "" tend to isolate themselves to give birth to a cryptic calf that adopts a solitary hiding defence strategy over the first 2–3 weeks of life. In the Fringe-eared Oryx this behaviour was observed consistently in radio-collared wild animals, and in the semi-captive Galana herd it was notable that adult "" showed a sharp peak in tendency to run away back to the wild in the month of calving. Most "" come into postpartum oestrus within 1–2 weeks, and so often attract the attention of adult !!. Small group sizes of 1–3 individuals (typically some combination of a lactating cow, neonate and consorting adult !) are most typical (but not invariable) in this period and calving interval data indicate that there is a high probability that healthy adult "" will conceive within this period. Females may also intermittently join herds while the calf lies out (Kingdon 1982). During the period 2–6 months postpartum, adult "" are most likely to be found in mixed-sex herds (10–15% adult and subadult !!) of variable size, with a range of calves and younger age groups also present. Crèching among calves and cohorts of young may be apparent, and on occasion a lactating cow may move as far as 10–12 km away from her own calf and back again in the course of a single day when, for example, travelling to a water source to drink in the dry season. During the period 6–9 months postpartum, adult "" become more likely to be found in mixed groups composed mainly of adults and subadults only, with a marginally higher proportion of !! (20–30%). This is not an absolute change, but it appears that progressively less time is spent in the immediate company of her own or other young in the period of later pregnancy leading up to isolation for birth of the next calf. A tendency of Fringe-eared Oryx herds to form group types characterized by the presence or absence of young was also noted independently in Tsavo East N. P. (Leuthold & Leuthold 1975b). The majority of maturing, and some fully adult, !! are also found in mixed-sex herds and information from individuals suggests that their ranging and foraging behaviour operates on a similar wide scale to that seen in "" (see above). All-male groups of Fringeeared Oryx bulls were only identified on six occasions in the course of a three-year study (setting aside the common phenomenon of solitary adult !!). Known radio-collared territorial bulls were present on five of these occasions, confirming the group was a temporary association. On the sixth occasion, the largest all-male group recorded (five individuals found walking to the river) included a tagged wild ! who was known to have been a member of a mixedsex group of 96 animals the day before and who was next seen in a mixed-sex group of 57 two weeks later.

However, some adult !! behave very differently, distinguished by spending the majority of their time alone and in comparatively restricted areas. In the Galana–Tsavo border zone, the best documented solitary male home areas (territories) were placed along the ecotone between red and grey soils, straddling grassland and woodland habitats preferred by herding oryxes. Two radiocollared solitary bulls in this area restricted their movements to separate 5–8 km2 zones over study periods of 1–3 years. They were seen to patrol boundaries in parallel with neighbouring adult !! with some evidence for a degree of spatially determined reciprocal dominance between neighbours. These !! make visual displays by a ritualized sequence of squatting defecation, associated only with adult male oryxes (castrated adult male oryxes did not develop this behaviour on Galana Ranch). This involves walking directly to a known dung site, scenting the ground and/or older dung piles, scraping the ground or dung pile with a front hoof before squatting low on the haunches while defecating, in the process creating a strikingly vertical visual display apparent at long range to other oryxes. Dung piles were not usually dropped exactly on top of one another, but scattered around a patch of bare soil. Sometimes 3–4 dung sites may be visited and re-marked in direct succession in the course of a morning ‘patrol’. Midden sites (n = 61 observations of active dung site use at 50 different dung site locations) mapped for one radio-collared bull were found to be scattered throughout his area of residence, with no indication of clustering towards the margins (Wacher 1986). More than 80% of sites had been used previously with 3–6 older dung piles (maximum ten) evident at most sites when inspected immediately after use. Tellingly, the most common social grouping of solitary !! when not alone was to be seen in groups with a single adult " (with or without new calf present), although larger mixed-sex groups move through territories fairly frequently, when the local resident may or may not join them temporarily. Consortships between solitary !! and oestrous "" are normal, often lasting 3–5 days and sometimes as long as three weeks. Radio-collared cows with neonates mostly consorted with 2–3 different !! over the postpartum period. Solitary bulls attempt to herd and control movement of consorting "" using passive body positions and posture, and active horn threats, chases and lunges. This usually results in the " remaining near the centre of the territory, but not always. Oestrous "" can and do escape to another solitary bull, or even back to herds on some occasions. Under these conditions, with dams in view on a neighbouring territory, it is notable that on three separate occasions one well-known, solitary, radio-collared bull was seen vigorously herding and threatening single calves (on one occasion galloping at full speed to head off the running calf) to prevent them moving towards their dams and retain them on his own side of the boundary. On one of these occasions a " was seen returning to her calf on the radio-tracked male’s territory. Adult !! have frequently been noted to be attentive to young calves, whether encountered in small or larger groups. In most situations this takes the form of herding and nudging the calf towards its mother (but in Arabian Oryx calves have been seen tossed through the air with a rough sweep of the adult male’s horns). Whilst these activities may confer a certain protective advantage to these calves in normal situations, it seems probable that a prime motivation of the ! is to manipulate the calf as a means of influencing movements of the mother while she is receptive to mating.

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Relations between solitary and herd-living !! visiting the territory can be surprisingly calm and tolerant in the absence of receptive "". Territorial bulls may exert dominance to newly arrived !! by means of a head-low march followed by ritualized lateral head-nod or ‘ear-point’ display across the front of the recipient (Walther 1978c, and see full description below). On other occasions they may not even bother to approach a mixed-sex visiting group in plain view on the territory. Prolonged and serious clashes between adult !! do occur, but are relatively rare. In the Galana study, this was only seen between pairs of !! unknown to the observer. A limited number of observations of a well-known resident solitary bull dealing with mixed-sex groups on his area that included a sexually receptive " suggested that the resident bull had little difficulty retaining social dominance and probably mating advantage also over the visiting herd !!. On such occasions, active herding and chasing is directed primarily at the ", but also at other members of the herd by the resident bull. Visiting !! may also attempt to court and mount the " while the resident is occupied or elsewhere. On one radio-collared male’s territory, mountings by all adult male oryxes made during herd visits to the territory were scored as ‘successful’ or ‘unsuccessful’ depending on whether the " involved stood still or walked forward as intromission was attempted. Summing over five different occasions when receptive "" were courted by both the resident and visiting !! in groups on the territory, the resident bull was scored ‘successful’ on 5/22 mountings observed while the visiting !! achieved 0/20 successful mountings and were commonly seen to be submissive when approached and threatened by the resident bull. The circumstances of these observations raise interesting questions about female options to control mating and mate choice. However, because the social system is dynamic and flexible it seems very probable that not all "" mate exclusively with solitary territorial !!. Receptive "" in large mixed-sex groups are sometimes observed, and the level of harassment and chasing by several !! on one cow can be intense on such occasions. This suggests that solitary territoriality is unlikely to be the only mating strategy used by oryx at Galana. In drier areas, used by Beisa Oryxes in N Kenya and Ethiopia, and at the margins of the oryx range, it seems even more probable that alternative, more mobile, or herd-based mating strategies might be expected to predominate. In Serengeti N. P., one herd of 15 Fringe-eared Oryxes travelled 17 km in one day, and a lone bull walked 4 km in a straight line within one hour; on the other hand, another Serengeti herd remained localized in a 20 km² area for three weeks (Walther 1978c). In these conditions it is possible that !! may attempt to detain "" groups when local grazing conditions allow, but may alternatively attach themselves to more determinedly mobile groups if food or other resources do not attract sufficient isolated calving "" to their chosen areas. It is therefore possible that oryxes are able to adjust social relations to resident or nomadic conditions and observed mating activities appear to exhibit features of both resource-defence and mate-defence options. It is also noted that in more recent studies of reintroduced Arabian Oryxes in harsh desert conditions of S Saudi Arabia, the oldest adult !! developed very large-scale spatial segregation relative to each other, where they were comparatively sedentary, not emulating younger !! who wandered widely in search of dispersing "" following rare rainfall events. These

observations hint at an unproven possibility that male oryxes may adjust their mating strategy according to circumstance, age and perhaps body size. Studying free-ranging groups of Fringe-eared Oryxes in a semidomesticated and resident state, Stanley Price (1978b) found that some of these groups had a closed membership that was of long standing. In such groups there was an alpha-male and often a betamale immediately below this dominant individual. All adult "" were arranged in a hierarchy below these !!, but they were dominant over any other adult !!. The ability of "" to hold social status in this way allows them to control their immediate grazing space effectively among conspecifics in mixed-sex herds. Besides providing a level of defence against predators (see below) this may be a further selective advantage for the existence of fully developed horns on "" in oryxes generally. Oryxes are notable as a group for specializing in habitats where resources can be scattered and rare, "" are adapted to breed at a high rate with simultaneous lactation and early pregnancy commonplace, and effective intraspecific defence of energy resources via social status may be at a premium. The vivid markings may well facilitate social interactions in that even the slightest gestures are amplified by the patterning.Thus, when an adult ! expresses routine dominance to subordinates using the ‘head nod’ or ‘ear point’ display he walks across the front of the target animal with head and horns held high, slightly turned to one side and ears forward.At the last moment the horns are tilted over in a ritualized (but undelivered) slow motion lateral ‘blow’ to the air in front of the recipient, the main gesture being created by movement of the brightly marked head, with profile of neck and shoulders clearly shown off.The ritualized submissive response also emphasizes head movement; the receiving animal stands at first with head low, horns pressed protectively down along the neck, before half jumping, half turning away with a shake of the head as the stylized ‘blow’ is delivered. Captive "" in this situation have been seen to lie down and bawl (Kingdon 1982). In addition to these ritualized exchanges, low-key social signalling is an ongoing feature at a range of intensities among oryx groups. Individuals may use a selection of simple stares, positional displacements and avoidance, horn and head gestures in incremental intensities while interacting during grazing or movement. In this context it is notable that adult "" make use of horns and gestures to maintain status and position while grazing in a similar way to !!. The occurrence of dramatic prancing ‘tournaments’ is described above. At close range, frequent soft grunts may be heard, notably from cows approaching young calves, or between courting pairs, while calves may bawl vociferously when denied suckling. In conflict, horn-to-horn clashes may develop with increased intensity among older animals. Fighting takes place face to face with antagonists alternatively ‘kneeling’ and standing to push against each other between sharply delivered head blows and horn swipes. With rising intensity fighting oryxes may swing round and lean together in a side-to-side position with horns crossed and over-the-shoulder stabbing actions aimed at the neck and shoulder region of the opponent. Fights usually end with one individual breaking away and running, pursued for a short distance by the victor who often then turns away and may indulge in a squat defecation display a short time later. But fatal outcomes do occur in other oryx species (Arabian Oryx) and may also be expected as a rare event in Beisa and Fringe-eared Oryx. 583

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Courtship consists of prolonged bouts of nose-to-tail circling, the " initially repeatedly stepping away from the male’s attempts to press his forequarters to her hindquarters, sniff and rub the female’s perineal area, kick (laufschlag) or mount. Females may also respond by producing urine, which the courting !! will sample through typical flehmen. Unreceptive "" may on occasion repel the ! with repeated frontal clashes or may also lie down to discourage attention. But !! may also initiate courtship bouts by pawing at lying "" to get them to their feet during rest periods. As circling intensifies the pair break into occasional excursive and rather slow circular or figure-of-eight marches led by the " to and from a more or less fixed point, as the bull follows directly behind. Mounting is rapid, with the male’s head held high to avoid the female’s horns and intromission is achieved with a single firm thrust. Reproduction and Population Structure Births occur throughout the year, but well-defined birth peaks may emerge in some conditions (Leuthold & Leuthold 1975b). As !! tend to mate "" during their postpartum oestrus, and the gestation period is in the region of 8.5 months (see below), the timing of birth peaks remains difficult to understand (Kingdon 1982). Birth months were not synchronized among six radio-collared breeding "" captured from the same population on Galana Ranch where newborn calves were found in all seasons and nearly all months (Wacher 1986). Stanley Price (1978b) found that cows matured at about two years. Ages at first mating recorded from five captive, but partially freerange, female Fringe-eared Oryxes at Galana Ranch ranged from 2 years 49 days to 2 years 227 days. Age at first calving from this sample ranged between 2 years and 339 days and 3 years and 194 days, with at least one of these "" failing to conceive at first oestrus. Modal gestation period indicated by counting last recorded mating to parturition was 255–259 days (8.5 months, n = 24 pregnancies) on Galana Ranch (Wacher 1986). The modal interval from birth to postpartum oestrus (indicated by mating) was 15–19 days after parturition (n = 32 births recorded from 18 individuals). The modal inter-calving interval from a sample of 68 births recorded from 19 individual female Fringe-eared Oryxes was 275–279 days (9.1 months); individual "" were recorded maintaining a ca. 9 month inter-calving interval under semi-captive natural range conditions over a period of six years on Galana Ranch (Wacher 1986). A radiotracked wild " was detected calving precisely at a 288-day interval, well within the typical range of calving intervals recorded from tame oryx. Observations of four other radio-collared wild "" also supported maintenance of a 9-month inter-calving interval (Wacher 1986). Based on semi-captive Fringe-eared Oryxes at the Galana Ranch, male calves average 10.5 kg at birth (n = 12) and females 11.5 kg (n = 7) (R. K. Ngowah & M. Stanley Price pers. comm., Galana Game Ranch Research Project). Predators, Parasites and Diseases Cheetahs Acinonyx jubatus are known to run down young animals, and hyaenas, Leopards and smaller predators are likely to take calves. Lions kill adults where both species are common, but such large predators are generally scarcer in the very arid areas that these oryxes prefer (Kingdon 1982). African Wild Dogs Lycaon pictus have also been recorded hunting Fringe-eared Oryxes in the Tsavo–Galana area of E Kenya with variable results. A radio-collared solitary adult bull

Beisa Oryx Oryx beisa in subordinate head-low posture.

known to be blind in one eye was killed and eaten by a pack of seven African Wild Dogs. A pack of five African Wild Dogs were observed approaching a herd of 21 oryxes in open grassland, including a single small calf around one month of age. They ran up to the oryxes and began walking intently forward, focusing their gaze on the calf located near the centre of the group. The adult oryxes formed a concave semi-circle facing the dogs; when the dogs were no more than 10 m from the oryx herd, the largest adult male oryx suddenly leapt forward at the lead dog making a tossing lunge with his horns. The dogs retreated at once, circled past the herd and continued on their original direction without further interest. The adult bull in this case was also radio-collared, from which it was known he was usually alone on his territory while the other 20 animals with him were newly arrived in the area and at best only intermittent associates. In a third incident, the fresh skeletonized carcass of an adult female oryx was found in front of a fallen Commiphora tree, which had clearly been used as a protective background support while attempting to defend herself with horns when turning to face attackers after a period of pursuit (a characteristic defence strategy of hippotragines). The oryx skull was notable for the fact that the incisiform dentition was worn down to an incomplete row of rounded pegs level with the gum, suggesting a very old animal probably poorly fed. The intact body of a dead African Wild Dog was photographed lying a few yards from the oryx skeleton, marked only by a deep stab wound consistent with an oryx horn entering the body cavity at the angle

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of the hindleg. A pack of 12 live and heavy-bellied African Wild Dogs was located resting in the shade 500 m away. Virus neutralizing antibodies to malignant catarrhal fever virus were found in the blood sera of 50 Fringe-eared Oryxes (Mushi & Karstad 1981). Antibodies to Brucella abortus bacterial infection have also been reported (Paling et al. 1988). Conservation IUCN Category: Near Threatened (O. b. beisa – Near Threatened; O. b. callotis – Vulnerable C1). CITES: Not listed. Although this species has declined markedly in numbers and range, it remains common in areas where livestock densities are low, with several large populations surviving in protected areas, such as Awash N. P. (in the Awash Valley), and Omo and Mago National Parks (in the south of Ethiopia), Sibiloi, Meru and Tsavo National Parks (Kenya) and Tarangire N. P. and Mkomazi G. R. (Tanzania) (East 1999). Boma N. P. is the only protected area for this species in SE Sudan (Fay et al. 2007). Where they have been hunted traditionally it has usually been for the good quality of their meat and the practical value of the very tough hide. The species remains susceptible to the effects of hunting and competition with livestock, since so many remaining populations are outside protected areas: according to East (1999), only 17% of Beisa Oryxes and about 60% of Fringe-eared Oryxes occur in protected areas. The most recent review of local population status within N Kenya compared aerial survey data and found continued substantial (>50%) reductions in density at important population centres in Samburu District over the period of the mid-1990s to 2008, though noting a more stable situation within Laikipia (Woodfine & Parker 2011). More effective protection and management of the remaining populations in areas where the species still occurs in substantial numbers, but is in decline, such as the Awash Valley, Omo-Mago-Chew Bahir, N Kenya and Tsavo, would greatly enhance the long-term survival prospects of this species (East 1999). The absence of karyotypic sterility barriers (Kumamoto et al. 1999) suggests that, in captivity, all oryx taxa may be vulnerable to hybridization, which from a conservation perspective is highly undesirable among these distinctive and naturally allopatric antelopes. Genetic evidence supporting the historical isolation between O. b. callotis and O. b. beisa led to a direct recommendation that conservation efforts should be directed towards preserving the genetic integrity of each group on the grounds that they may have distinct evolutionary potential (Masembe et al. 2006). As noted already, hybridization in captivity between this species and the Gemsbok has produced ‘viable offspring’ (Gray 1972). Opportunities for either subspecies to hybridize with the Scimitar-horned Oryx Oryx dammah and Arabian Oryx and for mixing between the two subspecies of Beisa Oryx exist in some captive collections. Comparatively small zoo-managed populations of both the Beisa and Fringe-eared Oryxes exist in zoos outside Africa, mainly in Europe and North America (EEP/SSP). Given the downward trend of wild Beisa populations in particular, and the successful role captive breeding has played for two other members of the taxon (Arabian and Scimitar-horned), the efficient long-term management and planning of captive Beisa groups may prove to be important in the future, especially if this is set up within range states where metapopulation management of divided and semi-captive populations in conjunction with National Parks may become an increasing necessity.

Beisa Oryx Oryx beisa adult male in confrontational posture showing linear geometry of patterning.

Measurements Oryx beisa O. b. callotis HB (!!): 1800 mm, n = 1 HB (""): 1700 mm, n = 1 Sh. ht (!!): 1100 (1050–1150) mm, n = 8* Sh. ht (""): 1110 (1050–1150) mm, n = 20* HF c.u. (!!): 380 mm, n = 1 HF c.u. (""): 410 mm, n = 1 Galana Ranch, Kenya (T. Wacher pers obs., Galana Game Ranch Research Project) *Semi-captives (all animals >36 months of age) The average weight at maturity (ca. 3–4 yr of age) of semi-captive Fringe-eared Oryxes has been estimated at 165.0 kg for adult !! (n = 19) and 146.0 kg for "" (n = 34) (Carles et al. 1981). The 585

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maximum weight recorded for a " on Galana Ranch is 180.0 kg; the heaviest semi-captive ! weighed in excess of 210 kg (T. Wacher pers. obs.). Ledger (1968) recorded average weights of wild O. beisa as 176.4 kg (range 167.8–209.4, n = 10) for !! and 161.5 kg (range 116.0–188.4, n = 10) for "". Available data from semicaptive Fringe-eared Oryxes suggest that full male body size and neck development are attained after 4–5 years of age, well after the age of physiological sexual maturity. This poses further interesting questions for potential differences in reproductive strategies operating in

individuals of differing body size in this complex and fascinating species Maximum recorded horn length for O. b. beisa is 109.2 cm from Ethiopia; the record for O. b. callotis is 110.2 cm for a pair of horns from L. Magadi in Kenya (Rowland Ward) Key References East 1999; Cobb 1976; Kingdon 1982; Lewis 1975; Stanley Price 1978b; Wacher 1986. Tim Wacher & Jonathan Kingdon

Oryx dammah SCIMITAR-HORNED ORYX (SCIMITAR ORYX) Fr. Oryx algazelle (Algazel); Ger. Säbelantilope (Nordafrikanischer Spießbock) Oryx dammah (Cretzschmar, 1827). In: Rüppell, Atlas zu der Reise im nördlichen Afrika von Rüppell, pt 1, Säugethiere, p. 22. Sudan, Northern Kordofan Prov., ‘bewohnen die grossen Steppen von Haraza [vicinity of Jebel Haraza]’. The year of original publication is 1827, not 1826 (see Grubb 2005).

Lateral view of skull of Scimitar-horned Oryx Oryx dammah.

Scimitar-horned Oryx Oryx dammah.

Taxonomy Monotypic. Has frequently been called Oryx leucoryx (which is restricted to the Arabian Oryx), and O. algazel, which is invalid (see Grubb 2005). Ansell (1972) and Corbet (1978) discuss use of O. dammah over O. tao. Synonyms: algazel, bezoartica, ensicornis, nubica, senegalensis, tao. Chromosome number: 2n = 56–58 (Wurster 1972, Gallagher & Womack 1992, Claro et al. 1994, Kumamoto et al. 1999). Majority of specimens have 2n = 58 with fixed centric fusion of chromosomes 1 and 25; variation in diploid number (i.e. 2n = 56–57) is the result of centric fusion polymorphisms of chromosomes 2 and 15 (Kumamoto et al. 1999). Sex chromosomes, X and Y, are acrocentric (Kumamoto et al. 1999). Scimitar-horned Oryx and Addax Addax nasomaculatus hybrids are known and reported to resemble the Addax in European zoos (Ruhe 1993). Hybrids born at Bou-Hedma N. P. in Tunisia (believed to be sired by an oryx !) were large-bodied like oryx, with typical oryx horn shape, but with obvious Addax body colouration and body shape (R. Molcanova pers. comm.).

