Mammals of Africa Volume III: Rodents, Hares and Rabbits 9781472926937, 9781408189924

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mammals of africa volume III

rodents, hares and rabbits

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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 III

rodents, hares and rabbits edited by david c. d. happold

Colour and pencil illustrations by Jonathan Kingdon Pen and ink illustrations by Meredith Happold

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First published in 2013 Copyright © 2013 by Bloomsbury Publishing Copyright © 2013 illustrations by Jonathan Kingdon and Meredith Happold 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) 978-1-4081-2253-2 ISBN (epdf) 978-1-4081-8992-4 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: Happold, D. C. D. (ed.) 2013. Mammals of Africa.Volume III: Rodents, Hares and Rabbits. Bloomsbury Publishing, London. Chapter/species profile: e.g. Emmons, L. H. 2013. Protoxerus stangeri Forest Giant Squirrel; pp 90–91 in Happold, D. C. D. (ed.) 2013. Mammals of Africa:Volume III. 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 Acknowledgements16 Acknowledgements for Volume III17 Mammals of Africa: An Introduction and Guide – David Happold, Michael Hoffmann, Thomas Butynski & Jonathan Kingdon

18

Order RODENTIA Rodents – M. D. Carleton & D. C. D. Happold

27

Family SCIURIDAE Squirrels – R. W. Thorington, Jr

38

Genus Allosciurus Slender-tailed Squirrel – P. Grubb Allosciurus aubinnii Slender-tailed Squirrel (Aubinn’s Squirrel) – C. E. Schennum & R. W. Thorington, Jr

40

Genus Atlantoxerus Barbary Ground Squirrel – S. Aulagnier Atlantoxerus getulus Barbary Ground Squirrel – S. Aulagnier, P. Gouat & M. Thévenot

42

Genus Epixerus Western Palm Squirrel – L. H. Emmons Epixerus ebii Western Palm Squirrel – L. H. Emmons

44 45

41

43

Genus Funisciurus Rope Squirrels – P. Grubb 46 Funisciurus anerythrus Thomas’s Rope Squirrel (Redless Tree Squirrel) – L. H. Emmons 49 Funisciurus bayonii Lunda Rope Squirrel – R. W. Thorington, Jr, L. A. Pappas & C. E. Schennum 50 Funisciurus carruthersi – Carruthers’s Rope Squirrel (Carruthers’s Mountain Squirrel) – J. Kingdon 51 Funisciurus congicus Congo Rope Squirrel (Striped Tree Squirrel) – R. W. Thorington, Jr, L. A. Pappas & C. E. Schennum 52 Funisciurus duchaillui du Chaillu’s Rope Squirrel – D. Brugière 54 Funisciurus isabella Lady Burton’s Rope Squirrel – L. H. Emmons 55 Funisciurus lemniscatus Ribboned Rope Squirrel – L. H. Emmons 56 Funisciurus leucogenys Red-cheeked Rope Squirrel – J. C. Ray 57 Funisciurus pyrropus Fire-footed Rope Squirrel (Redlegged Rope Squirrel) – L. H. Emmons 58 Funisciurus substriatus Kintampo Rope Squirrel – R. W. Thorington, Jr & C. E. Schennum 60 Genus Heliosciurus Sun Squirrels – P. Grubb Heliosciurus gambianus Gambian Sun Squirrel – D. C. D. Happold Heliosciurus mutabilis Mutable Sun Squirrel – D. C. D. Happold

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61 62 64

Heliosciurus punctatus Punctate Sun Squirrel (Small Sun Squirrel) – R. W. Thorington, Jr & C. E. Schennum Heliosciurus rufobrachium Red-legged Sun Squirrel – L. H. Emmons Heliosciurus ruwenzorii Rwenzori Sun Squirrel – J. Kerbis Peterhans & R. W. Thorington, Jr Heliosciurus undulatus Zanj Sun Squirrel – C. E. Schennum & R. W. Thorington, Jr Genus Myosciurus African Pygmy Squirrel – L. H. Emmons Myosciurus pumilio African Pygmy Squirrel – L. H. Emmons

65 66 68 69 70 71

Genus Paraxerus Bush Squirrels – P. Grubb 72 Paraxerus alexandri Alexander’s Bush Squirrel – J. Kingdon 74 Paraxerus boehmi Boehm’s Bush Squirrel – J. Kingdon 75 Paraxerus cepapi Smith’s Bush Squirrel – L. A. Pappas & R. W. Thorington, Jr 77 Paraxerus cooperi Cooper’s Bush Squirrel (Cooper’s Mountain Squirrel) – R.W. Thorington, Jr & C. E. Schennum 79 Paraxerus flavovittis Striped Bush Squirrel – C. E. Schennum & R. W. Thorington, Jr 80 Paraxerus lucifer Black-and-red Bush Squirrel – C. E. Schennum & R. W. Thorington, Jr 81 Paraxerus ochraceus Ochre Bush Squirrel – R. W. Thorington, Jr & C. E. Schennum 82 Paraxerus palliatus Red Bush Squirrel – R. W. Thorington, Jr, L. A. Pappas & C. E. Schennum 84 Paraxerus poensis Green Bush Squirrel – L. H. Emmons 85 Paraxerus vexillarius Swynnerton’s Bush Squirrel – R. W. Thorington, Jr & C. E. Schennum 87 Paraxerus vincenti Vincent’s Bush Squirrel (Selinda Mountain Squirrel) – R. W. Thorington, Jr & C. E. Schennum 88 Genus Protoxerus Forest Giant Squirrel – P. Grubb Protoxerus stangeri Forest Giant Squirrel (African Giant Squirrel) – L. H. Emmons

89 90

Genus Sciurus Squirrels92 Sciurus carolinensis Grey Squirrel – D. C. D. Happold 92 Genus Xerus Ground Squirrels – J. M. Waterman Xerus erythropus Striped Ground Squirrel (African Ground Squirrel, West African Ground Squirrel, Geoffrey’s Ground Squirrel) – J. M. Waterman Xerus inauris Cape Ground Squirrel (South African Ground Squirrel) – J. M. Waterman Xerus princeps Damara Ground Squirrel (Mountain Ground Squirrel, Kaokoveld Ground Squirrel) – J. M. Waterman

93 94 96 99

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Contents

Xerus rutilus Unstriped Ground Squirrel (Pallid Ground Squirrel) – J. M. Waterman

100

Family GLIRIDAE Dormice – M. E. Holden

102

Genus Eliomys Garden Dormice – M. E. Holden Eliomys melanurus Large-eared Garden Dormouse – M. E. Holden Eliomys munbyanus Maghreb Garden Dormouse – M. E. Holden

104

Genus Graphiurus African Dormice – M. E. Holden Graphiurus angolensis Angolan African Dormouse – M. E. Holden Graphiurus christyi Christy’s African Dormouse – M. E. Holden Graphiurus crassicaudatus Thick-tailed African Dormouse – M. E. Holden Graphiurus johnstoni Johnston’s African Dormouse – M. E. Holden Graphiurus kelleni Kellen’s African Dormouse – M. E. Holden Graphiurus lorraineus Lorraine’s African Dormouse – M. E. Holden Graphiurus microtis Noack’s African Dormouse – M. E. Holden Graphiurus monardi Monard’s African Dormouse – M. E. Holden Graphiurus murinus Forest African Dormouse – M. E. Holden Graphiurus nagtglasii Nagtglas’s African Dormouse – M. E. Holden Graphiurus ocularis Spectacled African Dormouse (Namtap) – M. E. Holden Graphiurus platyops Flat-headed African Dormouse – M. E. Holden Graphiurus rupicola Rupicolous African Dormouse – M. E. Holden Graphiurus surdus Short-eared African Dormouse – M. E. Holden

109

Family DIPODIDAE Jerboas – D. C. D. Happold

105 107

Spalax ehrenbergi Middle East Blind Mole-rat (Ehrenberg’s Mole-rat) – D. C. D. Happold

145

Subfamily TACHYORYCTINAE African Root-rats – G. G. Musser & M. D. Carleton

147

Genus Tachyoryctes Root-rats – J. U. M. Jarvis Tachyoryctes macrocephalus Giant Root-rat (Giant Mole-rat) – D. W.Yalden Tachyoryctes splendens African Root-rat – J. U. M. Jarvis

148 149 151

Family NESOMYIDAE Pouched Rats and Mice, Swamp Mouse, Climbing Mice, Large-eared Mouse, Fat Mice, White-tailed Rat and Rock Mice – M. D. Carleton & G. G. Musser

153

113

Subfamily CRICETOMYINAE Pouched Rats and Pouched Mice – M. D. Carleton & G. G. Musser

153

114 116

Genus Beamys Long-tailed Pouched Rat – D. C. D. Happold 154 Beamys hindei Long-tailed Pouched Rat – D. C. D. Happold 155