Description A large, robust antelope, with long horns arched over the back in both sexes, and striking rufous and white coat colouration. Cream to whitish body pelage contrasts with reddishbrown colouration on neck, shoulders and upper legs. Seasonal variation in coat length and colour quite marked, with animals appearing whitest in short bright summer coat, while long winter coat is duller and creamier and may reveal faint reddish-brown lateral flank-stripe and reddish tones on rump. Head is elongated, cream with face ‘mask’ of reddish-brown across muzzle and blaze on forehead, and reddish-brown eye-stripe from base of horn across eye and cheek. Eyes, nostrils, lips and inner ears black. Ventral surface and insides of legs creamy-white, hooves black. Tail long (ca. 39% of HB), and cream with brown–black terminal hairs. Pedal glands present on all four feet, but there is also a small preorbital gland (not present in other Oryx species); inguinal glands absent. Sexes superficially similar, but adult !! larger, with heavier neck and shoulders and thicker horns at all ages. Individuals may be reliably identified by small variations in horn shape and at close range by variation in details of face-mask pattern. The distinctive horns are long, ridged (lower one-half or one-third marked by 30–60 annulated corrugations), sharp-tipped and curved backwards in a shallow arc (80–150 cm), giving rise to the common name. Female horns are longer and thinner, with less prominent annulations; male horns are noticeably thicker at the base, generally more robust and often shorter in older individuals. Differences in horn circumference at the base are apparent in calves at an early stage and can be useful for determining sex. Hard and sharp adult

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horn tips develop after sloughing off the outer keratinous sheaths that cover the horns of young animals. Geographic Variation None recorded. Similar Species Oryx beisa: allopatric in East and north-east Africa only. Horns more or less straight; neck and shoulders lacking reddish-brown colouration; preorbital gland absent. O. gazella: southern Africa only. Horns more or less straight; neck and shoulders lacking reddish-brown colouration; preorbital gland absent. Distribution Formerly widespread in semi-arid Sahelo-Saharan zones north and south of the Sahara Desert from the Atlantic Ocean to the Nile R., but not north of the Grand Atlas Mts. Range countries include Egypt, Libya, Tunisia, Algeria, and Morocco north of the Sahara and Mauritania, Mali, Niger, Senegal, Burkina Faso, Nigeria, Chad and Sudan in sub-Saharan Africa. Historical Distribution In Egypt, the historical range included most of the Western Desert, west of the Nile R., but mostly on oases and wadis. Oryxes were recorded in Siwa and Kharga Oases, El Faiyum area, western Giza and Wadi El Natrun (Osborn & Helmy 1980, Saleh 2001). They were reportedly hunted to extinction in the mid-1800s, with the latest records dating from the 1850s or 1860s; however, there is a record of an individual observed in 1975 in the north-west (Kock 1970, Osborn & Helmy 1980, Saleh 2001). In Libya, the species was described as widespread in the Fezzan (southwest) and Kufra (south-east) regions (Khattabi & Mallon 2001), near known populations in N Chad and the Western Desert of Egypt, respectively; there are unconfirmed historical reports of Scimitarhorned Oryx in NE Libya, from Wadi Jerari (Hufnagl 1972, Khattabi & Mallon 2001). They reportedly became extinct in the 1940s due primarily to illegal, motorized hunting and habitat degradation due to overgrazing by livestock (Newby 1988), although Hufnagl (1972) quotes a possible sighting from NW Libya in 1964. Late nineteenth-century accounts suggest that the Scimitarhorned Oryx may have occurred at least sporadically in S Tunisia, but some doubted that an established population still existed and none were able to provide first-hand information or reliably sourced specimens (Johnston 1898, Sclater & Thomas 1894–1900, Lavauden 1920, 1926b). All published descriptions, including the presence of a stuffed oryx calf in a private museum near Tunis in 1898 (said to have been received alive from the southern frontier), could equally be explained by contact with trans-Saharan traders or the occasional appearance of wandering individuals (Devillers & DevillersTerschuren 2005). In Algeria, oryxes inhabited the sub-desert and steppe regions north and south of the Sahara. They were reportedly hunted to extinction in the 1960s, and the last animals were shot in the Tassili region in 1987, though these may have been vagrants from the Sahel (De Smet & Smith 2001). Oryxes were widely distributed in S Morocco and the Western Sahara, south of the Oued Drâa (Loggers et al. 1992, Aulagnier et al. 2001).They were reported as extinct in Morocco in the 1930s and have not been sighted in Western Sahara since the 1950s, except for a single individual seen in 1973 (Le Houérou 1992).

Oryx dammah

Their historical range is not well documented in Senegal, but they probably occurred throughout the Sahel zone of N Senegal (East 1999), and were hunted to extinction by 1914 (Sournia & Dupuy 1990). In Mauritania, they formerly inhabited the south and western regions, along the Western Saharan frontier (Trotignon 1975, East 1999); they were hunted to extinction by the 1960s (Trotignon 1975, Newby 1988). In Mali, oryxes inhabited the sahel zone in C Mali and north into the desert zone (East 1999). They are presumed extinct since perhaps 1983 (Newby 1988), although a pair was sighted on the Burkina Faso border in 1986 (Duvall et al. 1997). In Burkina Faso, they ranged into the sahel zone in the north, but were hunted to extinction in the 1950s (East 1999). Scimitar-horned Oryxes were widely distributed in sub-desert and sahel zones of C and S Niger and probably occurred in the northern desert zone with seasonal rainfall and good pasture availability (East 1999). The last reported sighting of oryxes in Niger was in 1986 (Millington et al. 1991). Overhunting, the introduction of deep permanent water bores for livestock excluding them from prime habitat, and desertification of the Sahel probably caused their extinction by 1989 (Dixon et al. 1991). Recent surveys report no evidence of live animals in the Termit region and Tin Toumma (Wacher et al. 2004a, 2008, 2009, 2010, Claro 2004). Oryxes formerly inhabited only extreme NE Nigeria, possibly only as a seasonal vagrant (East 1999). Oryxes were historically abundant and widely distributed in the sub-desert and northern Sahel zones of C and N Chad, extending north into the desert zone (East 1999). Reports of migratory herds of hundreds or even thousands of animals in the sub-desert steppe habitat across NC Chad were common during the twentieth century (Gillet 1966a, Newby 1974). A population of >3500 survived under active protection in the Ouadi Rimé–Ouadi Achim Faunal Reserve between 1973 and 1978 (Bassett 1975, Newby 1980), but by 1988 it was reported that only a few dozen survived in the wild (Dixon et al. 1991) and repeated surveys since 1990 have failed to find any evidence 587

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of surviving animals (Monfort et al. 2004, Devillers & DevillersTerschuren 2005, Wacher et al. 2011). In Sudan, Scimitar-horned Oryxes were distributed throughout the entire Sahelian zone of the Darfur and the Kordofan (Devillers & Devillers-Terschuren 2005), and were abundant in the Wadi Howar region near the Chad border. They were reportedly extinct by the mid-1970s (Newby 1988). Current Distribution Today, the Scimitar-horned Oryx is listed as Extinct in the Wild in all former range countries (Devillers & DevillersTerschuren 2005). However, semi-captive populations have been established within areas of natural habitat (ranging from 20–80 km²) in several former range states (mapped), including Tunisia (Bou-Hedma N. P., Sidi Toui N. P., Oued Dekouk N. R. and Dghoumès N.P.), Morocco (Souss-Massa N. P.; extralimital) and Senegal (Guembeul Wildlife Reserve and Ferlo-Nord Wildlife Reserve). In Tunisia, Bou-Hedma N. P. covers 16,448 ha of mountainous and undulating steppe east of Gafsa, and has fenced Total Protection Zones, which are patrolled (Smith et al. 2001). Habitat restoration, particularly of the Acacia raddiana woodland, was very successful (Kacem et al. 1994). In 1985, ten captive-born, young oryxes (five !!, five "") were translocated from European zoos to a 10 ha acclimatization area. The animals were released into a Total Protection Zone (2400 ha) 18 months later (Bertram 1988, Gordon 1991). In 1999, an additional adult ! from Europe was added to the population, individually managed to ensure he bred with established "", with the aim of increasing the genetic diversity (Molcanová 2004). The population has steadily increased from ten in 1985 to 70 in 1996 (Smith et al. 2001), peaking at an estimated 130 in 2005 (T. Gilbert pers. comm.), but had fallen to 73 by 2007 and below 50 in April 2011 (Molcanova & Wacher 2011a). Sidi Toui N. P. lies in SE Tunisia near the Libyan border. The oryx area, established in 1993, covers 6135 ha encircled by a double fence-line. The landscape of low hills surrounded by open plains, small dunes and dry sandy wadis represents typical pre-Saharan steppe (Karem et al. 1993), characterized by a variety of shrubs and dwarf shrubs and a complete absence of tree cover. In 1999, 14 Scimitar-horned Oryxes selected from European zoos to represent a separate genetic line to those at Bou-Hedma N. P. were transported to Sidi Toui N. P. Ten animals (one !, 9 "") were released, while the remaining animals were sent to Bou-Hedma N. P and Oued Dekouk National Reserve (Molcanova et al. 2001). Initial growth of this population peaked at 33 in 2005, but subsequently declined from 2005–2008 coincident with an unexpected swing to a malebiased sex ratio in a classic small population effect. As of 2011, the population size stood at 31 animals (Molcanova & Wacher 2011b). Oued Dekouk N. R., located near Tataouine, is another fenced reserve covering 6000 ha. In 1999, three animals (one !!, two "") from European zoos were transferred to Oued Dekouk with two further "" transferred from Sidi Toui N. P. to Oued Dekouk in 2003 (Molcanova 2006). The population had grown to 21 in 2008 and 26 in 2011 (Molcanova & Wacher 2008, 2011a). In February 2007, eight Scimitar-horned Oryx were translocated from BouHedma N.P. to Dghoumès N.P. (8000 ha), close to Tozeur. In Morocco, a captive population has been established at Souss-Massa N. P., which covers 33,800 ha in the Atlantic oceanic Sahara region, south of Agadir, but which is outside the original distribution range of

the species. The fenced 1500 ha Arrouais Reserve consists of bush and tree savanna, stony and sandy ground and migrating sand dunes with 5 ha pre-release enclosures (Müller & Engel 2004). Three shipments consisting of a total of 29 oryxes (17 !!, 12 "") from European zoos were released between 1995 and 1997 (Engel et al. 2001, Müller & Engel 2004). The population had increased to 50 in 2000 (Müller 2002) and an estimated 240 in 2005 (T. Gilbert pers. comm.) In Senegal, captive populations have been established at Guembeul and Ferlo-Nord Wildlife Reserves. In 1999, eight Scimitar-horned Oryxes (three !!, five "") from the Hai-Bar Wildlife Reserve in Israel were transported to an 8 ha enclosure within the Guembeul Wildlife Reserve (720 ha) at Saint Louis in the north-west region of Senegal (Clark & Bovee 2000). The population had increased to 23 in 2002 with births and the addition of two imported oryxes from Paris Zoo (Clark 2002). Guembeul serves as an acclimatization site for release into the Ferlo-Nord Wildlife Reserve (487,000 ha of sahelian savanna grassland and bushland on gently rolling sandy terrain in NE Senegal). In 2003, eight oryxes were transferred from Guembeul Wildlife Reserve to a fenced core area of 600 ha in Ferlo-Nord (Gueye 2004); all eight animals were born in Senegal and have since produced calves. In 2004, the estimated reintroduced population in Senegal was 30, 18 at Guembeul and 12 at Ferlo-Nord (Gilbert 2004b). Habitat Climatic changes during the last 5000 years left the Scimitar-horned Oryx population divided north and south of the central Sahara Desert. The African Sahel, the sub-Saharan steppe of hardy grasses, shrubs and drought-resistant trees that fringed the southern limits of the desert from Sudan to Senegal, constituted the most important habitat (Newby 1988). A gregarious species, its herd size varied with season, food availability, reproductive activity and human presence. Oryxes exhibited a seasonal, cyclical pattern of movement, driven by the requirement for grazing, water and shade. Grazing resources were dependent entirely on an unpredictable and variable rainfall (100–400 mm annually). In countries south of the Sahara, during the single Sahelian wet season (Jun–Sep), Scimitarhorned Oryxes used to follow the rains, migrating north to graze on freshly sprouted grasses and to areas with access to water. During the cooler months (Nov–Feb), the oryxes reached the Saharan Desert fringes and their winter grazing pastures.With the onset of the hot dry season (Mar–Jun) the grasses desiccated and the oryxes migrated south to the Sahelian dunes in search of perennial species, such as Panicum turgidum and Aristida mutabilis, fallen Acacia pods and persistent foliage of shrubs and herbs (Gillet 1966a, Newby 1974, Dragesco-Joffé 1993). The wooded wadis (ephemeral desert streams) and inter-dunal depressions provided essential shade during the hottest periods, when shade temperatures could reach 40–45 °C, and, as a result, peak feeding activity was at dawn and dusk. Dense shade trees, such as Maerua crassifolia, were particularly sought after, while in sparsely wooded regions even a clump of Panicum turgidum could provide shade (Gillet 1966a). In Chad, the pattern of seasonal movements consisted of annual migrations greater than 600 km (Newby 1988). Abundance Formerly abundant throughout the Sahelian zone, there were probably more Scimitar-horned Oryxes south of the Sahara than in the northern range countries (Newby 1988). Historically, aggregations of several hundred or even thousands congregated at favourable pasture or during the seasonal migrations

588

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(Brocklehurst 1931, Edmond-Blanc 1955, Gillet 1966a, Newby 1974). Oryxes were still considered common throughout northern Africa in the mid-1900s, but herds have not been observed since the 1970s. As late as 1985 there were an estimated 500 animals in Chad and Niger, but by 1988 it was reported that only a few dozen survived in the wild and since then there have been no confirmed reports of any surviving wild oryxes. Sadly, the prediction by John Newby (1978b) that ‘the Scimitar-horned Oryx will be extinct in the wild before the end of the century’ turned out to be true. Adaptations Although the physiology of the Scimitar-horned Oryx has not been subject to investigation like that of its southern or East African counterparts, it is likely that many of the adaptations to survival in arid environments detailed for the Gemsbok by Knight in this volume also hold true for the Scimitar-horned Oryx. They have the ability to survive for months without drinking, obtaining most of their moisture requirements through their food, but will drink freely when water is available. Physiological adaptations and behavioural mechanisms help them control water balance. For example, the light-coloured coat helps reflect and reduce radiant heat and they are inactive during the heat of the day. In extreme heat they seek shade or lie in shallow scrapes in the sand (Newby 1974, R. Molcanova pers. comm.). The coat is short and thick, and grows longer during the winter (Dolan 1966b, Gordon & Wacher 1986). Studies on the related Beisa Oryx Oryx beisa found that dehydrated animals conserve moisture by storing heat during the day (body temperature may exceed 45 °C for eight hours with no apparent ill effect); during the cooler night the heat is transferred back to the environment (Taylor 1969). Body temperature fluctuations are normal in hippotragine antelopes (Kock & Hawkey 1988). Milk analysis of a sample taken three days postpartum consisted of 12.7% fat, 5.4% protein and 24.5% non-fat milk solids (Mayor 1983). Such highly concentrated milk is consistent with the requirement to conserve water in an arid environment. A nomadic species with movements adjusted to seasonal rains, oryxes used to walk long distances to exploit newly sprouted vegetation and water. They have a highly developed sense of sight and smell, sensing changes in the atmospheric humidity (Gillet 1966a) or responding to distant lightning and the ‘smell’ of the rains on the winds (Newby 1974). They have large hooves adapted to walking great distances over sandy or loose, stony terrain, and walk at an amble, nodding their heads while walking fast. Captive animals have adapted readily to life in temperate zones. Foraging and Food Scimitar-horned Oryxes are ruminants and show a pronounced preference for grazing, but also browse. Gagnon & Chew (2000) in their review of the dietary preferences of African bovids classified them as variable grazers. They graze predominantly on a wide variety of grasses, legumes and perennials that vary seasonally in their availability (Gillet 1966a, b, Newby 1974, R. Molcanova pers. comm.). Plants with a high moisture content are selected, such as the leaves and fruit of the wild desert melon Citrullus colocynthis, a characteristic species of the Sahelian sub-desert steppes and an important forage species with its leaves and stems staying green well into the hot season (Newby 1974). The animals graze in the early morning, evening and at night, when plants have been observed to collect small droplets of moisture on the leaves (Gillet 1966a, b).

Plants of the genera Aristida, Brachiaria, Dactyloctenium, Echinochloa, Fagonia, Indigofera, Panicum, Requienia, Stipagrostis and Tephrosia were important food sources in Chad, as were high protein Acacia pods, particularly for lactating "" (Gillet 1966a, b, Newby 1974). Sahelian populations also fed on foliage from persistent shrubs, including Cornulaca monacantha, Chrozophora senegalensis and Cassia italica, and a few herbs, such as Heliotropium strigosum (Newby 1974, Dragesco-Joffé 1993). Animals returned to Tunisia fed primarily on grasses, including the drought-resistant Cenchrus sp., used their horns to hook down Acacia branches and explored a range of small herbs and shrubs. They showed exploratory tasting behaviour of a range of plant species that were unfamiliar when they were first introduced to natural range. They also avoided the toxic species Peganum harmala and Pergularia tormentosa (Gordon 1991, T. Wacher pers. comm.). A list of species known to be eaten by the Scimitar-horned Oryx both in the wild and at release sites for reintroduction programmes is provided by Gilbert & Woodfine (2004). Social and Reproductive Behaviour The most detailed observations of Scimitar-horned Oryx behaviour are those of Gillet (1966a, b) and Newby (1974, 1988) in the wild in Chad and more recent studies in semi-captive conditions in Tunisia (Molcanova & Wacher 2010, 2011a,b, Molcanova et al. 2011). Herd sizes in Chad typically ranged from 10–30 (Talbot 1960, Gillet 1966a, Newby 1974, Wilson 1980). The average of 37 herds observed was 14.8 (range 6–28). Typical herds were mixed sex, comprising at least one old !, with subadult !!, "" and juveniles (Newby 1974). Observations from Tunisia show a similar predominantly mixed-sex structure with a mean herd size of 12 (range 2–30) (Molcanova & Wacher 2011b). Within herds, linear social hierarchies exist, typically with the largest adult !! dominant to adult "", but herd movements are usually initiated by an older dominant " (Newby 1974, Knowles & Oliver 1975, Pfeifer 1981, Mayor 1983, Gordon et al. 1989, Engel 1997, Molcanova et al. 2001, C. Morrow pers. obs.). Generally tolerant relations between !! in mixed-sex herds have been reported in Chad (Newby 1974) and Tunisia. Dominance is primarily established with ritualistic displays. Both !! and "" use spatial displacement, erect posture and lateral horn presentation, often combined with tilting the head to aim the horns toward an opponent. Submission is expressed by turning away with a brief lowering of the head (or, in extreme cases, lying down). Displaying !! may thrash the ground or vegetation with horns from a standing or kneeling position. If ritualized displays are insufficient, equally matched !! fight initially face to face, sometimes kneeling, or swinging round side by side, enabling vigorous over-theshoulder stabbing (Molcanova & Wacher 2011b), the loser breaking off to run away. Fighting may cause horn breakages. Serious injury and/or death has been recorded (Gordon 1989, Blumer et al. 1992, Molcanova & Wacher 2007, 2011a,b). Damage or loss of horns results in reduction in social rank in some circumstances, although some long term territory-holders in Tunisia were also one-horned. Solitary wild !! were encountered in Chad; although not known to patrol or demarcate territorial boundaries, they did make vigorous attempts to keep "" together as a herd (Newby 1974). In semicaptive conditions (Tunisia), territorial behaviour by solitary !! has been described in detail by Molcanova & Wacher (2010, 2011a,b). Territorial !! occupied restricted areas (typically around 8 km2), in 589

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some cases held for periods of more than six years. These !! were frequently alone and areas were defended against !! of similar status. On territory they expressed dominance by characteristic squat defecation displays; on rare excursions off territory (for example, to visit the single drinking point), they defecated in normal standing posture. When herds move onto territories, resident !! are comparatively tolerant of submissive subordinate !! in the absence of receptive "", using ritualized displays to confirm status. Cases of sub-adult !! transitioning to territoriality, associated with a change from herd-based to a more solitary existence and development of restricted movement in an exclusive area have also been recorded. Courtship follows a highly ritualistic pattern: the ! approaches the " with head high, sniffs the anogenital region and may perform flehmen. A non-oestrous " will continue grazing or walk away, the ! losing interest. A " coming into oestrus may lower her head and run a short distance, the ! following. Oestrous "" walk and vocalize more, eat less and solicit attention from the !. The ! will perform flehmen in response to urination (sampling mid-stream or from the ground). The pair engage in lateral head-to-tail circling (‘mating circles’) and the ! will repeatedly perform laufschlag (‘foreleg lift’) to test for female receptivity before attempting to mount. Laufschlag may be performed from directly behind the ", from the side or on an angle, with the ! facing the female’s tail. The ! will also lean heavily and/or push on the female’s hindquarters to test for receptivity. If the " stands during the hindquarter pushing and laufschlag he will perform repeated mounts (82 ± 13, mean ± SEM, range 3–155 in artificially synchronized captive "") followed by a strong pelvic thrust in which his hindfeet may leave the ground. Bouts of courtship activity (4.7 ± 0.7, mean ± SEM, range 1–8 in synchronized "") are interspersed with other activities (grazing, walking, resting). Females separate from the herd for a few days prior to and following parturition. During labour, "" exhibit increased walking and alternating periods of standing and lying. Calves can stand and follow their mothers within hours of birth. Ano-genital licking and grooming is the most common mother–calf interaction and occurs independently of suckling. If in good condition, "" frequently come into post-partum oestrus during this period of isolation. They are usually joined by (or find their way to) an adult ! and form mating consortships (usually with territorial !!) over several days in semi-captive populations. Courtship bouts are frequent and the ! may also tend the calf assiduously at this time, working to keep the mother-calf pair within his territory. Female oryx may choose to associate with different !! during the post-partum period. This can involve running to escape blocking manoeuvres by the !, and may not be without risk. Fresh stab injuries have been noted on "" following successful transition between territorial !! (Molcanova & Wacher 2011b). Between periods of suckling, calves spend most of their time lying out hidden in vegetation with other calves of similar age for 3–8 weeks (Newby 1974, Gill & Cave-Browne 1988). Females typically return to herds with their calves within 2–3 weeks. Older calves may kneel on carpal joints while suckling. Social play in captive calves of both sexes from 2–15 weeks of age consists of running in circles, leg kicking, head tossing, sparring, circling and pawing the ground (Pfeifer 1985). Once returned to herds, older calves often associate together, forming distinctive crèches (for example, resting at one

side while adults graze), or two or three calves may keep company with one adult " (Molcanova & Wacher 2011a). Scimitar-horned Oryxes make a variety of sounds: !! and "" vocalize more when "" are in oestrus and "" are more vocal with calves at foot; !! grunt. Six vocalizations were distinguished from sonograms: adult contact, juvenile contact, adult alarm snorts, calf ‘moans’ and mother ‘purr’ calls (Gill & Cave-Browne 1988). Reproduction and Population Structure Limited data on reproduction exist for Scimitar-horned Oryxes in the wild (see Table 9), but comprehensive research on captive populations has provided data on ovarian cycles, seasonality and assisted reproductive technologies (ovulation induction, semen collection and cryopreservation procedures) (see Morrow 1997, Morrow et al. 1999, 2000 for reviews). Captive oryxes are seasonally polyoestrus, spontaneous ovulators with a 23.4 ± 1.3 day ovarian cycle (luteal phase 18–20 days) and intermittent, short, 8–12 day cycles; oestrus lasts 3–41 hours (Morrow 1997). A loosely synchronized spring anovulatory interval of 36–95 days was recorded in a small captive herd (Morrow et al. 1999). Ultrasonography and elevated periovulatory and luteal phase oestrogen concentrations suggests follicular recruitment throughout the ovarian cycle. Gestation ranges from 222 to 257 days (median 250 days) in captivity and 258 to 274 days in the wild (Table 9). Twin births are uncommon (0.7%; North American Studbook), and twin litters have a high mortality rate (56%), reflecting anatomical limitations imposed by a duplex uterus. Postpartum oestrus occurs in wild (Newby 1974), reintroduced (Gordon 1991, Molcanova & Wacher 2007) and captive (Knowles & Oliver 1975, Nishiki 1992) populations. Oryxes in Chad and those re-established in Tunisia exhibit asynchronous breeding in favourable climatic and nutritional conditions with births every 8–10 months (Gillet 1966a, Newby 1988, Gordon 1991, R. Molcanova pers. comm.). At Sidi Toui N. P., over the course of seven years (n = 40 births), calvings peaked in Mar and Oct, with births recorded in all except the hottest two months of Jun and Jul (Molcanova 2006). In captivity, calves have been born every month of the year (Table 9) and it is recommended that the breeding ! be removed at certain times of the year to manage births. However, analysis of North American Studbook records of unmanaged breeding herds confirm an 8–11 month reproductive periodicity; "" that calved in the winter/spring had a longer interbirth interval than "" that calved in the summer/autumn (Morrow et al. 1999). Median inter-birth interval was 277 days with 75% of intervals less than 11 months. Calves average about 8–9 kg at birth (range 5–11 kg), and are a light gingery brown colour for the first 2–3 months, with a narrow whitish area on the belly, and faint hint of adult face pattern. The cryptic colouring serves as an anti-predator adaptation during the lying-out phase. Horn buds are visible at birth. Adult colouration is attained between three and 6 months of age, and weaning occurs between six and ten months (Molcanova & Wacher 2010). Estimates of age of sexual maturity range between 10 and 27 months for "" and 22 and 30 months for !! (Table 9). Although no comparative data exist for wild animals, in a semi-captive population a " reproduced at 17 years (Molcanova & Wacher 2011b) and captive "" in good health can continue to reproduce past the age of 20 years. The high reproductive potential is due to a combination of low

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Table 9. Reproductive data for Scimitar-horned Oryx. Oestrous cycle (d)

Popn

Location

Latitude

Birth season

Wild Wild

Sudan Chad

14° N 13–16° N

Wild

Chad

14–17° N

Wild

Ennedi, Chad Bou-Hedma, Tunisia

15–18° N

In situ

Sidi Toui, Tunisia

32° N

Ex situ Captive Captive

Hai Bar, Israel Edinburgh, UK Germany

30° N 55° N 52° N

May Jul–Aug 8–10 month peaks Jan–Feb, Sep–Oct, May–Jun Jan–Apr Mar, Apr, Jul, Sep, Nov Dec, Mar, Apr, Aug, Sept Aug–Apr All year

Captive

London, UK

51° N

Captive

Ohio, USA

41° N

Captive

Virginia, USA

38° N

Mar–Aug

Captive Captive

Tokyo, Japan San Diego, USA

35° N 32° N

Nov–Mar

Captive

New Zealand

43° S

In situ

34° N

All year a

Gestation (d)

Sexual maturity F M

274 274 258

Brocklehurst 1931 Gillet 1966a, b >24

277, 289 222–253 242–256

R. Molcanova pers. comm. 18–25 27

247, 249

11–27

247.6

22 11

253, 257 b

> 12 a

21–22

mortality, adult longevity and an 8–11 month inter-birth interval. A captive animal was still alive after 27 years (Weigl 2005). Although sex ratios in small captive populations have been reported to be nearly 1 : 1 (Yoffe 1980, Gill & Cave-Browne 1988, Nishiki 1992), an analysis of 1129 births suggested that sex ratios in North America (1985–94) were male-biased (1 : 0.84) (Morrow et al. 1999). Predators, Parasites and Diseases Few large predators inhabited the arid environment of the Scimitar-horned Oryx, but there were once overlaps in range. Spotted Hyaenas Crocuta crocuta, Striped Hyaenas Hyaena hyaena, African Wild Dogs Lycaon pictus, Cheetahs Acinonyx jubatus, Golden Jackals Canis aureus and Lappetfaced Vultures Torgos tracheliotus were likely predators of young and infirm animals (Gillet 1966a, Newby 1974). Death of adults and calves from hunger, heat exhaustion and disease was common during droughts (Newby 1988). Although survival of calves in the wild was high, calf abandonment was also high during drought conditions (Newby 1988). In reintroduced semi-captive populations, calf deaths have resulted from starvation (mismothering or insufficient lactation; Gordon 1991, Molcanova et al. 2001), predation by Golden Jackals (Molcanova et al. 2001, Molcanova 2002) and aggression by dominant !! (Gordon 1991). In captivity, calves have in some cases suffered high mortality attributed to inbreeding (Mace & Pemberton 1988, Nishiki 1992). Normal haematological and serum chemistry values have been examined in captive populations (Bush et al. 1983, Hawkey & Hart 1984, Ferrell et al. 2001, Flach 2004a). Deaths from Parelaphostrongylus tenuis (meningeal worm) infection (Ferrell et al. 1997), disseminated intravascular coagulation (Pearce et al. 1985), perforation of the large bowel (Mayor 1983), bovine spongiform encephalopathy (Kirkwood

Newby 1974

Gordon 1991

247

22.7 ± 1.1a

22– 30

Edmond-Blanc 1955

24.4 ± 2.2 b

23.8 ± 1.3; short 8–12

References

Yoffe 1980 Gill & Cave-Browne 1988 Dittrich 1972 a Zuckerman 1953, Anonymous 1961, Kirkwood et al. 1987, bShaw et al. 1995 Pope et al. 1991, Morrow et al. 2000 Morrow & Monfort 1998, Morrow et al. 1999, 2000 Nishiki 1992 Durrant 1983 a Bowen & Barrell 1996, b Garland et al. 1992

& Cunningham 1999), and serious internal injuries from horn punctures (C. Morrow pers. obs.) have been reported for captive individuals. Tuberculosis is also a potential problem in captivity (Greth et al. 1994). Clinical malignant catarrhal fever, bovine viral diarrhoea virus, yersiniosis, gastrointestinal helminths, coccidial oocysts, acidosis, copper deficiency and hoof problems have been reported (Flach 2004a). Reported deaths of Scimitar-horned Oryxes at Hai-Bar N. R., Israel, have included coccidiosis, mouth tumour, parturition and worms (Yoffe 1980). Immobilization and anaesthesia protocols are well established (Kock & Hawkey 1988, Pearce & Kock 1989, Roth et al. 1998, Morrow et al. 2000, Flach 2004b). Because of the ability of desert antelopes to accumulate body reserves, individuals may be prone to obesity in captivity. Common veterinary treatments include trauma (broken horns, puncture wounds, lacerations) and hoof/joint disorders (lameness). Conservation IUCN Category: Extinct in the Wild. CITES: Appendix I. CMS: Appendix I. The reasons for the catastrophic decline of the Scimitar-horned Oryx are well documented and include habitat loss (desertification due to repeated droughts, development of permanent boreholes and resulting competition with domestic stock and human disturbance), civil war and exploitation through motorized hunting (Newby 1978b, 1980, Mallon & Kingswood 2001a, Devillers & DevillersTerschuren 2005). Their meat and hide was prized by local people, the thick hide being used for ropes, bags, shoes and shield coverings. They were ruthlessly hunted for their magnificent horns by trophy hunters (Gillet 1966a, Barrett 1967, Newby 1978b). Hunting parties using all-terrain vehicles and modern firearms replaced the 591

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traditional method relying on spears and camels, horses and/or dogs (Brocklehurst 1931, Talbot 1960). Between 1973 and 1978 a wild population flourished (>3500) under active protection in Ouadi Rimé–Ouadi Achim Faunal Reserve in Chad (Bassett 1975, Newby 1980), but between 1978 and 1986 protection in the Reserve was abandoned because of political unrest. The last known photograph of wild Scimitar-horned Oryx, a group of four, was taken by John Newby in the Aïr–Ténéré of Niger in May 1980 (Newby et al. 2004). As with other ‘last observations’ in Algeria and Egypt (see Distribution), it seems likely that these individuals may have been wandering far from normal locations. Despite sporadic casual reports, no wild Scimitar-horned Oryxes have been confirmed alive in the last 15-odd years. Extensive surveys dedicated to detection of Sahelo-Saharan antelopes carried out in Chad and Niger have produced only five fragments from former haunts in the Ouadi Rimé–Ouadi Achim Faunal Reserve and the central Termit Mts of Niger (Newby et al. 2004, Wacher et al. 2004a, 2011). No evidence of living oryxes has been obtained. The Scimitar-horned Oryx has thrived in captivity, and in 2008 it was estimated that there were 1675 animals in 204 institutions (Gilbert 2008). There are perhaps more than 4000 in private collections. However, despite being well represented in captive populations, almost all individuals are derived from a genetic base of fewer than 40–50 founders captured in Chad in the 1960s (Bertram 1988, Dixon et al. 1991). A large population of Scimitar-horned Oryxes (>2000) in private collections in the United Arab Emirates are descended from animals captured in N Sudan representing a separate genetic line than the European Endangered Species Programme (EEP) and American Zoos and Aquaria’s Species Survival Plan (SSP) populations (R. Molcanova pers. comm.). In captivity, small, fragmented populations are vulnerable to losses in genetic diversity and fluctuations in size, age and sex ratios. This vulnerability was recognized in the early 1980s, and breeding plans were instituted in North America (SSP; Species Survival Plan), the United Kingdom and Europe (EEP; Europäisches Ehrhaltzungszucht Programm), and Australasia (ASMP; Australasian Species Management Plan) to maximize genetic variation from different founder lineages. Despite being well organized on a regional and global basis, the conservation of this species presents a paradox to zoo managers and conservation biologists. Existing genetic management strategies necessitate the exchange of valuable breeding stock; however, insufficient funding and enclosure space can preclude breeding among genetically valuable pairs. Reproductive technology (e.g. artificial insemination) has proven potential for overcoming the risk and expense of transporting live animals, and for optimizing the use of limited enclosure space, while simultaneously preserving gene diversity (Garland et al. 1992, Morrow et al. 2000). Largely due to the success of rearing this species in captivity, populations of Scimitar-horned Oryxes have been re-established in protected, fenced enclosures in Tunisia, Senegal and Morocco (details provided in Distribution). While successful, these populations represent an intermediate stage between captivity and true reintroduction in range countries. Conservation of aridland antelopes

by necessity requires protection of large areas that could never be fenced to accommodate the seasonal migratory behaviour to pastures following the rains. Human encroachment, overgrazing, reduction of tree cover (shade) and the lack of resources have hampered efforts to create protected areas for Scimitar-horned Oryxes (Wacher 2001, Devillers & Devillers-Terschuren 2005). Plans to establish founder populations and a metapopulation management plan within range countries with the ultimate goal of restoring free-living, self-sustaining populations outside of protected areas are gaining momentum (Houston 2003, Wakefield 2003, Woodfine & Engel 2004). The conservation of this species and other Sahelo-Saharan antelopes was the focus of a workshop convened by the Secretariat of the Convention on the Conservation of Migratory Species and the Institut Royal des Sciences Naturelles de Belgique at Djerba,Tunisia in 1998. This workshop developed an Action Plan for the Conservation and Restoration of Sahelo-Saharan Antelopes and adopted the Djerba Declaration for improving the conservation status of these species (UNEP/CMS 1998, 1999). In 2003, a second seminar on the Conservation and Restoration of Sahelo-Saharan antelopes and their habitats was held at Agadir, Morocco, to review and update the activities and projects in the previous five years (UNEP/CMS 2004). The Sahelo-Saharan Interest Group (SSIG) was established in 2000 as a network of institutions and individuals keen to conserve SaheloSaharan wildlife and the habitats they require for survival (Monfort 2000).The SSIG works to maintain living, healthy deserts that sustain both the wildlife and people who rely on those ecosystems for their livelihood and survival (Monfort & Correll 2004). The SSIG holds an annual meeting and members are actively involved in conservation, research and training across the Sahelo-Saharan regions. In 2005, the SSIG unveiled the Sahara Conservation Fund, an international, nongovernmental organization committed to conserving the wildlife of the Sahara and bordering Sahelian grasslands. SCF is spear-heading an initiative to reintroduce Scimitar-horned Oryxes to their former stronghold in Chad, Ouadi Rimé–Ouadi Achim Faunal Reserve. Measurements Oryx dammah HB (""): 1360 (1200–1620) mm, n = 31 T (""): 530 (440–600) mm, n = 26 Sh. ht (""): 1150 (1060–1210) mm, n = 31 WT (""): 151.0 (129.0–177.0) kg, n = 19 WT (!!): 153.0 (144.0–166.0) kg, n = 3 North American captive population (C.J. Morrow pers. obs.; calf data NZP/CRC records 1980–1992) Maximum recorded horn length is 127.3 cm for an animal from Fada, Chad (Rowland Ward) Key References Devillers & Devillers-Terschuren 2005; Gilbert & Woodfine 2004; Gillet 1966a, b; Mallon & Kingswood 2001a (and chapters therein); Morrow et al. 1999, 2000; Newby 1974, 1988. Catherine Morrow, Renata Molcanova & Tim Wacher

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Tribe CAPRINI Sheep, Goats Caprini Gray, 1821. London Med. Repos. 15: 307.

The Caprini are a large tribe of bovids with 34 living species (Grubb 2005).They are largely native to Eurasia and the Rocky Mts, and form a monophyletic unit in morphological and molecular phylogenies (Hernández Fernández & Vrba 2005, Ropiquet & Hassanin 2005a, Hassanin et al. 2012).The latter authors have constructed a molecular tree that presents an improved picture of how the species and genera are related to one another within the tribe. Some taxonomists prefer to place caprines in their own subfamily, but we have followed previous practice and Ropiquet & Hassanin (2005a) in assigning a tribal level to this diverse and interesting group. In earlier treatments, as a subfamily, several tribal subdivisions were recognized. The most recent classification by Grubb (2005) recognized four tribes: (1) Naemorhedini, including Capricornis (serows) and Naemorhedus (gorals); (2) Ovibovini, including Ovibos (Muskox) and Budorcas (Takin); (3) Caprini, including Capra (goats and ibexes), Ammotragus (Barbary Sheep), Hemitragus (tahrs), Pseudois (Blue Sheep), Ovis (sheep), Rupicapra (chamois) and Oreamnos (Rocky Mountain Goat); and (4) Pantholopini, including Pantholops (Chiru or Tibetan Antelope). However, the molecular results of Ropiquet & Hassanin (2005a) indicate that all tribes classically defined in the literature are not monophyletic, thereby supporting the inclusion of all caprine species into a unique enlarged tribe Caprini (and see Hassanin et al. 1998, 2012, Hassanin & Douzery 1999). This phylogeny also supports the inclusion of the Chiru Pantholops hodgsoni in the caprines (and see Gatesy et al. 1997, Vrba & Schaller 2000). Only three caprines are found in Africa, namely Nubian Ibex Capra nubiana, Walia Ibex Capra walie and the Aoudad or Barbary Sheep Ammotragus lervia. Of these, only the Walia Ibex can be regarded as nonPalaearctic. Himalayan Tahr Hemitragus jemlahicus, a species native to the Himalaya of China, N India and Nepal, were introduced to the Western Cape Province of South Africa around 1930 (see below). Outside Africa, caprines range in size from around 30 kg (Naemorhedus spp.) to as much as 350 kg (Muskox Ovibos moschatus); "" may attain only about 60% of the body weight of adult !!. As with other bovids, the presence of horns is marked by strong interspecific, inter-sexual and individual variation; male horns are often

long or large, notably in the sheep and goats. Sinuses within the frontal bones spread far up the length of the horn cores. This is an obvious weight-reducing benefit for agile animals in which the !! fight by ramming and clashing their horns (Schaffer & Reed 1972), although similar fighting techniques also occur in living Bovini (cattle). Pelages include fine undercoats that become the woollen fleece of domestic sheep or the ‘pashm’ of Kashmir (cashmere) goats. Males often have ruffs, manes, fringes or beards. There are no face, pedal or inguinal glands, and "" have a single pair of inguinal nipples. The molar teeth have a simple clear-cut occlusal pattern. They are extremely similar in sheep and goats despite the former being primarily a closecropping grazer and the latter more catholic in its diet. Metapodials are often shortened and the body conformation is stocky. Chromosome numbers differ among the various species karyotyped to date, ranging, for example, from 2n = 60 in Capra, 2n = 52–58 among members of the genus Ovis (and 2n = 58 in the Barbary Sheep Ammotragus) and 2n = 42 in the Mountain Goat Oreamnos americanus (e.g. Schmitt & Ulbrich 1968, Wurster & Benirschke 1968, Nadler et al. 1974). Schaller (1977) described caprines as generalized and flexible feeders, living in habitats of simple structure and in areas of low primary productivity. He pointed out that they are often the only ruminant in a particular area or habitat: thus, caprines have a demonstrable association with mountain ranges, high plateaux, precipitous cliffs, uneven hilly country or jebels. Some of these habitats may be wooded, but they are more often barren, rocky, open and of low productivity. Each regional caprine tends to be an adaptable, medium-sized herbivore with a broad niche in a simple, impoverished habitat. Caprines have not flourished in the richer Eurasian habitats, where they are generally replaced by deer (which have made ecological radiations comparable to those of some African antelopes). Some Eurasian caprine distributions look like a relict pattern resulting from withdrawal in the face of human exploitation. While this may often be true, the animals’ anatomical peculiarities and agility on rocky terrain reflect long-term adaptation to such habitats. Their pre-human distributions were more likely an evolutionary response to competition from other ungulates (Kingdon 1982).

Bovine fighting techniques: Syncerus (below right) compared with Ovis (top right) and Nubian Ibex Capra nubiana (left).

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Fossils of Caprini are reasonably well known from Eurasia and sometimes occurred additionally in North Africa. It would seem that from about 12 mya onwards, different bovids throughout Eurasia, for example Aragoral and Protoryx, were gradually acquiring characters of the diverse Caprini that later inhabited the same land mass (Gentry 2000). Parallel evolution was evidently rife in the acquisition of characters such as anterior keels on the horn cores or shorter metapodials. Extinct relatives of Ovibos were widespread in the later Pliocene, even into North America, and Caprini had entered Africa by the start of the Pliocene. Better-known examples include the South African later Pliocene ovibovine Makapania broomi, a Pliocene species of Budorcas from Ethiopia, and the puzzling Nitidarcus asfawi from the Pleistocene of Ethiopia (Vrba 1997). Although Pliocene and Pleistocene sheep are not very common and are unknown in Africa, goats or goat-like forms are known from Ethiopia in the middle Pleistocene (Bouria anngettyae; Vrba 1997) and from North Africa in the later Pleistocene. It is thought that humans began exerting some control over herds of wild Caprini many millennia ago, perhaps around 8000 BC in Africa, and evidence of domestic sheep is known from the southern tip of Africa about 500 years BC. In the early 1930s, Groote Schuur Zoological Gardens, situated on the slopes of Devil’s Peak, Table Mountain, near Cape Town in the

Western Cape Province of South Africa, obtained a pair of Himalayan Tahrs from the National Zoological Gardens, Pretoria. The pair escaped from their enclosure and established themselves either on Devil’s Peak or on Table Mountain. In 1972, there were an estimated 330 individuals (Lloyd 1975), by which time Himalayan Tahrs occurred across much of the north/north-west and south/south-east faces of both Devil’s Peak and Table Mountain (and the Saddle linking them) and the north-west faces of the Twelve Apostles range. This uncontrolled increase in the numbers of Himalayan Tahrs, within a Nature Reserve and especially a reserve with a unique flora, posed a serious threat and the local conservation authority, Cape Nature Conservation, initiated a research study to investigate the problems involved. This survey recommended that, as total eradication was probably impossible, there should be a drastic reduction in the population (Lloyd 1975). Between 1975 and 1981 over 600 tahrs were removed by Parks and Forestry officials and members of Cape Nature Conservation. A survey conducted by the Mountain Club and officials of the Parks and Forestry showed that at the end of this period, 88 tahrs were still living on Table Mountain (see Skinner & Smithers 1990 for further discussion). Michael Hoffmann & Jonathan Kingdon

GENUS Ammotragus Aoudad Ammotragus Blyth, 1840. Proc. Zool. Soc. Lond. 1840: 13.

Ammotragus includes only the Aoudad Ammotragus lervia, which inhabits montane desert habitats in northern Africa. The classification of the Aoudad has changed over the years since it was first included in the genus Antilope, with its affinities with Ovis and Capra, in particular, having been widely discussed in the literature. Several taxonomic works have included the species in the genus Capra (Ansell 1972, Corbet 1978), with which it shares a series of morphological traits. Aoudads may interbreed with domestic goats and produce live and fertile offspring (Petzsch 1957, Van Gelder 1977b), although this hybridization does not occur readily. However, Aoudads also share a series of sheep-like characteristics with Ovis (Valdez & Bunch 1980), and one author has suggested it is related to the common ancestor of both goats and sheep (e.g. Geist 1971), due to its unique morphological and behavioural characteristics, which, along with fossil remains, would place it near the origin of the Palaeartic sheep lineage, close to the rupicaprids. Analysis of proteins has only served to confuse matters, with seroprotein (Schmitt 1963) and immunoglobulin (Curtain & Fudenberg 1973) analyses revealing a closer relationship to Ovis than to Capra, while the sequence of amino acids of several haemoglobin chains (Manwell & Baker 1975) showed a closer relation to Capra as well as some unique characteristics. Immuno-diffusion studies by Hight & Nadler (1976) paradoxically establish a closer relationship between Ovis and Capra than between either of them and Ammotragus.

A study on phylogenetic relationships based on the comparison of 12S rDNA sequences between eight caprine species, including Ammotragus, Pseudois, Capra aegagrus and five Ovis species, show two distinct clusters, one formed by all Ovis species and another grouping the other three genera together (Ludwig & Fischer 1998). A complete estimate of the phylogenetic relationships in the Ruminantia unequivocally supports the grouping of Ammotragus with the goats and tahrs (Hemitragus) (Hernández Fernández & Vrba 2005), although its own morphological, biological, ecological and behavioural characteristics suggest that it should be placed in its own genus Ammotragus. A molecular phylogeny of caprines groups Ammotragus with Capra, Hemitragus and Pseudois (Ropiquet & Hassanin 2005a), with Ammotragus most closely allied with the Arabian Tahr Arabitragus jayakari (Ropiquet & Hassanin 2005b). Common names referring to either sheep (genus Ovis) or goats (genus Capra) are misleading and names of an Arabic origin should preferentially be used, for example, Aoudad or Arrui. Ammotragus differs from goats and sheep in its long face and sharply bent-down braincase, long neck fringe and relatively long tufted tail. It differs additionally from sheep in that the wrinkling on horns does not form prominent ridges, the horns are not keeled, in the presence of a beard and the absence of preorbital and interdigital glands. Peter Grubb & Michael Hoffmann

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Ammotragus lervia AOUDAD (BARBARY SHEEP, ARUI) Fr. Mouflon à manchettes; Ger. Mähnenspringer (Mähnenschaf) Ammotragus lervia (Pallas, 1777). Spicil. Zool. 12: 12. ‘Africae borealori propria’; restricted to Algeria, Department of Oran (Harper 1940).

Aoudad Ammotragus lervia male.

Aoudad Ammotragus lervia female.