118

Genus Cricetomys Giant Pouched Rats – J. C. Ray & J.-M. Duplantier Cricetomys emini Emin’s Giant Pouched Rat (Forest Giant Pouched Rat) – J. C. Ray Cricetomys gambianus Gambian Giant Pouched Rat – J.-M. Duplantier & L. Granjon

110 112

120 123 124 126 128 130 131

Genus Saccostomus Pouched Mice – D. C. D. Happold Saccostomus campestris Cape Pouched Mouse (Southern African Pouched Mouse) – M. Perrin Saccostomus mearnsi Mearns’s Pouched Mouse (East African Pouched Mouse) – F. Keesing

157 158 159 161 162 164

Subfamily DELANYMYINAE Delany’s Swamp Mouse – G. G. Musser & M. D. Carleton

165

135

Genus Delanymys Delany’s Swamp Mouse – F. Dieterlen Delanymys brooksi Delany’s Swamp Mouse – F. Dieterlen

166 166

Genus Allactaga Jerboas – D. C. D. Happold Allactaga tetradactyla Four-toed Jerboa – D. C. D. Happold

136 136

Subfamily DENDROMURINAE African Climbing Mice – G. G. Musser & M. D. Carleton

168

Genus Jaculus Jerboas – D. C. D. Happold Jaculus jaculus Lesser Egyptian Jerboa – D. C. D. Happold Jaculus orientalis Greater Egyptian Jerboa (Oriental Jerboa) – D. C. D. Happold

137 138

Genus Dendromus African Climbing Mice – F. Dieterlen Dendromus insignis Montane African Climbing Mouse – F. Dieterlen Dendromus kahuziensis Kahuzi African Climbing Mouse – F. Dieterlen Dendromus lovati Lovat’s African Climbing Mouse – D. W.Yalden Dendromus melanotis Grey African Climbing Mouse – A. Monadjem Dendromus mesomelas Brants’s African Climbing Mouse – A. Monadjem Dendromus messorius Banana African Climbing Mouse – D. C. D. Happold

Family SPALACIDAE Blind Mole-rats, African Root-rats, Zokors and Bamboo-rats – G. G. Musser & M. D. Carleton

133

141

143

Subfamily SPALACINAE Mole-rats – G. G. Musser & M. D. Carleton

144

Genus Spalax Mole-rats – D. C. D. Happold

145

169 171 172 173 174 176 177

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Contents

Dendromus mystacalis Chestnut African Climbing Mouse – A. Monadjem Dendromus nyasae (kivu) Kivu African Climbing Mouse – F. Dieterlen Dendromus nyikae Nyika African Climbing Mouse – D. C. D. Happold Dendromus oreas Cameroon African Climbing Mouse – F. Dieterlen Dendromus vernayi Vernay’s African Climbing Mouse – F. Dieterlen Genus Dendroprionomys Velvet Climbing Mouse – D. C. D. Happold Dendroprionomys rousseloti Velvet Climbing Mouse (Congo Tree Mouse) – C. Denys Genus Malacothrix Long-eared Mouse – D. C. D. Happold Malacothrix typica Long-eared Mouse – D. C. D. Happold

178 179

182 183 184

210

Genus Microtus Voles – D. C. D. Happold Microtus mustersi Musters’s Vole – D. C. D. Happold

211 211

Subfamily LOPHIOMYINAE Maned Rat – M. D. Carleton & G. G. Musser

212

185 186 186

Family MURIDAE Gerbils, Jirds, Rats and Mice – M. D. Carleton & G. G. Musser

216

Subfamily DEOMYINAE Spiny Mice, Link Rat and Brushfurred Rats – M. D. Carleton & G. G. Musser

217

189

Genus Steatomys Fat Mice – A. Monadjem Steatomys bocagei Bocage’s Fat Mouse – D. C. D. Happold Steatomys caurinus North-western Fat Mouse – B. Sicard & J.-M. Duplantier Steatomys cuppedius Dainty Fat Mouse – J.-M. Duplantier & B. Sicard Steatomys jacksoni Jackson’s Fat Mouse – D. C. D. Happold Steatomys krebsii Krebs’s Fat Mouse – A. Monadjem Steatomys opimus Pousargues’s Fat Mouse – D. C. D. Happold Steatomys parvus Tiny Fat Mouse – A. Monadjem Steatomys pratensis Common Fat Mouse – A. Monadjem