Taxonomy Polytypic species, with six subspecies described (Allen 1939, Ansell 1972, Gray & Simpson 1980), though not without controversy (see, for example, Ellerman & Morrison-Scott 1951). The somewhat vague morphological differences between them (see below), the fact that most of the subspecies were defined from a few specimens, and the presence of several potential areas of hybridization, makes it difficult to expound reliably on the ranges of the different subspecies. A reassessment of subspecies boundaries is clearly necessary. Synonyms: angusi, barbatus, blainei, fassini, jaela, ornata, sahariensis, tragelaphus. Chromosome number: 2n = 58, with a large acrocentric X chromosome and a minute bi-armed Y chromosome (Nadler et al. 1974, Bunch et al. 1977), identical to that of Ovis aries cycloceros and O. a. vignei.

of these glands lends support to the argument that Ammotragus is a distinct genus. Both sexes have horns, relatively large and moderately long (even in "", compared with goats and sheep), with a somewhat high spiral angle. Horns are elliptical and keeled in cross-section, with a broad frontal surface, and have numerous shallow and uniform sulci as well as periodic growth checks or annuli (Ogren 1965, Schaffer & Reed 1972). The horns curve outwards, backwards and point inwards towards the neck, tending to converge over the nape in mature !!. The skull is distinguished by the placement of the foramen magnum beneath the horn bases, shaping the cranium accordingly, and the presence of sinuses covering most of the brain as well as complex septa. Aoudads share this advanced skull pattern with the Himalayan Tahr Hemitragus jemlahicus, bharals Pseudois spp. and Ovis ammon. Also, along with the Himalayan Tahr, Aoudad horn bases are displaced behind the eye orbit, while in most Caprini they are usually placed above its posterior half. Finally, the species is characterized by extensive cornual sinuses. All these characteristics are less pronounced or moderate in "" compared with !! (Schaffer & Reed 1972).

Description The Aoudad is goat-like in proportions, with head relatively long, legs short and stocky, and tail long and naked underneath. A characteristic feature is the mane, which extends down from under the throat along the front of the neck to the chest, and continuing down each of the forelegs in mature animals. This hair pattern on the legs is referred to as leggings or chaps. A mane is also found in wild sheep such as Ovis orientalis and O. aries, but chaps are unique to Aoudads. They do not have a typical goatee, and, like some wild sheep, they have a haired chin and a short erect dorsal fringe extending from the neck to the middle of back. The genus name Ammotragus means ‘sand goat’, probably in reference to the colour of the pelage, which is pale, tawny-brown grading to a whitish underside with dark brown areas about the head and shoulders, although variability in colour tones is notable among the subspecies (see below). Unlike sheep, the Aoudad lacks preorbital, interdigital and flank glands, but it has subcaudal glands. The lack

Geographic Variation A. l. lervia (Atlas or Moroccan Aoudad): Morocco (and possibly Western Sahara), N Tunisia and N Algeria. Horns scarcely, if at all, depressed; face with an indistinct dark median stripe; beard uniform sandy; body mid-tawny. A. l. ornata (Egyptian Aoudad): formerly quite widespread throughout the Eastern and Western Desert of Egypt. Horns strongly depressed, turning sharply downwards before bending 595

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Lateral view of skull of Aoudad Ammotragus lervia.

backwards; face with no stripe; beard uniform sandy; body sandy rufous. A. l. sahariensis (Saharan Aoudad): C and S Algeria, SW Libya, Mauritania, Mali and Tibesti Massif and Ennedi Plateau in Chad; presumed to be the subspecies in Western Sahara, but limits with A. l. lervia unclear. Horns like A. l. ornata; face with no stripe; ears with a white patch below; beard uniform sandy; body pale sandy rufous. A. l. blainei (Kordofan Aoudad): E Sudan. Horns strongly depressed but not bent backwards so much as in previously named subspecies; face with no stripe, but dark owing to admixture of blackish hairs; beard on sides of lower jaw nearly black; body brownish-grey, less rufous than any previously named subspecies; mane brownish. A. l. angusi (Aïr Aoudad): originally in Aïr Massif (Niger) and Termit Massif. Horns much more upright on head than previously named subspecies and curving further backwards and inwards; face with no stripe; beard on sides of lower jaw cinnamon-rufous; body very deep rufous, darker than A. l. ornata, dorsal fringe mixed with black, more strongly on front half; chaps sparse. A. l. fassini (Libyan Aoudad): originally in extreme S Tunisia and Libya. Horns less depressed than A. l. ornata and A. l. sahariensis and more backward-turned than A. l. blainei; face with no stripe, but dark due to mixture of brownish and rufous hairs; ears with a dark patch below; beard black, and on lower jaw mixture of tawny and brown; body light tawny-reddish, more rufous than A. l. sahariensis and A. l. blainei. Similar Species Capra nubiana. Confined to Egypt, Sudan and Eritrea, its distribution may overlap with that of the Aoudad, particularly in some mountainous areas of the western coast of the Red Sea: at the edge of the Egyptian Eastern Desert, and a few areas in Sudan (see below). Similar in size to the Aoudad (!! weighing around 90 kg), but shows a characteristic long goatee in mature !!, lacking any mane or the like; male horns are long (up to 1400 mm in some individuals), projected upwards and forming a semi-circle over its back, whereas the " of this species has shorter horns that grow only up to 380 mm in length. Distribution

Endemic to North Africa.

Historical Distribution Formerly widespread in any rugged and mountainous terrain from deserts and semi-deserts to open forests in North Africa (Brentjes 1980, Shackleton 1997). Known also within its present range from late Pleistocene fossils, Holocene rock art and Ancient Egyptian mummified remains.

Ammotragus lervia

Current Distribution The distribution of the six subspecies, while imperfectly known, can be summarized as follows. Atlas or Moroccan Aoudads are found in the mountains of Morocco, except the western half of the Rif (Aulagnier & Thévenot 1997, F. Cuzin pers. comm.), as well as the northern part of Algeria (Kowalski & Rzebik-Kowalska 1991, De Smet 1997b) and Tunisia (De Smet 1997a), where more than 30 years ago the Aoudad was approaching extinction (Schomber & Kock 1960). The boundary between this subspecies and the Saharan Aoudad in Morocco/Western Sahara is unclear. It has been presumed to be the subspecies imported to European zoological gardens at the end of last century, and from there to American zoos about 1900 (Gray 1985). This subspecies would then form the basis of free-ranging populations inhabiting the S USA (Gray 1985). However, no conclusive evidence to support the notion that this was the sole subspecies used in extralimital transplants has been found (J. Cassinello pers. obs.). On the contrary, Ogren (1965: 9) stated that the pelage of some of the individuals introduced to New Mexico resembles each of the six recognized subspecies, which suggests the existence of hybrid forms. Gray (1985) also supposes that this is the subspecies introduced in the Sierra Espuña mountains of Murcia (Spain), a population that has expanded throughout the whole south-east of the country (Cassinello 2000, Cassinello et al. 2004). The Egyptian Aoudad is native to deserts in Egypt (see Gray 1985), and was assumed to be extinct (Amer 1997). However, there are reports of its continued presence in remote areas, including herds located in the extreme southern region of the Egyptian Eastern Desert (mainly within the boundaries of the Gebel Elba Protected Area) and the Gebel El Uwienat and Gilf El Kebier Plateau (Western Desert, at the extreme southern corner of the country) (M. A. Saleh pers. comm.). In addition, Wacher et al. (2002, and references therein) also report evidence of the presence of Aoudads in both the Elba Protected Area and the Western Desert between 1997 and 2000. Manlius et al. (2003) and Manlius (2009) provide further information on their distribution in these remote areas.

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The Saharan Aoudad has a very large geographic distribution that, according to Gray (1985), includes parts of S Morocco and Western Sahara (the boundary with the Atlas Aoudad is unclear; see also Aulagnier & Thévenot 1997), the Sahara of S Algeria (Kowalski & Rzebik-Kowalska 1991; although De Smet 1997b only accounts for A. l. lervia in Algeria), SW Libya (see also Shackleton & De Smet 1997), the mountains of the Adrar de Iforas in Mali (Lamarche 1997a), N Niger (although Magin & Newby 1997 only account for A. l. angusi in Niger), and Mauritania (Lamarche 1997b). According to Mekonlaou & Daboulaye (1997), this subspecies used to be widespread in Chad, but today it is probably restricted to the sandstone massifs in Ennedi (Alados et al. 1988 reported this population as A. l. blainei) and the Tibesti Massif in NW Chad. The species was recently reported from Western Sahara (Cuzin 2003), where there have been no reliable reports of Aoudad since the surveys of Valverde (1957). This is presumably the subspecies that has been successfully breeding in captivity in a public research institute in SE Spain since 1975 (EEZA, CSIC, Almería, Spain, see, for example, Alados et al. 1988). The Kordofan Aoudad was once relatively widespread from W Sudan to the Red Sea coast, but currently is probably restricted to the Red Sea hills of E Sudan (Nimir 1997). According to Alados et al. (1988), and contrary to Mekonlaou & Daboulaye (1997), this is the subspecies that may occur in the Ennedi and Uweinat mountains in NE Chad. This may be the form in SE Libya. In 1923, A. l. blainei was introduced into the Sabaloka reserve on the Sixth Cataract of the Nile (Gray 1985), where it no longer survives. The Aïr Aoudad inhabits the Aïr Massif in Niger (Magin & Newby 1997). There is a small but apparently viable population that inhabits the Termit Massif region in Niger, isolated from others geographically and probably since long ago, thus of extreme interest both from conservation and taxonomic perspectives (J. Newby pers. comm., T. Rabeil pers. comm; see also Claro & Sissier 2003, Claro 2004, Wacher et al. 2008). According to Alados et al. (1988) this is the form in the Tibesti in Chad, not sahariensis (see above). The Libyan Aoudad is found in Libya (Hufnagl 1972, Shackleton & De Smet 1997) and in the extreme southern part of Tunisia (Gray 1985, De Smet 1997a). Like the Saharan Aoudad, this subspecies was introduced into the EEZA, but the whole population was moved into the Barcelona Zoo, also in Spain, in the late 1980s. As noted above, Aoudads have been successfully introduced as a game species in mountainous desert regions of Texas, New Mexico and California (USA), N Mexicao and S Spain. Habitat Aoudads tend to inhabit rocky and precipitous terrain, from near sea level up to the summits of the Aïr Mts in Niger, which may reach 2100 m (Magin & Newby 1997) and to snow-free areas at about 4100 m in the Atlas Mts (Cuzin 2003). They also require at least some tree cover for shade, and might wander far from water sources for long periods of time. No detailed studies have been carried out in Africa, but empirical data obtained from populations introduced in USA (Johnston 1980) and Spain (Cassinello 2000) revealed the following habitat preferences according to season: open lands and protective rocky slopes during the breeding season (spring), woodlands during summer and grasslands during rutting (autumn) and winter, although in dry regions Aoudads can also be seen in dry forest areas during rutting. A recent study on habitat suitability of the exotic free-ranging population inhabiting SE

Spain shows an impressive expansion potential and adaptation to different environmental conditions, but constrained by low winter precipitation, high altitude, high terrain slope and the presence of forest (Cassinello et al. 2006). Abundance There are no total population estimates available, but numbers are probably in the order of 5000–10,000 individuals. The total population in Morocco has been estimated at between 800 and 2000 individuals (Cuzin et al. 2007), which is higher than that suggested by Aulagnier & Thévenot (1997) at 800–1000. Several thousand individuals survive in Algeria (De Smet 1997b), while low numbers occur in Chad (Mekonlaou & Daboulaye 1997), the Adrar des Iforas in Mali (Lamarche 1997a), and in Mauritania (Lamarche 1997b). In Niger, some 3520 Aoudads were estimated to be present in Aïr Ténéré National N. R. in 1989 (making this the most important stronghold for wild Aoudad on the continent), and 670 outside the limits of the reserve (Poilecot 1991). Numbers are suspected to be increasing in protected areas in the Aïr mountains, but declining elsewhere in Niger (Magin & Newby 1997). Aoudads seem to be locally numerous in the Eastern and Western Deserts of Egypt (M. A. Saleh pers. comm.), where they were once thought extinct. There are no estimates for Lybia (Shackleton & De Smet 1997) or Tunisia (De Smet 1997a); there also are no estimates for Sudan, where they are very rare and declining (Nimir 1997). Small groups scattered irregularly over large ranges are the typical pattern of the distribution of the species in the wild, so that obtaining population estimates in the field is difficult. Freeranging populations in USA and Spain vary in group composition and density both monthly and yearly (Gray & Simpson 1982a, Cassinello 2000). Adaptations This is a mountain-dwelling caprine perfectly adapted to inhabiting rocky and steep terrains.There is no convincing explanation for the existence of its conspicuous mane, but given that it is a sexually dimorphic character, the most likely explanation is that the mane is used for display purposes by !! and it is possible that it may act as a dispenser of scent from squirts of urine. Ogren (1965) postulates that it might provide protection against sand storms, by covering the face; it would also prevent flies from disturbing the animal while foraging. Finally, the mane might also contribute to display in agonistic and sexual contexts; captive adult !! have been seen waving their manes by twisting their heads, raised on their hindlegs while laying the forelegs on fences confronted with other all-male enclosures (T. Abáigar pers. comm.). They use their horns as shovels when sand bathing, a common behaviour probably related to protection against ectoparasites or, as Ogren (1965) suggests, to homeothermy. Foraging and Food Little data on food habits have been reported from the wild (see Poilecot 1991, Chaveas 2000). This species is a generalist herbivore combining grazing with browsing. In Spain their diet comprises all kind of shrubs, succulent and non-succulent forbs, creepers, dwarf shrubs and grasses, depending on seasonal availability (Miranda et al. 2012). In the Aïr Massif, and probably elsewhere, the green seed pods of acacia trees (Acacia tortilis raddiana, A. ehrenbergiana) are much sought after by Aoudads; they are even reported butting tree trunks to make the pods fall (J. Newby pers. 597

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comm.). Aoudads can survive without drinking water for long periods, eating just succulent forbs, but if they have the chance they will drink daily from ponds or wells. They probably make small migratory movements in relation to food availability, but there is no specific information. Social and Reproductive Behaviour The Aoudad is a gregarious species. There is limited information from the wild, but their behaviour is well documented in exotic, free-ranging herds in the USA and Spain. Gray & Simpson (1982a) found that "" lead the group when adults of both sexes are present, while group composition and group size vary seasonally. Six group types can be distinguished in free-ranging populations: solitary; nursery (females, young and juveniles); mixed; all-male; all-female; and alljuvenile (Gray & Simpson 1982a, Poilecot 1991, Cassinello 2000, Chaveas 2000). Mixed groups are mostly found during rutting, and nursery groups during spring and summer. Adult "" and juveniles are the sex–age classes most commonly observed.Yearly fluctuations in group dynamics in the same areas are evident, but generally are characterized by a prevalence of adult "", followed by subadults and finally a lower percentage of adult !! (Cassinello 2000). However, the percentage of adult !! observed in the Tirghist Reserve in Morocco is slightly higher than that of subadults (Chaveas 2000). From the meagre information available (e.g. Dupuy 1966, Chaveas 2000), group size in populations in the wild resembles that in free-ranging populations in USA and Spain. In their Spanish range, herds tend to be small, the majority of groups consisting of less than 11 individuals (Bigalke 1986, Cassinello 2000). In the Texan population, mean group size can oscillate from 5 to 20 individuals (Gray & Simpson 1982a). The mean home-range size ranges from 259 to 3367 ha, and dispersal is particularly accentuated in summer (Simpson et al. 1978, Dickinson & Simpson 1980). The species is not territorial, and !! that reach adulthood leave maternal herds to create their own all-male group or to join existing ones. Social rank determines each individual’s position in a hierarchical order. In captivity, the rank of an adult " may vary according to proximal factors, such as mating, parturition and weaning of young (Cassinello 1995). Aoudad fighting techniques present a variety of forms (Katz 1949, Haas 1959, Ogren 1965, J. Cassinello pers. obs.), some of them ancestral in Caprini, from reverse-parallel pushing to horn-locking; although horn clashing is also present, they do not rise on their hindlegs (as goats do) and they usually clash from short distance, so that collision is not as violent as in wild sheep. Typically, the contestants approach with heads lowered in a threat posture, and when closer they bring the heads down even further, thus directing the bases of the horns forward. Contestants then run toward each other and collide, the blow being delivered with the base of the horns, the force of the collision producing a loud sound that can be heard as far as 300 m away (Habibi 1987). Even infants display all fighting forms when playing (Haas 1959, J. Cassinello pers. obs.). Courtship is carried out by the most dominant !! and follows a conspicuous and ritualized display, similar to that of other caprines, with low-stretch posture, tongue-licking and female chasing; also, a sort of guarding behaviour has been observed in dominant !! following copulation with a " (see Habibi 1987). Females in good

condition bias their investment towards their sons, allocating their resources preferentially (the suckling rate of sons is significantly higher than daughters) and producing more sons than daughters (Cassinello 1996, Cassinello & Gomendio 1996). Mother–offspring conflict during weaning has been investigated. There seems to be a progressive cessation of investment, with no behavioural conflict arising between mother and offspring, except when she resumes sexual activity before weaning has taken place (Cassinello 1997a). Sporadic allosuckling events have been reported in captivity, carried out by alien young of a similar age to that of the allo-mother’s true offspring (Cassinello 1999). Aoudads may produce up to six types of vocalizations: sheep and goat-like bleats (the only sound youngsters produce), short nose snorts, mouth blows, rasping screeches, and grunts (see Gray & Simpson 1982b). Reproduction and Population Structure Males and "" can be regarded as sexually mature when aged 14 months and 9 months, respectively (Cassinello 1997b). The mating season peaks from Sep to Nov, and the average gestation period is 22 weeks (Gray & Simpson 1980). In captivity, twins may be produced as frequently as once every four or five births; triplets have been reported in freeranging populations (ARMAN 1991). Young weigh about 4.5 kg at birth. Singletons are heavier than twins, and high-ranking mothers tend to produce heavier young than low-ranking mothers. Birthweights also increase with maternal age (see Cassinello 1997b, Cassinello & Alados 1996, Cassinello & Gomendio 1996). Interbirth intervals average 10 months in captivity, and weaning takes place at a mean age of 8.2 months. The population sex ratio seems to be 1 : 1. Studies in captivity have shown that variation in female reproductive success is determined first by longevity, followed by fecundity, offspring survival at one month and age at first parturition. Among the factors that affect the components of reproductive success, longevity depends on physical condition, fecundity is higher in top-ranking "", offspring survival increases with birth-weight, and age at first parturition increases with population density and inbreeding coefficients (Cassinello & Alados 1996). Aoudads may live up to 20 years in captivity (Ogren 1965, Weigl 2005), but longevity probably rarely exceeds 10 years in the wild (Gray & Simpson 1980). Predators, Parasites and Diseases Presumably, Lions Panthera leo and Leopards Panthera pardus were among the natural predators of the species in the past, but today Striped Hyaenas Hyaena hyaena, Golden Jackals Canis aureus and feral domestic dogs may be the most common natural predators. Also, raptors may predate newborns and young, as reported in Aoudad populations in Spain (accounts from forest rangers of the Golden Eagle Aquila chrysaetos taking young). The free-ranging populations in S USA seem to be remarkably free of disease and parasites (Ogren 1965, Pence 1980). Conservation IUCN Category:Vulnerable C1. CITES: Appendix II. The lack of surveys carried out in North Africa make determining the precise current status of the species difficult. In its native habitat,

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the major threats are habitat loss, competition with livestock and poaching (Alados & Shackleton 1997), resulting in a steady decrease in numbers throughout its range and increasing isolation of populations. Some populations or subspecies are at serious risk of extinction, and the confirmation of the presence of A. l. ornata in Egypt (Wacher et al. 2002; M.A. Saleh pers. comm.) lends support to the dire need for population surveys in other areas, and clarification on the taxonomic status of the recognized subspecies. The degree to which Aoudads are legally protected across their range varies. For example, Auodads have been fully protected in both Morocco and Tunisia since 1966, while in Niger hunting has been banned since 1964; on the other hand, they receive no formal protection in Mali, and although they are listed under Schedule II as a protected species in Sudan, they can still be shot by anyone with the appropriate licence. Aoudads occur in a number of protected areas, including: Takherkhort Hunting Reserve (established in 1967 specifically to conserve this species), the adjacent Toubkal N. P., and Eastern High Atlas N. P. (Morocco); Belezma, Tassili n’Ajjer, and Ahaggar National Parks, and in Djebel Aissa State Forest (Algeria); Fada-Archei Faunal Reserve (Chad); Adrar Mouflon Partial Faunal Reserve (Mauritania); and the vast Aïr Ténéré National N. R. (Niger) (see Shackleton 1997 and chapters therein). The population in Djebel Chambi N. P. in Tunisia was reintroduced in 1987; some animals escaped and a wild population has survived (De Smet 1997; K. de Smet pers. comm.). A few Aoudads are held in captivity in Djebel Bou-Hedma N. P. (De Smet 1997a), and the species was also released in Oued Dekouk N. R. (K. De Smet pers. comm.). Aoudads were introduced into Tripoli N. R. in Libya (Shackleton & De Smet 1997). In contrast to the situation in the wild, introduced US and Spanish populations are either well established or steadily

increasing (Gray 1985, Cassinello et al. 2004). However, it is a priority to register and keep information on subspecific origins of captive and introduced populations and prevent undesirable hybridization events, particularly if reintroduction programmes are to be implemented. Measurements Ammotragus lervia HB (!!): 1490 (1050–1760) mm, n = 18 HB (""): 1290 (1040–1500) mm, n = 41 T (!!): 190 (170–210) mm, n = 3 T (""): 170 (160–190) mm, n = 4 Sh. ht (!!): 960 (890–1050) mm, n = 3 Sh. ht (""): 780 (760–800) mm, n = 4 WT (!!): 86.0 (50.0–132.0) kg, n = 17 WT (""): 42.0 (12.0–68.0) kg, n = 39 Free-ranging population, New Mexico, USA (Ogren 1965), except HB and WT, which correspond with sexually mature individuals (!! older than 1.5 yr; "" >1 yr) from the captive population at EEZA (CSIC), Almería, Spain (Cassinello 1997b) Maximum recorded horn length is 88.0 cm for a pair of horns from the Ennedi Mts, Chad; however, record horn length from an extralimital population introduced in Colorado, USA, is 90.1 cm (Rowland Ward) Key References Cassinello 1998, 2000; Cassinello et al. 2004, 2006; Chaveas 2000; Gray & Simpson 1980, 1982a, b; Ogren 1965; Shackleton 1997 (and chapters therein). Jorge Cassinello

GENUS Capra Ibexes Capra Linnaeus, 1758. Syst. Nat., 10th edn, 1: 68.

Capra is a polytypic genus that includes eight species: Markhor C. falconeri, Goat C. hircus (wild populations commonly called C. aegagrus) and the ibexes (Nubian Ibex C. nubiana, Walia Ibex C. walie, Siberian Ibex C. sibirica, Alpine Ibex C. ibex, Spanish Ibex C. pyrenaica and Tur C. caucasica) (Grubb 2005). There is no agreement on the true number of species recognized, and as few as two (hircus and falconeri; Haltenorth 1963) and as many as nine have been recognized (the Tur sometimes being split into two species, C. caucasica and C. cylindricornis) (Heptner et al. 1961). The genus is distributed from the Iberian Peninsula, the Alps and the Caucasus Mts through the Middle East, south into Egypt, Sudan, Ethiopia and the Arabian Peninsula and east across Iran, Afghanistan and Pakistan into the Hindu Kush, Pamirs, Tien Shan and Altai Mts of central Asia, in upland precipitous habitats. There is a substantial Pleistocene fossil record in Europe; only the supposed Pliocene C. primaeva (doubtfully in this genus) is recorded from Africa. Only two species, the Walia Ibex and Nubian Ibex, occur in Africa: the former is found only in and around Simien Mountains N. P. in Ethiopia, while the latter is found in Egypt, east of the Nile, NE Sudan, N Eritrea,

Israel, Jordan, SE Oman, Saudi Arabia and SE Yemen (and extinct in Lebanon and Syria). Analysis of mitochondrial cytochrome b and Y chromosome DNA sequences suggests that the two species form a monophyletic clade, although the Walia Ibex potentially has been isolated for up to 0.8 million years from Nubian Ibex (Gebremedhin et al. 2009). Members of the genus are robust animals with stout limbs, short hoofs and short tail. The rhinarium is hairy, and there is a beard on the chin. Long hair extends over forequarters in the Markhor, but not in other species. Males and "" are strongly sexually dimorphic; !! are larger with large horns, "" with relatively small, slender horns. Horns grow during life but growth is seasonal, forming rings or checks during the quiescent phase, which aids ageing individuals in the field. There is a wide range in size between species and sexes (50–130 kg). The skull is characterized by a short narrow facial region, but is broad across the frontals, with tubular orbits; the skull is strengthened by secondary bony deposition leading to disappearance of the nasofrontal suture, though the naso-maxillary suture remains open. 599

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Species are differentiated by size, form of the horns in !! (much variation between species), length of pelage and colour pattern. Along the anterior surfaces of the horns, ibexes have prominent bosses (secondarily lost in some populations of Tur and Spanish Ibex). African ibexes (Nubian and Walia) have contrasting blackand-white leg markings and horns that are long and semi-circular in outline, placed further forward on the skull compared with other ibexes, associated with the development of a swelling on the frontal bones just below the horn bases. They differ from each other in size and horn features (slender or robust, strongly or slightly laterally compressed). Ansell (1972) included the Barbary Sheep or Aoudad Ammotragus lervia in this genus, but this has not been followed here and is in conflict with molecular data (Ludwig & Fischer 1988, Ropiquet & Hassanin 2005a). Peter Grubb & Michael Hoffmann Nubian Ibex Capra nubiana male with tightly curved horns.

Capra nubiana NUBIAN IBEX Fr. Bouquetin de Nubie; Ger. Nubischer Steinbock Capra nubiana (F. Cuvier, 1825). In: E. Geoffroy Saint-Hilaire and F. Cuvier, Hist. Nat. Mammifères 6, pt 50, ‘Bouc sauvage de la Haute-Egypte’, p. 2, pl. 397. Egypt, ‘de la Haute-Égypte … ou de Nubie’; Nubia (Lydekker 1913: 153; G. M. Allen 1939: 549) or Upper Egypt (Ellerman & Morrison-Scott 1951: 407), which are virtually synonymous; restricted by Grubb (2005) to Sudan, Northern Prov., Nubian Desert, east of Nile R.

Taxonomy Monotypic. Considered a subspecies of Capra ibex (Ellerman & Morrison-Scott 1951, Ansell 1972, Corbet 1978, Harrison & Bates 1991), but treated as distinct by Uerpmann (1987) and Grubb (1993c, 2005). Synonyms: arabica, beden, mengesi, sinaitica, typical. Chromosome number: 2n = 60 (Schaller 1977).

Nubian Ibex Capra nubiana.

Description Tan to greyish, mid-sized ungulate of mountainous desert terrain, with prominent horns and striking black and white leg markings. Mature !! much larger and heavier than "", with dark beards and scimitar-shaped horns. During the rut, breeding !! develop enlarged necks and dark pelage on chest, shoulders and flanks. Both sexes have whitish belly, inner legs and buttocks, and short dark tails. Nostril, mouth and orbits outlined in black. Eyes have amber irises and horizontal black pupils. Chin and hairs surrounding mouth and nostrils are usually whitish. Darker band extends from eye to edge of mouth. Nasal area may be rufous-brown. Ear backs grey, and fronts have black centres with contrasting white edges. Summer body pelage of adults and young comprises short, tan guard hairs (replaced in winter by brown to grey guard hairs with underfur). Mid-dorsal band of longer hairs extends from nape to base of tail, and is dark in !!. Belly whitish, partially edged with a dark band in many adults. Fronts of upper and lower foreleg black, with white band above grey knee callus. White fetlocks and pasterns contrast with black hooves on fore- and hindlegs. Dark band above fetlock extends to leading edge of hindleg. The flattened face of the male horns has as many as 30 prominent horizontal knobs; female horns are short and slender and bear narrow growth rings, but no knobs. Male horn dimensions vary among populations (Habibi 1994).

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Capra nubiana

Geographic Variation None recorded. Similar Species Capra walie. Restricted to the Simien Mts in Ethiopia. Larger and stouter body, more robust horns and darker pelage. Distribution Once widely distributed in north-east Africa, east of the Nile R., Sinai, the Near East and the Arabian Peninsula, but decimated by relentless hunting and habitat encroachment in much of their original range (see, for example, Manlius 2001). Nubian Ibexes in Africa exist mainly as small isolated populations in the Eastern Desert and Red Sea Mts of Egypt, and in the Red Sea hills of Sudan. Locations of confirmed recent occurrences in Egypt’s Eastern Desert include: Gebel Ataqa, Gebel El Galâla El Baharîya (North Galala Plateau), and Gebel El Galâla El Quiblîa (South Galala Plateau) (M. Saleh pers. comm.). Nubian Ibexes also occur throughout the Red Sea Mts of Egypt, from Gebel Ghârib in the north to Elba near the Sudan border. Krausman & Shaw (1986) rediscovered a population surviving in the Wadi el Assuti and Wadi Habib, where ibexes had not been seen since 1927. Recent records in Egypt’s Sinai Peninsula are from Gebel El Maghara, Gebel El Halal and Gebel Yalag in N Sinai, the Gebel El Tîh in C Sinai and at many sites in the mountains of S Sinai (Saleh & Basuony 1998). Manlius (2001) provides a detailed discussion of the historical ecology and biogeography of the Nubian Ibex in Egypt from 1800 to the present day. In Sudan, small Nubian Ibex populations were observed around Erkowit, Jebel Ashat and Jebel Sherik Gebel during a 1990 survey (Nimir 1997), and the species likely occurs elsewhere in the country’s Red Sea hills as far north as the Gebel Elba (Nimir 1997, I.M. Hashim pers. comm.). No confirmed recent records are available for Eritrea, where the species had been recorded near the Sudan border (Hillman &Yohannes 1997, D. Zinner pers. comm.). Nubian Ibexes had been reported in

Lateral, palatal and dorsal views of skull of female Nubian Ibex Capra nubiana.

extreme N Ethiopia (Yalden et al. 1984), but the only recent record is an unconfirmed report from the Tendaho Estates area in EC Ethiopia near the Djibouti border (Hillman et al. 1997). Extralimital to Africa, Nubian Ibexes occur in Israel, Jordan, Oman, Yemen and Saudi Arabia, but are now extinct in Lebanon and Syria (Shackleton 1997, and chapters therein; Grubb 2005). Habitat Mountains, cliffs, hills and associated plateaux, canyons and wadis. Supports arid and semi-arid vegetation, including annual forbs and grasses dependent on seasonal rainfall, geophytes, woody shrubs and stands of Acacia, Populus and Pistacia trees. Dense herbaceous vegetation at runoff sites and springs are important foraging habitat. Steep slopes provide vital escape cover, especially for "" and young (Kohlmann et al. 1996). Springs or other open freshwater are essential. Human pressures force Nubian Ibexes to remote and inaccessible locations, and livestock grazing degrades foraging and watering sites (Alados & Shackleton 1997). Abundance Recent African records are from opportunistic observations by biologists (e.g. Krausman & Shaw 1986) and reports by local inhabitants. Three populations in Sinai comprised 300–400 animals in the 1970s (Baharav & Meiboom 1981). About 200 Nubian Ibexes inhabit the Gebel El Galâla El Baharîya Plateau (M. Saleh pers. comm.), and small groups are reported elsewhere in E Egypt (Amer 1997). Numbers in Egypt appear stable (M. Saleh pers. comm.). Isolated bands exist in the Red Sea hills of Sudan, but populations may be declining (Nimir 1997, I.M. Hashim pers. comm.). Adaptations A stocky body, short legs and exceptional equilibrium enable Nubian Ibexes to negotiate rugged terrain. Wide 601

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hooves with large flexible pads may promote stability on narrow ledges and absorb shock when they jump. They respond to threats by rapidly scaling or descending steep slopes, and routinely use narrow, precarious trails. They have excellent distance vision, and group living enhances vigilance (D. Saltz pers. comm.). Their pelage is very cryptic in most desert habitats. Nubian Ibexes lack strong physiological adaptations for water conservation or thermoregulation. They obtain preformed water from plants, but require non-saline drinking water (Shkolnik et al. 1979). During hot summers, Nubian Ibexes are active during early morning and dusk. They avoid activity, and seek shade, during mid-day (Baharav & Meiboom 1982), and may also forage at night. In cool winters they shift their daily activity to sunlit slopes, and are active longer during the day. Nubian Ibexes have a low basal metabolic rate, perhaps reducing their energy requirements in arid environments (Chosniak et al. 1984). Foraging and Food Nubian Ibexes consume a wide array of herbaceous and woody plants. Specific diet varies by locality and season. Woody browse consumed in Egypt included Acacia raddiana, Lindenbergia sinaica, Lycium shawii, Capparis spinosa and Ficus pseudosycomorous (Osborn & Helmy 1980). Herbaceous species eaten were Phragmites australis, Imperata cylindrica, Juncus rigida, Alhagi manifera and excavated roots of Lotus arabicus. In Sinai, important perennial plant foods in winter were: Globularia arabica, Helianthum lipii, Ephedra foliata, Zilla spinosa, Thymus decussatus, Gymnocarpus decander and Echinops glaberrimus (Baharav & Meiboom 1982). Protected Nubian Ibex populations have eaten landscape plantings and agricultural crops in Israel (Hakham & Ritte 1993). In the Negev Desert, mature !! relied on low-quality fibrous browse during most of the year (Gross 1991). They took large bites, and had high intake rates and long rumination bouts (Gross et al. 1995b, 1996, S. Kohlmann pers. comm.). By contrast, the smaller "" and young !! sought higher quality, but scarcer, herbaceous forage throughout the year. Smaller animals chewed food thoroughly, and spent more time actively foraging than did the large !!. Social and Reproductive Behaviour Nubian Ibex are highly social (Habibi 1994, Gross et al. 1995a), typically occurring in female-based bands that include young and male juveniles to three years of age. Mature !! more than seven years of age represent a second group type, while !! 4–6 years old may form more transitory associations of six or more individuals. In a protected Negev population, adult "" maintained a stable linear dominance hierarchy throughout the year (Greenberg-Cohen et al. 1994), and female-based groups averaged more then 20 individuals (Gross et al. 1995a). Nubian Ibex bands are significantly smaller in the face of intense hunting pressure and habitat degradation. In hunted Egyptian and Saudi Arabian populations, groups averaged less than six animals (Osborn & Helmy 1980, Habibi & Grainger 1990). Mature !! disperse in late summer, and may travel long distances to join female groups during the 6–8 week autumn rut. A travel corridor between Israel populations used by breeding !! was ≥60 km long (Shkedy & Saltz 2000). They court "" using characteristic approach behaviour, with neck outstretched, tail-up and displaying flehmen (Habibi 1994). Unreceptive "" ignore, threaten or flee persistent suitors.Younger !! also court "", but are easily displaced by mature !!. Closely-matched breeding !!

engage in dramatic horn clashes in which winning !! often attain higher elevations to increase the downward force of their attack (S. E. Kohlmann pers. comm.). Neonates remain hidden for about a week after birth, and then follow their mothers who rejoin female-based groups. Kids are adept climbers and gambol and play in precarious sites. In protected populations, Nubian Ibexes have formed temporary nurseries of many young associated with only few "" (Hakham 1985, Levy & Bernadsky 1991).Young have been confined for several days or more in a natural topographical ‘trap’ in which they were visited by their mothers until they managed to escape (Müller et al. 1995). Reproduction and Population Structure Females breed by two years of age. Following a 20-week gestation period, they give birth in spring (Mar–Apr) to one or two offspring at secluded sites in rough terrain. Nubian Ibexes are short-lived, rarely exceeding 12 years of age in the wild, even in protected populations (P. Alkon pers. obs.). Sex ratios at birth are even, but adult "" usually outnumber !! (Habibi 1994). Annual fluctuations in protected populations are probably regulated by yearly changes in environmental conditions as well as density-dependent mechanisms (Alkon & Kohlmann 1990). The dispersion of breeding !! among distant female-based groups during the rut (Shkedy & Saltz 2000) may promote high genetic diversity in protected populations (Stuwe et al. 1992). Predators, Parasites and Diseases On the African continent the only likely predator is the Leopard Panthera pardus, although, on the Arabian Peninsula, Grey Wolves Canis lupus may occasionally prey on them (Harrison & Bates 1991, G. Ilani pers. comm.). Attacks by large raptors are rare. Parasites of wild Nubian Ibexes include nematode lungworms, coccidia and arthropods (Solomon et al. 1996, Yeruham et al. 1999). Captive animals have succumbed to sarcoptic mange (Yeruham et al. 1996). Conservation IUCN Category: Vulnerable C1+2a(i). CITES: Appendix II. The main threats to the Nubian Ibex include competition with livestock and camels in Egypt, where the availability and distribution of waterholes is likely to be a major factor in the condition of populations (Amer 1997). Hunting is also a threat in Egypt and Sudan (Amer 1997, Nimir 1997), and is implicated in the disappearance of this species from some parts of its range in Egypt (Manlius 2001). Egypt and Sudan have enacted protective wildlife laws and established wildlife reserves, but resources for surveys and enforcement are limited. Armed conflicts also have disrupted conservation measures in parts of north-east Africa. In Egypt, Nubian Ibexes are legally protected and their hunting is totally forbidden under articles of the Agricultural Law. Nature Reserves in which they occur are Gebel Elba Conservation Area (disputed Sudan Government Administration area) and the Wadi el Assuti in the Eastern Desert, and Gebal Mûsa and Gabel Katerîna Wildlife Reserve in south central Sinai (Amer 1997). Nubian Ibexes appear to have maintained their number and range in Egypt over the past several decades (M. Saleh pers. comm.). In Sudan, Nubian Ibexes are a Schedule II species under the Wildlife Conservation Act. Hunting was banned between 1989 and 1992, since when the ban has been lifted and animals can be hunted under special permit (Nimir 1997).

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Protected areas in which they occur include Tokar G. R., SinkatErkawit Game Sanctuary and Erkawit Game Sanctuary, totalling 829,500 ha. No hunting of wildlife has been recently permitted in Eritrea. The Yob Wildlife Reserve (2658 km2) along the Sudanese border encompasses past Nubian Ibex range in that country (Hillman & Yohannes 1997). Future conservation measures should include placing the Nubian Ibex on Schedule I of theWildlife Conservation Act in Sudan to ensure full protection (Nimir 1997), government control of the area that includes the Wadi el Assuti and its junction with the Wadi Habib (one of the few places outside the Sinai where good numbers of Nubian Ibexes survive) and making Gebel Elba a National Park (requiring cooperation between Egyptian and Sudanese governments) (Amer 1997, Nimir 1997). Nubian Ibexes respond strongly to effective protection, which bodes well for future conservation measures (Habibi 1994, 1997, Alkon 1997, M. Saleh pers. comm.). Within a generation, newly protected Nubian Ibexes reduced flight distances from humans, returned to abandoned range and formed larger groups. The largest extant Nubian Ibex populations (≥1600 animals) are in Israel, where they are very tolerant of humans following four decades of protection.

Measurements Capra nubiana HB (!!): 1370 (1190–1590) mm, n = 20 HB (""): 1092 (920–1210) mm, n = 25 T (!!): 116 (80–170) mm, n = 20 T (""): 91 (65–160) mm, n = 27 HF c.u. (!!): 290 (260–390) mm, n = 13 HF c.u. (""): 272 (240–300) mm, n = 20 E (!!): 168 (135–200) mm, n = 20 E (""): 153 (130–175) mm, n = 28 WT (!!): 64.2 (42.0–85.0) kg, n = 16 WT (""): 31.4 (25.0–39.5) kg, n = 30 Negev Desert, Israel (P. U. Alkon et al. unpubl.); ages of all !! ≥4 years, and all "" ≥3 years Maximum recorded horn length is 138.4 cm for an animal from Khartoum Zoo in Sudan (Rowland Ward) Key References Gross 1991; Habibi 1994; Harrison & Bates 1991; Manlius 2001; Osborn & Helmy 1980; Shackleton 1997 (and chapters therein). Philip U. Alkon

Capra walie WALIA IBEX (ETHIOPIAN IBEX) Fr. Bouquetin d’Ethiopie (Le Wali); Ger. Aethiopischer Steinbock (Walia-Steinbock) Capra walie Rüppell, 1835. Neue Wirbelt. Fauna Abyssin. Gehörig, Säugeth. 1: 16; 40 pp, 14 pl. ‘die höchsten felsigten Gebirge Abyssiniens … in den Provinzen Simen und Godjam’; restricted to Ethiopia, mountains of Simien (Lydekker 1913: 156).

valie, wali. Chromosome number: not known, but probably 2n = 60, as for other karyotyped Capra species.

Walia Ibex Capra walie.

Taxonomy Monotypic. Originally named as a species (Rüppell 1835, Lydekker 1913, Harper 1945), it has been considered a subspecies of Capra ibex (Haltenorth 1963, Schaller 1977) and of Capra nubiana (Heptner et al. 1961). Now usually treated as a distinct species (Ansell 1972, Yalden & Largen 1992, Grubb 1993c, 2005), which appears to be supported by genetic data that reveal that the differentiation between C. walie and C. nubiana is similar to the differentiation observed between the Alpine Ibex C. ibex and the Spanish Ibex C. pyrenaica (Gebremedhin et al. 2009). Synonyms: vali,

Description Mid-sized ungulate of robust build with outstanding rock-climbing abilities. Less massive and sleeker than the Alpine Capra ibex and the Siberian Ibex Capra sibirica, but stouter and heavier than its more elegant northern neighbour, the Nubian Ibex Capra nubiana. Back and upper body chestnut-brown, the chin, throat, belly and the inner surfaces of the legs whitish for both sexes. A black stripe extends down the front of each leg, a white band just above the knee cuts across the black stripe on the forelegs. This black and white pattern is less pronounced in young !! and "" than it is in !! aged three years or older. Males aged 4–6 years have small beards, which turn into long beards from about the seventh year, at which age their chest darkens and the black streak on their back becomes more pronounced. Females and young !! do not have beards (Nievergelt 1981, Nievergelt et al. 1998). Adult !! are larger than "" and have semicircular, massive horns with transverse knobs on the front. Females have small and slightly curved horns. Geographic Variation

None recorded.

Similar Species Capra nubiana. Occurs in Egypt, Sudan and W Eritrea, extending into the Near East and the Arabian Peninsula. Slender and smaller, dominant colour of coat bright and sandy, and horns of !! thinner and more curved. 603

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The steep and rocky areas in the alpine and ericaceous belts along the escarpment as well as the mountain forest belt are the main habitat of the Walia Ibex.War in this region gave rise to indiscriminate hunting mainly in the western parts of the National Park, and the Walias adapted to living and ranging in adjoining habitats as well. There is reasonable evidence to suggest that Walias not only found refuge among rocks and in the denser mountain forest in lower altitudes, but that most animals of the small population moved into the escarpment towards the slightly higher and less disturbed eastern region (although it lies outside the National Park). The areas south of Bwahit Mountain were the main retreat during this critical period. They included parts of the escarpment, but also cliffs, steep slopes with almost no forests and an extensive high plateau, where the alpine steppe is so scarce that there is almost no competition from livestock. The use of flat, plateau-like areas as feeding grounds is rather untypical for most ibexes, but its use by the Nubian Ibex has been observed. This habitat preference of the Walias in the Bwahit area indicates that rich alpine steppe on the slightly lower plateaux within the National Park are suitable Walia Ibex habitat, too, provided there is no disturbance by man or livestock (Nievergelt 2012). Capra walie

Distribution Endemic to Ethiopia. The Walia Ibex is found only in and around Simien Mountains N. P. in the North Gonder Administrative Zone of NW Ethiopia. As far as historical records go, its range seems to have always been very restricted and the total population remains small and vulnerable (Nievergelt 1981, Yalden & Largen 1992, Hillman et al. 1997). Considering available records and looking at the topographic and climatic features of the heterogeneous and often fractured country, it may be assumed that the extremely deep and arid lowland-channel with the Tekeze R. is a natural border in the north and east. In the north-west, west and south, Walias probably occurred in the spurs of high Simien, ranging from Adi Arkay towards Debark and perhaps Dabat. Habitat The Simien Mts are remarkable for the impressive escarpments that separate the hilly high plateaux from the terraced lowlands, the original flora of which is a dense and diverse mountain forest with species such as Juniperus procera, Olea africana, Syzygium guineense up to roughly 3000–3200 m. An ericaceous belt dominated by the tree heather Erica arborea and St John’s Wort Hypericum revolutum occurs at 3200–3800 m, and the whole of the vegetation above the timberline consists of afroalpine grassland with the megaphytic Lobelia rhynchopetalum. Inatye Mountain within the National Park reaches 4070 m, and the adjoining mountains Bwahit and Ras Dejen are 4430 and 4543 m, respectively. Seasonal changes in vegetation are determined by the wet season, from Jun until Sep/Oct, but variations in diurnal temperature exceed by far the variations entailed by seasonal changes. Today, farmers have turned a large part of the original vegetation of the lowlands into terraced fields (Klötzli 1977, Nievergelt 1981, Hurni 1982, 1986, Nievergelt et al. 1998, Hurni & Ludi 2000) and livestock have also encroached into the forest. The higher escarpments and plateaux are intensively used as pasture for cows, sheep, goats and horses but they are too cold for agriculture.

Abundance The population size of Walia Ibex numbered 150–300 animals between 1966 and 1969, increasing slowly until 1983 when there were possibly more than 500 animals, and then decreasing again during the period of civil unrest during the early 1990s. In 1994 and 1996, the population was estimated at 200– 250 animals, but it subsequently increased, reaching about 500 animals in 2004 (Nievergelt 2012). The earlier recovery of the population after 1971 can be regarded as a positive response to the introduction of an effective control system with several outposts of game wardens. Along with this recovery, the animals slowly became less shy, and were even seen in habitats close to footpaths. In addition, a notably higher proportion of older !! was observed. After the war the animals became extremely shy again, hiding in naturally well-protected places (Hillman et al. 1997, Nievergelt et al. 1998). The more recent recovery took place exclusively in the eastern corner of the National Park, the Chennek area, and was even more pronounced in the adjoining Bwahit–Mesarerya– Digowa range outside the National Park (Ludi 2005, 2006) where, in the 1960s and 70s, no Walias could be observed at all. The most recent estimate (2009) puts the total population at ca. 750 animals (Alemayeh et al. 2011). Adaptations As with all members of the genus Capra, the Walia Ibex is an outstanding rock-climber with strong and relatively short legs; it is more sure-footed than almost any other African ungulate, and moves safely up and down sheer cliffs. The "" give birth in extremely inaccessible places and the young learn to climb immediately. The characteristic afroalpine climate requires adaptation to cold, but this regime is also the basis of rich pastures and – between roughly 2300 and 2700 m – the dense and protective evergreen montane forest. The diurnal activity pattern shows a peak of foraging and social activities in the morning and again towards the evening. Foraging and Food The diet of the Walia Ibex includes a great variety of herbaceous and woody plants (Dunbar 1978, Nievergelt

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1981). Among the abundant trees and shrubs dietary staples include Erica arborea (twigs) and Lobelia rhynchopetalum, but they seem to be partial to a number of green herbs, such as Simenia acaulis, Alchemilla rothii, Arabis alpina, Swertia sp. and Scabiosa columbaria. There is little evidence of them foraging on members of the family Papilionaceae, and no reports of them foraging on Labiatae. Most herbs are foraged mainly around their respective flowering seasons. According to direct observation, fresh plants were usually selected by !! at the beginning of their flowering period, while "" favoured them at the end of this period when the plants produce highly nutritious fruits. Social and Reproductive Behaviour Like other ibexes,Walias are highly social animals (Dunbar & Dunbar 1974b, 1981, Nievergelt 1974, 1981). They have varied social units, such as female-based groups that include young and juvenile !! until they are about two or three years, groups of adult !! and mixed groups. As can be expected from the low density, most groups are relatively small. In earlier studies, the largest group recorded included 24 animals ("" and young, including !! up to three years); in 2004, in the open area east of the National Park borders, a herd of 54 Walias was observed (Ludi 2006). Sexual segregation is pronounced, mostly during parturition and lactation. In keeping with the rutting peak (see below), the highest occurrence of mixed groups can be observed from Mar to May. The clear rank order amongst !! correlates closely with their age, body size and horn length. The linear dominance hierarchy observed in several Capra species can be expected in Walia "" as well. Open groups and established hierarchies give the social system its structure. Individuals are occasionally involved in fights or play-fights if the sparring partner is of similar age or rank. It is remarkable that quite a number of fights were observed between "" and !! of one or two years, when the !! attain or surpass the body weight of adult "" and start to leave the female-based groups. As has been observed in other ibexes, there seems to be no sign of territoriality. Based on seven individually known animals it is presumed that individuals or small groups usually remain for several months in areas of less than 0.5 km2, but adult !! often range in larger areas. Courting !! lower their horns while approaching receptive "", and display flehmen, lifting their tail into an almost horizontal position. Newborns stand within the first hours after birth, and follow their mother closely. Within the female-based groups, the young remain in close contact with their mothers. Reproduction and Population Structure In contrast to the ibex species living in temperate latitudes, where winter dictates a discrete annual reproductive cycle, rutting behaviour and newborns can be observed throughout the year. However, there is a clear rutting peak between Mar and May, and, in keeping with the gestation period of five months, births peak accordingly after the wet season in Sep/Oct. Older ibexes, both !! and "", have a more synchronized cycle. Although "" could well be both fertile and sexually receptive earlier, "" younger than four years of age are only exceptionally observed with a newborn (Nievergelt 1972). This late breeding corresponds to the slow growing process of this mountain ungulate. Data on life expectancy are lacking. Based on findings concerning other caprines (Geist 1971, Nievergelt 1966),

and the examination of horns, the maximum age in the wild was estimated to be around 15 years. Predators, Parasites and Diseases Leopards Panthera pardus and Spotted Hyaenas Crocuta crocuta are potential resident predators, but there have been no direct reports of attacks by these animals. A number of direct observations indicate that numerous large birds of prey such as Verreaux’s Eagle Aquila verreauxii, Tawny Eagle A. rapax and Martial Eagle Polemaetus bellicosus may take young (Nievergelt 1974). In samples of faeces the following intestinal parasites have been identified: Trichostrongyloides spp., Nematodirus spp., Strongyloides spp., Trichuris spp. and Bunostomum spp. The samples were collected after the reported death of five Walias, with symptoms of diarrhoea, in the eastern part of the range and at the end of the wet season in 2004 (Berhanu Gebre et al. 2004). Conservation IUCN Category: Endangered B1ab(iii), D. CITES: Not listed. Walia Ibexes are protected by Ethiopian law, and cannot be hunted except under a Special Permit for Hunting Game Animals for Scientific Purposes. The species has survived a period of war, and the main current threats to it now are posed by human and livestock pressure on the habitats in and around Simien Mountains N. P. (13,600 ha). The latter was gazetted by the Simen National Park Order No. 59 (1969) and has been a World Heritage Site since 1978 (when most of the people living in the lowland areas where resettled outside). The Park is administered by the Parks and Wildlife Administration Authority of the Regional Government, and has attracted increasing international attention since the World Heritage Committee placed the Park on the List of World Heritage in Danger in 1996 because of decline in the population of the Walia Ibex due to human settlement (most of the people resettled outside of the park in 1978 have returned once again and now reside either within or outside the Park), grazing and cultivation. Walia Ibexes are excluded by cultivation of the lowland terraces right up to the timberline and are further displaced by the practice of allowing domestic animals to graze at the higher altitudes. Despite the existence of national and regional legislation, the remoteness of the area, coupled with the existence of people living within and outside of the Park prior to its establishment as a Conservation Area, makes legislation difficult to enforce. Only the sheer inaccessibility of major parts of its habitat has protected the Walia from total extinction. Proposed conservation measures include: (1) extend the National Park area towards the Bwahit region so as to establish a buffer zone and increase the effective area of the Park with natural movement corridors along the escarpment range (as proposed by Hurni 1986); (2) reduce human and livestock impact in the National Park; (3) prohibit all hunting within the Park and enforce regulations effectively; (4) exclude all possibility of hybridization by prohibiting free-ranging domestic goats from the area; (5) initiate a captive-breeding programme to reduce the risk of extinction (there currently are no Walia Ibexes in captivity anywhere in the world); and (6) establish a monitoring programme that includes systematic assessment of conservation measures (Brown 1969b, Hurni 1986, Hillman et al. 1997, Shackleton 1997, Nievergelt et al. 1998). The Walia Ibex is used by the Ethiopian Wildlife Conservation Organization (EWCO) and the Ethiopian Wildlife and Natural 605

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History Society (EWNHS) as their emblem, and frequently features elsewhere in other Ethiopian symbolism (including the national football team, who are nicknamed ‘The Walya Antelopes’). Measurements No reliable body measurements are available, although Haltenorth & Diller (1980) report measures as: HB: 1500–1700 mm; T: 200– 250 mm; Sh. ht (!!): 1000–1100 mm; Sh. ht (""): 900– 1000 mm; and WT (!!): 100–125 kg; WT (""): 80–100 kg

Maximum recorded horn length is 118.1 cm for a pair of horns from Ethiopia (Rowland Ward) Key References Dunbar & Dunbar 1974b, 1981; Hillman et al. 1997; Nievergelt 1972, 1974, 1981; Nievergelt et al. 1998. Bernhard Nievergelt

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Glossary abbrev. = abbreviation adj. = adjective cf. = confer, compare with; as opposed to Lat. = Latin n. = noun pl. = plural q.v. = quod vide, ‘which see’ v. = verb acetabulum: the concave socket (fossa) in the pelvic bone in which the head of the femur articulates. acrocentric: describes a chromosome that has the centromere (q.v.) very near one end and that therefore appears to have only one arm (= telocentric (q.v.) for practical purposes). ad libitum: (Lat.) as much as one likes; having unrestricted access to a resource (e.g. water or food). aestivate: state of torpor (q.v.) induced by cold or drought; usually associated with a reduced metabolic rate and inactivity. aFN: the total number of chromosomal arms in the autosomal chromosome complement of a species (cf. fundamental number [FN], which includes the chromosomal arms of the sex chromosomes as well as those of the autosomal (q.v.) chromosomes). Each metacentric (q.v.), submetacentric (q.v.) or subtelocentric (q.v.) chromosome is given a value of 2; each acrocentric (q.v.) chromosome is given a value of 1. See also fundamental number. afroalpine: describes habitats and/or vegetation occurring above the treeline on African mountains. Includes montane grassland and heathlands. afromontane: refers to mountainous regions in Africa, e.g. afromontane forests and afromontane grasslands. agouti: having an even mixture of pale- and dark-tipped hairs on the pelage creating a grizzled, speckled or ‘pepper and salt’ appearance. Albertine Rift Valley: see Rift Valley (q.v.). alisphenoid: bone in the skull. Allee effect: a scenario in which populations at low numbers are affected by a positive relationship between population growth rate and density, which increases their likelihood of extinction. allele: an alternative form of a gene. A diploid organism carries two alleles (which may be same or different) for each gene locus. At any one locus, there may be several possible alleles (although only two are present in a single organism). Allelomimetic behaviour: behaviour in social animals in which each animal does the same thing as those nearby. Allen’s Rule: A rule that states that structures in endotherms such as limbs (which are more prone to heat loss) are reduced in size by means of natural selection over time in cooler climates (to reduce heat loss). allogrooming: grooming behaviour directed at another individual. cf. autogroom (q.v.).

allomothering: non-parental mothering; caring for young by individuals (male or female) that are not the parents of the young. allopatry (adj. allopatric): the situation where populations of the same or different species have non-overlapping geographic ranges; refers also to populations of the same, or different, species that are geographically separated. cf. sympatry (q.v.); syntopy (q.v.). allozyme: one of a number of forms of the same enzyme having different electrophoretic properties and that are encoded by alternate alleles at the same genetic locus. altimontane: collective term for the belts of ericaceous and afroalpine vegetation on the high mountains of tropical East Africa (White 1983). altricial: describes young born in an undeveloped state. cf. precocious (q.v.). altruism: behaviour that enhances the reproductive and genetic fitness of another individual at the expense of its own. alveolus (pl. alveoli, adj. alveolar): small cavity; socket that houses the root of a tooth. angular process: process at the posterior lower corner of the mandible; situated ventral to the coronoid process (q.v.). ante-orbital (= anteorbital): in front of the orbit (q.v.). antebrachial: anterior to the arm (forelimb). anterior palatal foramina: the two foramina (q.v.) on the ventral part of the skull. apomorphy (adj. apomorphic): situation in which a novel character evolves from a pre-existing character. In cladistics (q.v.), an apomorphic character shared among two or more species (synapomorphy [q.v]) indicates shared descent from a common ancestor and hence monophyly (q.v.). cf. plesiomorphy (q.v). arboreal: living above the ground (in trees and shrubs). cf. scansorial (q.v.); terrestrial (q.v.). auditory bulla: see tympanic bulla. auditory meatus (pl. auditory meati): the external opening of the ear; the passage leading from the tympanic membrane (ear drum) to the external ear. autapomorphy: derived trait uniquely characteristic of a taxon. autogroom: grooming behaviour in which an individual grooms itself. cf. allogroom (q.v.). autosomal: pertaining to any chromosome other than the sex chromosomes. bachelor herd: a herd comprised entirely of males, usually mature, but of mixed age. baculum (pl. bacula, adj. bacular): the os penis, or penis bone, which supports the penis in some mammals. bai (pl. bais): an opening or clearing. basal metabolic rate: metabolic rate required for survival in the thermal neutral zone (q.v.); a state that requires the lowest expenditure of energy when at rest. basicranium: the base of the skull. 607

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basisphenoid: cranial bone in middle of base of skull; the median posterior part of the sphenoid bone, forming part of the floor of the braincase. Bergmann’s Rule: The theory that the size of a warm-blooded animal in a single, closely related, evolutionary line, increases along a gradient from warm to cold temperatures. bicuspid: having two points or cusps (particularly of teeth). bifid: divided by a shallow notch. bilophodont; describes cheekteeth having two transverse ridges. bipedal: body supported by the two hindlimbs; movement not using the forelimbs. blastula: a hollow ball of undifferentiated cells (derived from a fertilized ovum by cell division), which represents one of the earliest stages of embryonic development. BP: (abbrev.) before the present. brachydont: describes a premolar or molar tooth with low crowns. cf. hypsodont (q.v). braincase (= cranium): that part of the skull housing the brain; the part of the skull posterior to the front line of the orbits. cf. rostrum (q.v.). buccal: On the cheek side of the mouth or teeth or penetrating to the cheek or sometimes used broadly as pertaining to the cavity of the mouth. bulla: see tympanic bulla. bunodont: describes molar teeth, entirely covered by enamel, that have low, rounded, hill-like cusps (as opposed to sharp, pointed cusps). (cf. hypsodont, lophodont). bushmeat: meat for human consumption derived from nondomesticated mammals, birds and reptiles taken from their natural habitats and domiciles. bushveld: savanna vegetation type characterized by a grassy ground layer and a moderately dense upper layer of shrubs and scattered trees. BZ: (abbrev.) Biotic Zone. C or c: (abbrev.) canine tooth; upper case denotes adult dentition, lower case denotes deciduous dentition (milk teeth). See also canine. c.u.: (abbrev.) (Lat. cum unguis = with nail) measurement of the hindfoot when length of the nail on the claw is included in the measurement. Usually hindfoot is measured without the claw because claws may be broken or worn. When length of claw is included, it is conventional to record as ‘HF c.u.’. cf. s.u. (q.v.) caecum (pl. caeca): a blind-ending pouch in the alimentary canal (often enlarged as a fermentation chamber) located at the junction of the small and large intestines. canine: the most anterior tooth on the maxilla bone and in a similar position on the mandible; situated immediately posterior to the incisors; if incisors are absent, the most anterior tooth in the jaw. Unicuspid; tall and pointed in most mammals. Never more than one canine on each side of each upper and lower jaw; absent in some taxa. caniniform: having shape and appearance of a canine tooth. carnassial shear: Found in some carnivores and formed by the blade-like cusps of the fourth upper premolar teeth and first lower molar teeth, which, occluding together like the blades of a pair of scissors, provide a shearing action for cutting through tough skin or bone. Sometimes referred to as the carnassials.

carotid: pertaining to the carotid artery located in the front of the neck though which blood from the heart flows to the brain. cauda epididymides: the ducts of the epididymides at the posterior end of the testes that carry sperm from the testes to the vas deferens, which, in turn, carries sperm to the penis. Sometimes used to store sperm prior to copulation. caudal: pertaining to the tail; in the direction of the tail. Cenozoic (= Cenozoic Era): geological era, ca. 65 mya to today, comprising the Quaternary and Tertiary Periods: the Age of Mammals. central Africa: Cameroon (south of the Sanaga R.), Central African Republic (but only south of ca. 7° N), Equatorial Guinea, Gabon, DR Congo (except SE). Mainly rainforest habitats and rainforest– savanna mosaics. centromere: the part of a chromosome where sister chromatids are linked together during mitosis. cerebellum: the part of the hindbrain that controls and coordinates motor movements, posture, balance and muscle tone. cerebrum (= cerebral hemispheres): the anterior part of the brain that is involved in voluntary movements, processing sensory information, olfaction, learning, memory, communication and other functions. cervical: pertaining to the neck. cf. (in general usage): compare or compare with. In the context of descriptions, implies a difference or contrast: e.g. ‘In Elephatulus edwardii, first lower premolar single-rooted (cf. E. myurus in which the first lower premolar is double-rooted).’ cf. (in taxonomy): precedes the specific name if there is uncertainty in the assignment. cheekteeth: the premolar (q.v.) and molar (q.v.) teeth combined. choana (pl. choanae): the openings of the internal nostrils on the skull, situated immediately posterior to the bony palate. chromosome: one of the thread-like bodies within the nucleus of a cell, which carry the genes (genetic material) in linear order; each chromosome is composed of one long molecule of DNA (and two long molecules at cell division). Chromosomes occur in pairs (one from each parent) and are visible as rod-like bodies in cells that are dividing. The total number of chromosomes in a cell is expressed as the diploid number (2n). CI: condyloincisive length; the length of the skull from the anterior end of the longest incisor tooth to the posterior end of the occipital condyles. cf. GLS. cingulum (pl. cingula): ridge around the base of the crown of a tooth. CITES (abbrev.): Convention on International Trade of Endangered Species of Wild Fauna and Flora; an international treaty set up to ensure that international trade in wild animals and plants does not threaten the survival of species in the wild. It accords varying degrees of protection to more than 33,000 species of animals and plants. Appendix I lists species that are the most endangered among CITES-listed animals and plants. Appendix II lists species that are not necessarily now threatened with extinction but that may become so unless trade is closely controlled. Appendix III is a list of species included at the request of a Party that already regulates trade in the species and that needs the cooperation of other countries to prevent unsustainable or illegal exploitation.

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clade: branch of a phylogenetic tree containing the set of all organisms descended from a common ancestor. cladistic (analysis): a methodology that provides a classification in which organisms are grouped in terms of the time when they had a common ancestor. cline (adj. clinal): in context of geographic variation, a gradual and sequential change of a character(s) without a significant break such as would justify division into separate subspecies or species. CMS: Convention on Migratory Species of Wild Animals (also known as the Bonn Convention). An intergovernmental treaty, concluded under the aegis of the United Nations Environment Programme, concerned with the conservation of migratory terrestrial, aquatic and avian migratory species throughout their range. Migratory species threatened with extinction are listed on Appendix I; Migratory species that need or would significantly benefit from international co-operation are listed in Appendix II. CNL: condylo-nasal length; measurement from the most anterior part of the nasal bone to the most posterior part of the occipital condyle (exoccipital) on the same side of the skull; a similar measurement to ‘greatest length of skull’. cochlea (pl. cochleae, adj. cochlear): a hollow structure, spirally coiled like a snail’s shell, situated in the skull and containing the internal organ of hearing. competitive exclusion: the principle that two different species cannot indefinitely occupy the same ecological niche. concatenation: a chain of linked elements. concave: having a curvature that curves inwards; having an outline or a surface curved like the interior of a circle or sphere. cf. convex (q.v.). concavity: a concave depression in an outline or surface. conceptus: embryo prior to implantation. conductance: in thermal biology, the rate at which heat passes across a temperature gradient, e.g. the density and thickness of the pelage affects the rate at which body heat passes from the body to the outside. Thick pelage, which traps and holds air, results in low thermal conductance. condylar process: process at the posterior upper corner of the mandible, which forms the lower hinge of the jaw articulation; fits into the glenoid fossa (q.v.) of the skull. condylarth (adj): as in the Condylarthra, an extinct order of mammals. condyle: a rounded process on a bone that articulates with a socketlike concavity in another bone. condylobasal length: the length of a skull, measured from the anterior points of the premaxilla (q.v.) to the posterior surfaces of the occipital condyles (q.v.). congeneric: belonging to the same genus. conspecific: belonging the same species. cf. heterospecific (q.v.). contiguous: touching; sharing a boundary (as in geographic ranges). convex: having a curvature that bulges outwards; having an outline or a surface curved like the exterior of a circle or sphere. cf. concave (q.v.). coprophagy: the eating of faeces. Includes the eating of an individual’s own faeces as they are voided from the anus. copulatory plug: plug formed in the vagina of the female after copulation; formed from seminal fluids of the male. Prevents or

reduces the chance of sperm from another male(s) entering the female reproductive tract if the female copulates again soon after copulation with the first male. coronoid canal: a foramen (canal) in the coronoid process (q.v.) of the mandible. coronoid process: angular pointed process on the upper margin of the mandible, situated anteriorly to the condylar process (q.v.); does not participate in the jaw articulation. corpus luteum (pl. corpora lutea): a glandular mass of tissue on the surface of an ovary, that develops after the extrusion of an ovum from a Graafian follicle (q.v.); secretes the hormone progesterone. cotype: originally synonymous with syntype but now used as synonym of paratype (q.v.). CR: (abbrev.) see crown–rump length. cranial profile: the shape of the cranium (the part of the skull that surrounds the brain) when viewed from the side. craniodental: pertaining to the skull and teeth. cranium: that part of the skull housing the brain. Also called braincase. crepuscular: at, active in, twilight, when light intensity is higher than at night but lower than during the day. cf. diurnal (q.v.); nocturnal (q.v.). Cretaceous Period: period (within the Mesozoic Era); 146–65 mya. crown: (1) top of head; (2) exposed part of a tooth (visible above gum), especially the grinding surface. crown–rump length (CR): distance from the crown of head to the rump of a foetus (i.e. maximum length of a foetus in its natural form). cuckolding: when an intruding male mates with an oestrous female without her mate being aware of the event. cursorial: pertaining to running. cusp (adj. cuspidate): a prominence or sharp point, such as on the occlusal surface of some teeth. See also t. cutaneous: (adj.) pertaining to the skin. Cyrenaica: a region of North-East Libya. Includes the Cyrenaican Plateau and that part of the Mediterranean Coastal Biotic Zone between the plateau and the sea, as well as drier terrain south of the plateau. cytochrome b: a protein involved in electron exchange in the mitochondria. It is the product of a gene in the mitochondrial genome. The sequence of this gene is often compared between species in phylogenetic studies to infer relatedness. cytogenetics (adj. cytogenetic): the study of the microscopic structure of chromosomes, especially the mapping of genes. cytonuclear: (adj.) pertaining to the nucleus and the cytoplasm of a cell. Dahomey Gap: the geographic region where savanna habitat extends southwards to the West African coast in E Ghana, Togo, Benin (formerly Dahomey) and extreme SW Nigeria. The presence of savanna forms a break (or gap) in the extensive Rainforest Biotic Zone, which extends along the West Africa coast from Sierra Leone to Cameroon. The Dahomey Gap is an important biogeographical barrier separating the faunas to the east and west of the Gap. deciduous teeth: see milk teeth (q.v.). 609

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Glossary

Dega: Ethiopian word for the temperate agricultural/economic altitudinal zone, about 2300–3000 m, warm enough for cerealbased agriculture. delayed implantation: a means of lengthening the interval between copulation and parturition by delaying the implantation of the blastula (q.v.), so that both copulation and parturition can occur in the most optimal seasons. Development to blastula stage is followed by a period of halted development lasting several weeks or months; then the blastula implants and embryonic development proceeds normally, usually without any further interruption, until the young is born. deme: a unit of population that is interbreeding and is separate from any other such population. dental formula: a simple numerical method of denoting the number of incisor (I), canine (C), premolar (P) and molar (M) teeth on one side of the upper jaw and lower jaw, and the total number of teeth. For example, the dental formula of a primitive mammal is I 3/3, C 1/1, P 4/4, M 3/3 = 44, which means there are three incisors, one canine, four premolars and three molars on each side of the upper jaw and also the lower jaw, making a total of 44 teeth. The formula may also be expressed in the form 3143/3143 = 44. Each incisor, premolar and molar is numbered according to its position in the tooth row; superscript numbers indicate upper jaw, subscript numbers indicate lower jaw (mandible) e.g. P4 (upper fourth premolar), M2 (lower second molar). dentine: the substance, also known as ivory, comprising tusks (q.v.) and the interior hard part of vertebrate teeth. Lies under the enamel of teeth (but may be exposed if the enamel wears) and surrounds the pulp chamber and root canals. diastema: space in the mouth between the incisor teeth and cheekteeth in those mammals that feed on grasses, herbs etc. (e.g. rodents, hares, rabbits, ruminants, etc.). dichromatism: condition in which members of a species show one of only two distinct colours or colour-patterns. dilambdodont: molar tooth with W-shaped ridges. cf. zalambdodont (q.v.). dimorphism: see sexual dimorphism. diphyly: the derivation of a taxon from two separate lines of descent. cf. monophyly (q.v.). diploid number (2n): total number of chromosomes (including sex chromosomes) in a somatic cell of an organism. distal: the end of any structure furthest away from the mid-line of the body or furthest from the point of its attachment. cf. proximal (q.v.). diurnal: at, active in, daytime; when light intensity is high. cf. crepuscular (q.v.); nocturnal (q.v.). DNA hybridization: technique of comparing the similarity between two DNA molecules by reassociating single strands from each molecule and determining the extent of double-helix formation. In phylogenetics, this technique is used to determine the relatedness of two or more taxa. DNA: (abbrev.) deoxyribonucleic acid; the very large self-replicating molecule that carries the genetic information of a chromosome; each molecule is composed of two complementary chains of DNA. dorsoventral (dorsoventrally): from dorsal to ventral surface; from back to belly of an animal.

E: (abbrev.) length of external (outer) ear (= pinna), measured from tip of ear to the posterior point of the ear conch). Length and shape usually affected by preservation. East Africa: Kenya, Uganda, Rwanda, Burundi and Tanzania. eastern Africa: SE Sudan, Ethiopia, Eritrea, Djibouti, Somalia, Kenya, Uganda, Tanzania, Malawi (but only south of L. Malawi and east of the Shire R. Valley) and Mozambique (but only east of Malawi and north of the Zambezi R.). echolocation: the use of reflected ultrasonic pulses of sound to perceive the surroundings (including obstacles, prey and other animals). ecotype: a genetically distinct geographic variety or population within a species, which is adapted to specific environmental conditions. ectoparasite: a parasite that lives on the exterior of an organism (e.g. ticks, fleas, lice). cf. endoparasite (q.v.). ectotympanic: a bony element within the middle ear that supports the tympanic membrane or eardrum. edaphic: influenced by conditions of soil or substratum. emargination: a distinct notch or indentation. embryo number: number of foetuses within the uterus or uteri of the female (as assessed by autopsy). Expressed as mean number (with range from minimum to maximum, and sample size). cf. litter-size (q.v.). enamel: hard material that forms a cap over the dentine component of a tooth; usually the most visable part of a tooth. encephalization quotient (EQ): a measure of comparative brain size or weight defined as the ratio of the actual brain weight to the expected brain weight predicted for an animal of a given body weight. endemic: restricted to, peculiar to, or prevailing in, a specified country or region. endoparasite: a parasite that lives in the interior of an organism (e.g. nematodes, cestodes, blood parasites). cf. ectoparasite (q.v.). entoconid cusp: the posterior cusp on the lingual (inner) side of a lower molar tooth. entotympanic: an independent ossification found in the floor of the tympanic cavity in various extant and extinct eutherian groups, including, for example, Scandentia, Chiroptera, Dermoptera, Hyracoidea, Pholidota, Xenarthra, Carnivora, and Macroscelidea. Eocene: geological Epoch (within the Tertiary Period); 55–38 mya. epiphysis (pl. epiphyses): any part of a long bone that is formed from a different centre of ossification and that later fuses with the bone to form its terminal part. epitympanic recess: a hollow located on the roof of the middle ear. erg: a large, relatively flat area of desert covered with wind-swept sand with little or no vegetation cover (sometimes referred to as a dune sea). evaporative water loss: the loss of water from the body through the skin and/or the lungs. A mechanism used by mammals to reduce Tb (q.v.) when Ta (q.v.) is high. Excessive evaporative water loss may lead to dehydration if free (drinking) water is unavailable. exfoliating: shedding flakes (e.g. of bark), or breaking into relatively thin slabs (e.g. of granitic rock). exoccipital condyles: a pair of projections from the occipital bone on either side of the foramen magnum (q.v.), which articulate with the first of the spinal vertebrae.

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extant: living at the present time. cf. extinct. F. R.: (abbrev.) Forest Reserve. facultative: having the capacity to switch from one mode of life or action to another depending on conditions or circumstances. cf. obligate (q.v.). female-defence polygyny: a mating system in which males control access to females directly, usually by virtue of female gregariousness (Emlen & Oring 1977). fenestra (pl. fenestrae): opening in a bone, or between two bones. flank: the side of the body of a mammal. flehmen: an act performed by many species of mammals whereby an adult male sniffs the vulva and urine of a female to test for oestrus. The head of the male is raised, the nose pointed upwards, the lips retracted and the nose wrinkled. This muscular contraction opens the ductus incisivus ensuring that scent molecules reach the Jacobsen’s organ for olfactory analysis. FN: (abbrev.) see fundamental number. foliaceous: (adj.) resembling the leaf of a plant. folivore (adj. folivorous): an animal that eats leaves. foramen (pl. foramina): an aperture (which is usually small, round or elliptical) in a bone, or between bones, for the passage of a nerve, blood vessel or muscle. foramen magnum: the large opening at the posterior end of the skull through which the spinal cord passes. form: a neutral term for a single individual or taxonomic unit that may be employed without reference to the formal taxonomic hierarch of categories; one of the varieties found in a polymorphic species. forest island: see relict forest. fossa (pl. fossae): a depression or hollow usually in a bone (e.g. glenoid fossa, preorbital fossa) fossorial: adapted for digging; burrowing. cf. subterranean (q.v.). founder effect: the loss of genetic diversity that occurs when a new isolated population is derived from a very small number of individuals. fovea: small pit or depression. frontal bone: one of a pair of bones forming the anterior part of the braincase. frugivorous: fruit-eating. fundamental number (FN): an ambiguous term sometimes defined as (1) the total number of chromosomal arms in the full chromosomal complement of an organism (i.e. including the sex chromosomes), or (2) the total number of chromosomal arms found in the autosomal chromosomes only (i.e. excluding the sex chromosomes). When only the autosomal chromosomes are included, some authors (but not all) use aFN instead of FN to avoid ambiguity. For further details, see aFN. fusiform: elongated and tapering at both ends. fynbos: the heath shrublands characteristic of the Cape Floristic Kingdom (within the South-West Cape Biotic Zone) of South Africa. Dominant plants are sclerophyllous, evergreen, low (2). subterminal: just below the end or tip. subterranean: living permanently below the ground; cf. fossorial (q.v.). suckling: the act of a mother giving milk directly from her breast (mammary glands) to her young. Mothers suckle; their young suck. sulcus (pl. sulci): a groove, fissure or furrow. superovulation: see polyovulation (q.v.). supinate: to turn or rotate the hand or forearm, or the hindlimb and foot. supracaudal: above the tail. supraoccipital crest: ridge of bone, orientated transversely across the back of the skull, at the junction of the parietal and/or supraoccipital bones and the occipital bone. Sometimes referred to as the lambdoid crest. supraorbital ridge: ridge of bone along upper rim of orbit (eyesocket). supraorbital: above (dorsal to) the orbit. supraordinal: describes a taxon above the level of the order. sympatry (adj. sympatric): the situation where populations of two or more different species have overlapping geographic ranges; refers also to populations of two or more species whose geographic ranges are partly or wholly overlapping. They may or may not interact. cf. allopatry (q.v.); syntopy (q.v.).

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symplesiomorphy: a primitive or ancestral character shared by two or more groups, which is inherited from ancestors older than the last common ancestor. synanthropic: associated with humans and/or their houses and other buildings. synapomorphy (adj. synapomorphic): situation in which a homologous character is present in two or more taxa and is thought to have originated in their most recent common ancestor. See also apomorphy. syndactyly: of digits; whole of part fusion of two or more digits (e.g. Digits 2 and 3 of the hindfoot in otter-shrews). synonym: one or more of different names for the same taxonomic unit. A synonym may be a ‘senior synonym’ (the oldest name), or a ‘junior synonym’ (a more recent name) that is no longer considered as valid. May be used to refer to all names that have been associated, at some time in the past, with the taxonomic unit as currently understood. syntopy (adj. syntopic): describes the situation where two or more species use the same or similar habitats and activity times. They may or may not interact. cf. allopatry (q.v.); sympatry (q.v.). syntype: any specimen, or one of a series of specimens, used to designate a species when a holotype (q.v.) and paratype(s) (q.v.) have either not been selected, or have been lost or destroyed. systematics: the science of arranging organisms in a way that reflects their evolutionary relationships; such relationships may be expressed as a phylogeny (q.v.). Often defined (somewhat incorrectly) as a synonym of taxonomy (q.v.). T: (abbrev.) length of tail, measured from anterior of the first caudal vertebra to the posterior end of the last caudal vertebra (excluding any tufts, bristles etc. at tip of tail). Ta: (abbrev.) ambient temperature; the temperature in which an animal is living. cf. Tb (q.v.). Ta: ambient temperature; the temperature in which an animal is living. cf. Tb (q.v.). talonid: heel at the posterior end of a lower molar tooth. tapetum lucidum: light-reflecting layer behind or in the retina of the eyes of some vertebrates that reflects light back through the retina thereby increasing the sensitivity of the eye to dim light. taxon (pl. taxa): any defined unit (e.g. family, genus, species, subspecies) in the classification of organisms. taxonomy: the science of biological nomenclature; the study of the rules, principles and practice of naming and classifying species and other taxa. Sometimes considered as an integral part (and near synonym) of systematics (q.v). Tb: (abbrev.) body temperature; the temperature of the core (central) part of an animal. cf. Ta (q.v.). telocentric: describes a chromosome that appears to have a terminal centromere (q.v.) and therefore only one arm. Modern studies have revealed that all chromosomes have two arms but the smaller arm of telocentric chromosomes is not visible under a light microscope. temporalis: a broad radiating muscle arising from the coronoid process (q.v.) of the lower jaw and attaching to the upper part of the skull. termitarium (pl. termitaria): a place where termites (Insecta: Isopoda) live. Often a large mound of modified hard soil.The shape and size of a termitarium is unique to each species of termite.

terrestrial: living on the ground. cf. arboreal (q.v.); scansorial (q.v.). territory: an area defended by an individual against certain other members of the species, usually by overt aggression or advertisement; territory is marked by the urine, faeces or glandular secretions of the territory’s owner. cf. home-range (q.v.). Tertiary Period: geological period, 65–2 mya, comprising five epochs: Palaeocene, Eocene, Oligocene, Miocene and Pliocene (q.v.); followed by the Quaternary Period (q.v.). testes: the male gonads, or testicles, in which spermatozoa are formed and in which the male hormone is produced. Tethys Sea: the sea separating the two supercontinents, Gondwana (q.v.) and Laurasia (q.v.) during much of the Mesozoic Era before the opening of the Indian and Atlantic oceans during the Cretaceous Period (q.v.). thermal conductance: a measure of the ability of substances (including pelage) to transfer heat. thermolability (adj, thermolabile): the ability of a homeotherm (e.g. camel) to allow its body temperature to vary over a 24-hour period, without either hibernating or aestivating. thermoneutral zone: the range of body temperatures within which an animal does not have to increase its metabolic rate to increase Tb (q.v.) (when Ta (q.v.) is low) and reduce Tb (when Ta is high). thermoregulation: regulation of body temperature, either by metabolic or behavioural means (or both simultaneously) so that Tb (q.v ) is kept more or less constant. thoracic: pertaining to, or situated upon, the chest. through-put time: time taken for food to pass through the digestive tract. tibia (pl. tibiae): one of the two bones forming the lower leg (the shin bone); part of hindlimb between knee and ankle. TL: (abbrev.) total length from tip of snout to posterior end of tail. Equivalent to the head and body length and tail length added together. See also HB (q.v.). and T (q.v.). toothrow: Generally, the row of teeth from the most anterior incisor tooth to the most posterior molar. In golden moles, the row of teeth from the canine to the most posterior molar. Sometimes used in contexts of specific types of teeth, e.g. premolar toothrow, molar toothrow. topotype: any specimen from the type locality (q.v.), i.e. the same locality as that from which the holotype (q.v.) was taken. topotypical: pertaining to the type locality (e.g. a topotypical population is one found at the type locality). torpor (adj. torpid): a state in which there is a (usually shortterm) reduction of metabolic rate and a lowering of Tb (q.v.) when Ta (q.v.) declines; arousal from torpor occurs when Ta increases and without high energy costs to the individual. Torpor is associated with a state of inactivity and reduced responsiveness to stimuli. Torpor lasts for only short periods of time (hours or days). cf. hibernation. tragus: a cartilaginous structure, usually small, projecting from the inner side of the external ear just anterior to the auditory meatus (q.v.). transverse: in a direction across the body from side to side. cf. longitudinal (q.v.). Triassic Period: period (within the Mesozoic Era); 248–208 mya. The first mammals appeared in this period. 619

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triconid: describes a molariform tooth having three cusps. tricuspid: having three points or cusps (particularly of teeth). trifid: divided into three by two emarginations (q.v.). tubercle: a small rounded protuberance. tusks: long, continuously growing incisor or canine teeth that protrude (usually in pairs) beyond the mouth in some mammals including elephants (in which the tusks are incisors), and warthogs and other pigs (in which the tusks are canines); comprised of dentine (ivory). Some mammals, e.g. hyraxes, have ‘tusk-like’ incisors. tympanic bulla (pl. tympanic bullae): one of a pair of usually rounded bony capsules, on underside of skull (one on each side), housing structures of the middle and inner ear in many mammals. Also called auditory bulla (q.v.). type description: the original description of a species; the original description of the holotype (and paratype[s] if included). type locality: the locality from which a holotype (q.v.), lectotype (q.v.) or neotype (q.v.) was collected. Also called topotypical locality. type population: the population from which the holotype was selected. type series: the holotype and all specimens collected at the same place and time and used, together with the holotype, to describe a new species. type species: usually the species that was the first to be described under the name of a new genus. Not all genera had a designated type species when they were first created; in such cases, other rules determine which species will be the type species. type specimen: see holotype. underfur: dense and often woolly layer of the pelage, situated close to the skin and below the soft hairs and guard hairs; usually short and present in those species that experience lower Ta. unicuspid: having one cusp or point (particularly of teeth). upper critical temperature: the highest ambient temperatures at which the animal must increase its metabolic rate to maintain a constant body temperature. If the ambient temperature increases above the upper critical temperature and the animal is unable to cool itself, it will enter hyperthermia and may eventually die. cf. lower critical temperature. uvula: the conical projection from the posterior edge of the soft palate that plays a role in the articulation of sounds and the closing the nasopharynx during swallowing. vagility: the ability to move about, disperse or migrate. vagrant: an individual that has been found well outside the normal geographic range of its species, e.g. a bat or bird that has been wind-borne, or an animal that has been transported as a stowaway on a ship, to a distant locality. vascularized: infiltrated with capillaries. vasoconstriction: constriction of the capillaries of the blood system near the surface of the skin in order to reduce the rate of heat loss through the skin; a mechanism used by many mammals to conserve heat when Ta (q.v.) is low. cf. vasodilation (q.v.).

vasodilation: the dilation (or opening) of the capillaries of the blood system near the surface of the skin in order to increase the rate of heat loss through the skin; a mechanism used by many mammals to cool themselves when Ta (q.v.) is high. cf. vasoconstriction (q.v.). veld: Afrikaans word, used mainly by southern African biologists, to refer to a wide variety of grassland vegetation types typically used for grazing. See also bushveld, highveld, lowveld. vertebra (pl. vertebrae): any of the bones that make up the backbone. vertebral formula: the number of vertebrae in each part of the spine, from anterior to posterior: the parts are cervical (C), thoracic (T), lumbar (L), sacral (S), caudal (Ca). vestigial: small and imperfectly developed; a structure having a smaller and more simple form than the corresponding structure in an ancestral species. vibrissa (pl. vibrissae): long stiff hairs on the face, especially around nostrils and lips; often associated with the perception of tactile sensation; ‘whiskers’. vlei: southern African term for a marsh or swamp, either permanent or seasonal. wadi: a desert valley, usually dry at the surface except after heavy rainfall. water turnover: the rate at which water (fluids) is utilized and replaced in the body per unit time (normally expressed as ml/ kg body weight/day); the amount of water an animal processes through its body each day. Water turnover is related to water availability, the urine concentrating ability of the kidney, amount of protein in the diet and Ta (q.v.). Water turnover rates are characteristically low in arid-adapted mammals when compared with non arid-adapted mammals. West Africa: ca. south of 18° N from Senegal to the Sanaga R. in Cameroon, and Bioko I. (Equatorial Guinea) (Rosevear 1965). WT: (abbrev.) weight (mass) of an individual, usually expressed in grams (g) or kilograms (kg). xiphisternum: The posterior segment, or extremity, of the sternum (sometimes called the xiphoid process). zalambdodont: cheekteeth with three main cusps connected by crests (ectolophs) forming a V-shape; largest cusp is at the apex of the V (on the lingual or tongue side of the tooth); assumed to be derived from the primitive tribosphenic teeth found in some extinct early mammals. cf. dilambdodont (q.v.). ZW: (abbrev.) see zygomatic width. zygomatic arch: one of a pair of cheekbones, formed of the maxillary process anteriorly, jugal bone medially and squamosal bone posteriorly. Ranges from massive, broad, widely flared and bony, to frail, slender and cartilaginous. When present, provides protection to the eyes and orbits. Also called zygoma. zygomatic width (ZW): greatest width between the outer aspect of one zygomatic arch to the equivalent position on the opposite zygomatic arch. See also GWS.

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Authors of Volume VI Philip U. Alkon 3303 East Street Las Cruces, NM 88005 USA email: [email protected] Jeremy Anderson International Conservation Services PO Box 594 White River 1240 South Africa email: [email protected] Peter Arcese Faculty of Forestry University of British Columbia 3rd Floor, Forest Sciences Centre #3041 - 2424 Main Mall Vancouver, BC V6T 1Z4 Canada email: [email protected] Peter Arctander Department of Evolutionary Biology Institute of Biology University of Copenhagen Universitetsparken 15 Copenhagen 2100 Denmark email: [email protected] Nico L. Avenant Department of Mammalogy National Museum Bloemfontein PO Box 266 Bloemfontein 9300 South Africa email: [email protected] Roseline C. Beudels-Jamar Conservation Biology Unit Institut Royal des Sciences Naturelles de Belgique 29, rue Vautier 1000 Bruxelles Belgium email: [email protected]

Jean-Renaud Boisserie Institut de Paléoprimatologie, Paléontologie Humaine: Evolution et Paléoenvironnements UMR 7262, CNRS & Université de Poitiers 5 rue Albert Turpain 86022 POITIERS cedex France email: [email protected] Mathieu Bourgarel CIRAD - Unité AGIRs - Animal et Gestion Intégrée des Risques Campus international de Baillarguet 34 398 Montpellier cedex 5 France email: [email protected] Anthony E. Bowland Department of Conservation and Horticulture Faculty of Health Business and Science Batchelor Institute of Indigenous Tertiary Education c/o Post Office Batchelor NT 0845 Australia email: [email protected] Justin S. Brashares Dept of Environmental Science, Policy, & Management University of California, Berkeley 130 Mulford Hall #3114 Berkeley, CA 94720 USA email: [email protected] Peter N. M. Brotherton Natural England 3rd Floor Touthill Close City Road Peterborough PE1 1UA UK email: peter.brotherton@naturalengland. org.uk

Thomas M. Butynski King Khalid Wildlife Research Centre Saudi Wildlife Commission PO Box 61681 Riyadh 11575 Kingdom of Saudi Arabia email: [email protected] Isabella Capellini Department of Biological Sciences University of Hull Cottingham Road Kingston-upon-Hull HU6 7RX UK email: [email protected] Jorge Cassinello Research Group on Behavioural Ecology and Conservation Biology of Ungulates Instituto de Investigación en Recursos Cinegéticos (IREC), CSIC-UCLM-JCCM Ronda de Toledo s/n 13071 Ciudad Real Spain email: [email protected] Guy Castley International Centre for Ecotourism Research Griffith School of Environment Griffith University, Gold Coast Campus Queensland 4222 Australia email: [email protected] Philippe Chardonnet International Foundation for Wildlife Management 58 rue Beaubourg 75003 Paris France email: [email protected] Isabelle Ciofolo Le Pré Commun 31160 – ASPET France email: [email protected]

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William Crosmary Département de Biologie Université Laval Pavillon Alexandre-Vachon 1045 avenue de la Médecine Québec G1V 0A6 Canada email: [email protected] David H. M. Cumming Percy FitzPatrick Institute, University of Cape Town and Tropical Resource Ecology Programme, University of Zimbabwe PO Box HG 400 Highlands Harare Zimbabwe email: [email protected] Fabrice Cuzin BP 1172 Bab Agnaw 40.000 Marrakech Morocco email: [email protected] Jean-Pierre d’Huart Conservation Consultancy Services SPRL 14 Rue du Monty 1320 Beauvechain Belgium email: [email protected] Tim Davenport Wildlife Conservation Society PO Box 922 Zanzibar Tanzania email: [email protected] Jeremy David 4 Wanderers Close Constantia Hills Cape Town 7806 South Africa email: [email protected] Pierre Devillers Conservation Biology Unit Institut Royal des Sciences Naturelles de Belgique 29, rue Vautier 1000 Bruxelles Belgium email: [email protected]

Patrick Duncan Centre d’Etudes Biologiques de Chizé CNRS-UPR 1934 79360 Beauvoir-sur-Niort France email: [email protected] Johan du Toit Department of Wildland Resources Utah State University 5230 Old Main Hill Logan, UT 84322-5230 USA email: [email protected] Paul W. Elkan International Programs Wildlife Conservation Society 2300 Southern Blvd Bronx, NY 10460 USA email: [email protected] S. Keith Eltringham Deceased Richard D. Estes 5 Granite Street Peterborough, NH 03458 USA email: [email protected] Elisabetta Falchetti Museo Civico di Zoologia Via U. Aldrovandi 18 Rome Italy email: elisabettamaria.falchetti@comune. roma.it François Feer CNRS-MNHN UMR7179 Laboratoire d’écologie 4, Avenue du Petit Château 91800 Brunoy France email: [email protected] Frauke Fischer Zoology III Biozentrum, Am Hubland 97074 Würzburg Germany email: [email protected]. de

Clare FitzGibbon Natural England 3 Moors Team (Bodmin Moor and River Camel) Pydar House Truro TR1 1XU UK email: Clare.Fitzgibbon@naturalengland. org.uk Hervé Fritz CNRS UMR 5558, LBBE Université de Lyon 1 43 bd du 11 nov 1918 69622 Villeurbanne Cedex France email: [email protected] Valerius Geist Faculty of Environmental Design 2500 University Drive The University of Calgary Calgary, Alberta T2N 1N4 Canada email: [email protected] Alan W. Gentry c/o Department of Palaeontology Natural History Museum Cromwell Road London SW7 5BD UK email: [email protected] Nina Giotto Laboratoire Comportement et Ecologie de la Faune Sauvage (CEFS) Institut National de la Recherche Agronomique (INRA) BP 52627 F-31326 Castanet-Tolosan Cedex France email: [email protected] L. Morris Gosling Newcastle Institute for Research on Sustainability University of Newcastle Newcastle upon Tyne, NE1 7RU UK email: [email protected]

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Colin P. Groves School of Archeology and Anthropology Australian National University Canberra, ACT 2601 Australia email: [email protected] Peter Grubb Deceased John M. Harris Vertebrate Paleontology Department Los Angeles County Natural History Museum 900 Exposition Blvd. Los Angeles, CA 90007 USA email: [email protected] John Hart Lukuru Foundation Projet Tshuapa-Lomami-Lualaba (TL2) 1235 Ave Poids Lourds Kinshasa Democratic Republic of Congo email: [email protected] Michael Hoffmann IUCN Species Survival Commission c/o United Nations Environment Programme - World Conservation Monitoring Centre 219c Huntingdon Rd Cambridge CB3 0DL UK email: [email protected] Reino R. Hofmann Trompeterhaus D-15837 Horstwalde Nr Baruth Brandenburg Germany email: [email protected] Bernd Hoppe-Dominik Wilhelmshöhe 14 38108 Braunschweig Germany email: bernd.hoppe-dominik@hondelage. de

Peter P. Hoppe Am Hauenstein 13 D-67157 Wachenheim Germany email: [email protected]

Hans Klingel Hackelkamp 5 38110 Braunschweig Germany email: [email protected]

Ibrahim M. Hashim Sudanese Wildlife Society PO Box 6041 Takamul Khartoum Sudan email: [email protected]

Michael Knight Conservation Services South African National Parks PO Box 76693 NMMU 6031 Port Elizabeth South Africa email: [email protected]

Michael Jacobs PO Box 445 Fortine, MT 59918 USA email: [email protected] Christine Janis Department of Ecology and Evolutionary Biology Box G-B207 Brown University Providence, RI 02912 USA email: [email protected] Richard Jeffery PO Box 32056 Lusaka Zambia email: [email protected] Richard Jenkins IUCN UK 219c Huntingdon Rd Cambridge CB3 0DL UK email: [email protected] Trevor Jones Udzungwa Elephant Project Box 99 Mang’ula Tanzania email: [email protected] Jonathan Kingdon Department of Zoology University of Oxford WildCRU, Tubney House Abingdon Road Tubney OX13 5QL UK

Thomas Künzel BirdLife Indochina – Cambodia Programme PO Box 2686 Phnom Penh Cambodia email: [email protected] Sally A. Lahm Ecology and Environment, Inc. 368 Pleasant View Drive Lancaster, NY 14086 USA email: [email protected] Alain Laurent BEIRA.CFP 49 rue Raymond IV 31000 Toulouse France email: [email protected] Yvonnick Le Pendu Universidade Estadual de Santa Cruz Departamento de Ciéncias Biologicas Campus Soane Nazaré de Andrade CEP 45662-900 Ilhéus, Bahia Brazil email: [email protected] Kristin Leus IUCN/SSC CBSG (Europe) Copenhagen Zoo p/a Annuntiatenstraat 6 2170 Merksem Belgium email: [email protected]

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Authors of Volume VI

Walter Leuthold Kinkelstr. 61 CH-8006 Zürich Switzerland email: [email protected] Rolf Lindholm PO Box 150 Snug Tasmania 7054 Australia email: [email protected] Peter Lloyd email: [email protected] Eline D. Lorenzen Department of Integrative Biology University of California Berkeley 1005 Valley Life Sciences Building Berkeley, CA 94720 USA email: [email protected] Janice May PO Box 150 Snug Tasmania 7054 Australia email: [email protected] Isabelle G. Michaux Chemin des Gandins F-38660 Saint Hilaire du Touvet France email: [email protected] Miranda Mockrin USDA Forest Service Rocky Mountain Research Station 2150A Centre Avenue Fort Collins, CO 80526 USA email: [email protected] Renata Molcanova 5 Wingate Road Totton Southampton SO40 3FU UK email: [email protected] Catherine Morrow Scimitar Scientific Ltd Hamilton 3281 New Zealand

Rory Nefdt UNICEF ESARO PO Box 44145 00100 Nairobi Kenya email: [email protected]

Andrew J. Plumptre Wildlife Conservation Society PO Box 7487 Kampala Uganda email: [email protected]

Dorothe Nett GIZ Tanzania Deutsche Gesellschaft für Internationale Zusammenarbeit (GIZ) GmbH 65 Ali Hassan Mwinyi Road PO Box 1519 Dar-es-Salaam Tanzania email: [email protected]

Herbert H. T. Prins Resource Ecology Group Department of Environmental Sciences Wageningen University Droevendaalsesteeg 3a 6708PB Wageningen The Netherlands email: [email protected]

John Newby rue des Tigneuses 2 1148 L’Isle Switzerland email: [email protected] Helen Newing Durrell Institute of Conservation and Ecology School of Anthropology and Conservation Marlowe Building University of Kent Canterbury Kent CT2 7NR UK email: [email protected] Bernhard Nievergelt Wildlife and Conservation Biology University of Zürich (retired) Burenweg 52 8053 Zürich Switzerland email: [email protected] Norman Owen-Smith School of Animal, Plant & Environmental Sciences University of the Witwatersrand Wits 2050 South Africa email: [email protected] Hubert P. Planton Chemin des Gandins F-38660 Saint Hilaire du Touvet France email: [email protected]

Ettore Randi Facoltà di Scienze Matematiche, Fisiche e Naturali Università di Bologna Via Selmi, 3 Bologna Italy email: [email protected] S. Craig Roberts Senior Lecturer School of Natural Sciences University of Stirling Stirling FK9 4LA UK email: [email protected] Philip T. Robinson Department of Laboratory Animal Resources University of Toledo 3000 Arlington Ave. MS 1004 Toledo, Ohio 43614 USA email: [email protected] Francesco Rovero Tropical Biodiversity Section Museo delle Scienze Via Calepina 14 38122 Trento Italy email: [email protected] Catherine Schloeder PO Box 445 Fortine, MT 59918 USA email: [email protected]

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Authors of Volume VI

Paul Scholte Gestion durable des forêts dans le Bassin du Congo Deutsche Gesellschaft für Internationale Zusammenarbeit (GIZ) GmbH B.P. 7814 Yaoundé Cameroon email: [email protected] Erik R. Seiffert Department of Anatomical Sciences Stony Brook University Health Sciences Center T-8, Stony Brook New York, 11794 USA email: [email protected] Armin H. W. Seydack Conservation Services Division South African National Parks PO Box 3542 Knysna 6570 South Africa email: [email protected] Russell Seymour International Giraffe Working Group email: [email protected] Hans R. Siegismund Department of Evolutionary Biology Institute of Biology University of Copenhagen Universitetsparken 15 2100 Copenhagen Denmark email: [email protected] Claudio Sillero-Zubiri Department of Zoology University of Oxford WildCRU, Tubney House Abingdon Road Tubney OX13 5QL UK email: [email protected] Alberto M. Simonetta Dip Biologia Animale e Genetica Firenze Italy

Anthony R. E. Sinclair Beaty Biodiversity Research Centre University of British Columbia 6270 University Blvd. Vancouver, B.C. V6T 1Z4 Canada email: [email protected]

Paul Vercammen Sharjah Breeding Centre for Endangered Arabian Wildlife PO Box 29922 Sharjah United Arab Emirates email: [email protected]

John D. Skinner Deceased

Savvas Vrahimis Free State Department of Tourism, Environment & Economic Affairs Scientific Support Services Private Bag X 20801 Bloemfontein 9300 South Africa email: [email protected]

James L.D. Smith Department of Fisheries, Wildlife and Conservation Biology University of Minnesota St. Paul, MN USA email: [email protected] Clive A. Spinage Wickwood House, Stanford Road Faringdon Oxon SN7 8EZ UK email: [email protected] Chris R. Thouless Conservancy Development Support Services Programme World Wildlife Fund PO Box 9681 Windhoek Namibia email: [email protected] Thomas L. Thurow Professor of Rangeland and Watershed Management Ecosystem Science & Management Department University of Wyoming Laramie, WY 82071 USA email: [email protected] Pim van Hooft Resource Ecology Group Lumen Building Wageningen University The Netherlands email: [email protected]

Tim Wacher Conservation Programmes Zoological Society of London Regent’s Park London NW1 4RY UK email: [email protected] Friedrich K. Wilhelmi IUCN SSC Antelope Specialist Group North-east African Subgroup email: [email protected] Andrew Williams Maliasili Initiatives PO Box 293 Underhill, VT 05489 USA email: [email protected] John Wilmshurst Jasper National Park Parks Canada PO Box 10 Jasper, Alberta T0E 1E0 Canada email: [email protected] Vivian J. Wilson Deceased Torsten Wronski King Khalid Wildlife Research Centre Saudi Wildlife Commission PO Box 61681 Riyadh 11575 Kingdom of Saudi Arabia email: [email protected]

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Derek W. Yalden School of Life Sciences University of Manchester Manchester M13 9PL UK email: [email protected]

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Indexes French names Addax 566 Algazel 586 Antilope de Bates 208 Antilope Harnaché 163 Antilope Hirola 491 Antilope musquée 214 Antilope royale 211 Beira 315 Biche-Robert 382 Blesbok 496 Bongo 179 Bontebok 496 Bouquetin de Nubie 600 Bouquetin d’Ethiopie 603 Bubale 511 Buffle d’Afrique 125 Buffle noir 125 Cépalophe rayé 245 Cépalophe zébré 245 Céphalophe à dos jaune 288 Cephalophe à flancs roux 265 Céphalophe à front noir 268 Céphalophe à ventre blanche 255 Céphalophe bai 294 Céphalophe bleu 228 Céphalophe couronne 235 Céphalophe d’Abbott 285 Céphalophe de Aders 248 Céphalophe de Grimm 235 Céphalophe de Harvey 261

Céphalophe de Jentink 299 Céphalophe de Maxwell 224 Céphalophe de Peters 279 Céphalophe de Rwenzori 253 Céphalophe de Weyns 275 Céphalophe d’Ogilby 272 Céphalophe du Cap 235 Céphalophe du Gabon 255 Céphalophe du Natal 258 Céphalophe noir 281 Cerfe de Berberie 117 Chevrotain aquatique 88 Cobe de Buffon 439 Cobe de Roseaux 431 Cobe onctueux 461 Damalisque 502 Dik-dik argente 325 Dik-dik de Günther 334 Dik-dik de Salt 323 Éland de cap 191 Éland de Derby 186 Gazelle à front rouge 357 Gazelle à longues cornes 352 Gazelle blanche 352 Gazelle Dama 382 Gazelle de Bright 375 Gazelle de Cuvier 349 Gazelle de Grant 374 Gazelle de Mongalla 369

Gazelle de montagne 349 Gazelle de Peters 376 Gazelle d’Eritrée 359 Gazelle de Soemmerring 380 Gazelle de Speke 346 Gazelle de Thomson 361 Gazelle de Waller 391 Gazelle dorcas 340 Gazelle du Clarke 388 Gazelle-Girafe 391 Gazelle leptocére 352 Gazelle leptocère 352 Gemsbok 572 Girafe 98 Gnou à queue blanche 528 Gnou bleu 533 Grand Koudou 152 Grysbok de Sharpe 308 Grysbok du Cap 304 Guib Harnaché 163 Hippopotame 68 Hippopotame nain 80 Hippopotame pygmée 80 Hippotrague rouan 548 Hylochère 42 Impala 480

Le Wali 603 L’Hippotrague noir 556 L’Oréotrague 470 Mouflon à manchettes 595 Nyala 148 Nyale des montagnes 159 Okapi 110 Oryx algazelle 586 Oryx beisa 576 Ourebie 406 Petit Koudou 142 Phacochère 54 Phacochère du déser 51 Potamochère 32, 37 Puku 445 Redunca 431 Redunca de montagne 422 Redunca grande 426 Rhebouk 417 Sanglier 28 Sitatunga 172 Springbok 398 Steenbok 311

Le Cobe de Madame Gray 455 Le Cobe lechwe 449 Le Dik-dik de Kirk 328

German names Abbottducker 285 Addax 566 Adersducker 248 Aethiopischer Steinbock 603 Batesbockchen 208 Beira 315 Berberhirsch 117 Bergnyala 159 Bergriedbock 422 Blauducker 228 Bleichbockchen 406 Blessbock 496 Bongo 179 Bright-Gazelle 375 Buntbock 496 Buschschwein 32 Damagazelle 382

Dorkasgazelle 340 Dünengazelle 352

Grosskudu 152 Grossriedbock 426

Echtgazelle 349 Elanantilope 191 Eritrea-Dikdik 323 Eritreagazelle 359 Eritrea-Spiessbock 583

Harveyducker 261 Hirschferkel 88 Hunters Leierantilope 491

Flusschschwein 37 Flusspferd 68 Frau Grays Wasserbocke 455

Kaapgreisbock 304 Kaffernbüffel 125 Kirkdikdik 327, 328 Kleiner Kudu 142 Kleinkudu 142 Kleinstbockchen 211 Klippspringer 470 Kronenducker 235 Kuhantilope 511

Gemeiner Riedbock 431 Gerenuk 391 Giraffe 98 Giraffengazelle 391 Grant-Gazelle 373, 374 Grasantilope 439

Jentinkducker 299

Lamagazelle 388 Leierantilope 502 Litschi 449 Mähnenschaf 595 Mähnenspringer 595 Maxwellducker 224 Mendesantilope oder 566 Mongallagazelle 369 Moorantilope 449 Moschusboeckchen 214 Natal-Rotducker 258 Nordafrikanischer Spießbock 572 Nubischer Steinbock 600 Ogilbyducker 272 Okapi 110

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English names

Petersducker 279 Peters-Gazelle 376 Pferdeantilope 548 Piacentini Dik-dik 325 Pinselohrschwein 37 Puku 445 Rappenantilope 556 Rehantilope 417 Reisenelenantilope 186 Riesenducker 288 Riesenwaldschwein 42 Rotflankenducker 265

Rotstirngazelle 357 Rwenzoriducker 253 Säbelantilope 572 Schirrantilope 163 Schwartzruckenducker 294 Schwarzducker 281 Schwarzfersenantilope 480 Schwarzstirnducker 268 Sharpegreisbock 308 Sömmerring-Gazelle 380 Spekegazelle 346 Springbok 398

Steinbockchen 311 Stelzengazelle 388 Streifengnu 533 Südafrikanischer Spiessbock 579 Sumpfantilope 172 Swahili Choroa 583

Weissnacken-Moorantilope 455 Weissschwanzgnu 528 Weusbauch Ducker 255 Weynsducker 275 Wildschwein 28 Wüstenwarzenschwein 51

Thomsongazelle 361 Tiefland-Nyala 148

Zebraducker 245 Zwergflusspferd 80 Zwerg-rüsselantilope 334

Walia-Steinbock 603 Warzenschwein 54 Wasserbock 461

English names Addax 566 Alcelaphines 488–543 Antelope, Bates’s Dwarf 208 Bates’s Pygmy 208 Dwarf 208 Roan 547, 548 Royal 211 Sable 547, 556 Antelopes 199–592 Dwarf 206–219 Gazelline 338–403 Horse-like 544–592 Spiral-horned 122, 137–198 Antilopines 120, 199–592 Aoudad 594, 595 Arui 595 Beira 302, 315 Blesbok 496 Boar, Wild 28 Bongo 179 Bontebok 496 Bovids 22 Bovines 120–198 Bovoids 120–606 Buffalo, African 122, 124, 125 Bushbuck 163 Bushpig 31, 32 Cetaceans 61 Chevrotain, Water 88 Chevrotains 22, 86–92 Damalisks 495–510 Deer 22, 115–119 Barbary Red 117 Old World 116 Red 117 Dibatag 387, 388 Dik-dik, Cavendish’s 329 Damara 329 Günther’s 334 Kirk’s 328 Kirk’s, Species Group 327 Naivasha 329 Piacentini’s 325 Salt’s 323 Silver 325 Thomas’s 329

Ugogo 329 Dik-diks 319–337 Duiker, Abbott’s 285 Aders’s 248 Banded 245 Bay 294 Black 281 Black-fronted 268 Blue 228 Bush 235 Common 235 Grey 235 Grimm’s 235 Harvey’s 261 Jentink’s 299 Maxwell’s 224 Natal Red 258 Ogilby’s 272 Peters’s 279 Red-flanked 265 Rwenzori Red 253 Walter’s 17 Weyns’s 275 White-bellied 255 Yellow-backed 288 Zebra 245 Duikers 220–301 Blue 223–234 Forest 244–301 Eland, Common 191 Giant 186 Lord Derby’s 186 Gazelle, Addra 382 Atlas 349 Bright’s 375 Clarke’s 388 Cuvier’s 349 Dama 382 Dorcas 340 Edmi 349 Grant’s 374 Grant’s, Species Group 373 Heuglin’s 359 Loder’s 352 Mongalla 369 Peters’s 376 Red-fronted 357

Rhim 352 Slender-horned 352 Soemmerring’s 380 Speke’s 346 Tana 376 Thomson’s 361 Waller’s 391 Gazelles, Greater 372–387 Ring-horned 356–372 Slender 339–355 Gedemsa 159 Gemsbok 572 Gerenuk 390, 391 Giraffe 94, 95, 96, 98 Gnu, White-tailed 528 Goats 593, 599–606 Grysbok, Cape 304 Sharpe’s 308 Grysboks 302, 303 Hartebeest 510, 511 Hippopotamus, Common 64, 68 Pygmy 78, 80 Hippopotamuses 22, 61–83 Hirola 490, 491 Hog, Forest 40, 42 Giant Forest 42 Red River 31, 37 Ibex, Ethiopian 603 Nubian 600 Walia 603 Ibexes 599–606 Impala 477, 479, 480 Klipspringer 469, 470 Kob 439 Kobs 437–468 Korrigum 502 Kudu, Greater 152 Lesser 142 Lechwe, Mrs Gray’s 455 Nile 455 Southern 449 Nyala 148 Mountain 159

Okapi 94, 95, 110 Oribi 404, 405, 406 Oryx, Beisa 576 Fringe-eared 576 Scimitar 586 Scimitar-horned 586 Southern 572 Oryxes 571–592 Pig, Eurasian Wild 28 Pigs 22–60 Old World 27 Puku 445 Reduncines 413–468 Reedbuck, Bohor 431 Common 431 Mountain 422 Southern 426 Reedbucks 421–436 Rhebok 417 Grey 416, 417 Vaal 417 Ruminants 84–606 Horned 93–606 Sheep 593–599 Barbary 595 Sitatunga 172 Springbok 398 Springbuck 398 Steenbok 302, 303, 311 Steinbok 311 Steinbuck 311 Suni 213, 214 Tiang 502 Topi 502 Tsessebe 502 Warthog, Common 54 Desert 51 Warthogs 49–60 Waterbuck 461 Wildebeest, Black 528 Common 533 Wildebeest 527–543

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Indexes

Scientific names Addax 566–571 nasomaculatus 566 Aepyceros 479–487 melampus 480 Aepycerotini 477–487 Alcelaphini 488–543 Alcelaphus 510–526 buselaphus 511 Ammodorcas 387–390 clarkei 388 Ammotragus 594–599 lervia 595 Ancodonta 62–83 Antidorcas 398–403 marsupialis 398 Antilopinae 199–606 Antilopini 338–403 Beatragus 490–495 hunteri 491 Bovidae 120–606 Bovinae 122–198 Bovini 124–136 Bovoidea 120–606 Capra 599–606 nubiana 600 walie 603 Caprini 593–606 Cephalophini 220–301 Cephalophus 244–301 adersi 248 callipygus 279 dorsalis 294 harveyi 261 jentinki 299 leucogaster 255 natalensis 258 niger 281 nigrifrons 268 ogilbyi 272 rubidus 253 rufilatus 265 silvicultor 288 spadix 285 weynsi 275 zebra 245 Cervidae 116–119 Cervinae 116–119 Cervoidea 115–119

Cervus 117–119 elaphus 117 Cetartiodactyla 22–606 Choeropsis 78–83 liberiensis 80 Connochaetes 527–543 gnou 528 taurinus 533 Damaliscus 495–510 lunatus 502 pygargus 496 Dorcatragus 315–318 megalotis 315 Eudorcas 356–372 albonotata 369 rufifrons 357 thomsonii 361 tilonura 359 Gazella 339–355 cuvieri 349 dorcas 340 leptoceros 352 spekei 346 Giraffa 96–110 camelopardalis 98 Giraffidae 95–115 Giraffinae 96–110 Giraffoidea 94–115 Hippopotamidae 63–83 Hippopotamus 64–78 amphibius 68 Hippotragini 544–592 Hippotragus 547–565 equinus 548 niger 556 Hyemoschus 88–92 aquaticus 88 Hylochoerus 40–49 meinertzhageni 42 Kobus 437–468 ellipsiprymnus 461 kob 439 leche 449 megaceros 455 vardonii 445

Litocranius 390–397 walleri 391 Madoqua 320–337 guentheri 334 (kirkii) 327 (kirkii) cavendishi 329 (kirkii) damarensis 329 (kirkii) kirkii 328 (kirkii) thomasi 329 piacentinii 325 saltiana 323 Madoquini 319–337 Nanger 372–387 dama 382 (granti) 373 (granti) granti 374 (granti) notata 375 (granti) petersii 376 soemmerringi 380 Neotragini 206–219 Neotragus 207–213 batesi 208 pygmaeus 211 Nesotragus 213–219 moschatus 214 Okapia 110–115 johnstoni 110 Okapinae 110–115 Oreotragini 469–476 Oreotragus 469–476 oreotragus 470 Oryx 571–592 beisa 576 dammah 586 gazella 572 Ourebia 405–413 ourebi 406 Ourebiini 404–413 Pecora 93–606 Pelea 416–420 capreolus 417 Phacochoerini 49–60 Phacochoerus 50–60 aethiopicus 51 africanus 54 Philantomba 223–234

maxwelli 224 monticola 228 walteri 17 Potamochoerus 31–40 larvatus 32 porcus 37 Raphicerini 302–318 Raphicerus 303–314 campestris 311 melanotis 304 sharpei 308 Redunca 421–436 arundinum 426 fulvorufula 422 redunca 431 Reduncini 413–468 Ruminantia 84–606 Suidae 25–60 Suiformes 24–60 Suinae 25–60 Suini 27–49 Suoidea 24–60 Sus 28–31 scrofa 28 Sylvicapra 235–243 grimmia 235 Syncerus 124–136 caffer 125 Tragelaphini 137–199 Tragelaphus 138–198 angasii 148 buxtoni 159 derbianus 186 eurycerus 179 imberbis 142 oryx 191 scriptus 163 spekii 172 strepsiceros 152 Tragulidae 87–92 Tragulina 86–92 Traguloidea 87–92 Whippomorpha 61–83

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