191 192

190

193 194 195 196 197 198 199

213 214

Genus Acomys Spiny Mice – F. Dieterlen Acomys airensis Aïr Spiny Mouse (Western Saharan Spiny Mouse) – L. Granjon & K. Bâ Acomys cahirinus Cairo Spiny Mouse (Northeast African Spiny Mouse) – F. Dieterlen Acomys cineraceus Grey Spiny Mouse (Heuglin’s Spiny Mouse) – F. Dieterlen Acomys ignitus Fiery Spiny Mouse – F. Dieterlen Acomys johannis Johan’s Spiny Mouse – L. Granjon Acomys kempi Kemp’s Spiny Mouse – F. Dieterlen Acomys louisae Louise’s Spiny Mouse – F. Dieterlen Acomys mullah Mullah Spiny Mouse – F. Dieterlen Acomys percivali Percival’s Spiny Mouse – S. Takata Acomys russatus Golden Spiny Mouse – F. Dieterlen Acomys spinosissimus Least Spiny Mouse (Southern Spiny Mouse) – D. C. D. Happold Acomys subspinosus Cape Spiny Mouse – E. R. Dempster Acomys wilsoni Wilson’s Spiny Mouse – S. Takata

217

Genus Deomys Rusty Link Rat – J .C. Ray & J. R. Malcolm Deomys ferrugineus Rusty Link Rat (Congo Forest Rat) – J. C. Ray & J. R. Malcolm

235

219 220 222 223 224 225 226 227 228 230 231 233 234

201

Genus Mystromys African White-tailed Rat – D. C. D. Happold 201 Mystromys albicaudatus African White-tailed Rat (Southern African Hamster) – M. Perrin 201

Genus Petromyscus Pygmy Rock Mice – C. G. Coetzee Petromyscus barbouri Barbour’s Pygmy Rock Mouse (Namaqua Pygmy Rock Mouse) – C. G. Coetzee Petromyscus collinus Pygmy Rock Mouse – C. G. Coetzee Petromyscus monticularis Brukkaros Pygmy Rock Mouse (Short-eared Pygmy Rock Mouse) – C. G. Coetzee Petromyscus shortridgei Shortridge’s Pygmy Rock Mouse (Kaokoveld Pygmy Rock Mouse) – C. G. Coetzee

Subfamily ARVICOLINAE Voles, Lemmings and Muskrats – M. D. Carleton & G. G. Musser

Genus Lophiomys Maned Rat – D. C. D. Happold Lophiomys imhausi Maned Rat (Crested Rat) – D. C. D. Happold

Genus Prionomys Bates’s Climbing Mouse – C. Denys Prionomys batesi Bates’s Climbing Mouse (Dollman’s Tree Mouse) – C. Denys

Subfamily PETROMYSCINAE Pygmy Rock Mice – M. D. Carleton & G. G. Musser

210

181

Genus Megadendromus Bale Mouse – D. W.Yalden 188 Megadendromus nikolausi Bale Mouse (Nikolaus’s African Climbing Mouse, Giant Climbing Mouse) – D. W.Yalden 188

Subfamily MYSTROMYINAE White-tailed Rat – M. D. Carleton & G. G. Musser

Family CRICETIDAE Hamsters, Voles and New World Mice and Rats – M. D. Carleton & G. G. Musser

203 204 204 205 207 208

237

Genus Lophuromys Brush-furred Rats – F. Dieterlen 238 Lophuromys brevicaudus Short-tailed Brush-furred Rat – L. A. Lavrenchenko 240 Lophuromys chrysopus Golden-footed Brush-furred Rat (Ethiopian Forest Brush-furred Rat) – L. A. Lavrenchenko 242 Lophuromys cinereus Grey Brush-furred Rat – F. Dieterlen 243 Lophuromys dieterleni Dieterlen’s Brush-furred Rat – F. Dieterlen 244 Lophuromys eisentrauti Eisentraut’s Brush-furred Rat – F. Dieterlen 245 Lophuromys flavopunctatus Yellow-spotted Brush-furred Rat (Buff-spotted Brush-furred Rat) – F. Dieterlen 246 9

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Contents

Lophuromys huttereri Hutterer’s Brush-furred Rat – F. Dieterlen Lophuromys luteogaster Buff-bellied Brush-furred Rat (Hatt’s Brush-furred Rat) – F. Dieterlen Lophuromys medicaudatus Medium-tailed Brush-furred Rat (Western Rift Brush-furred Rat) – F. Dieterlen Lophuromys melanonyx Black-clawed Brush-furred Rat – D. W.Yalden Lophuromys nudicaudus Fire-bellied Brush-furred Rat – F. Dieterlen Lophuromys rahmi Rahm’s Brush-furred Rat – F. Dieterlen Lophuromys roseveari Rosevear’s Brush-furred Rat (Mount Cameroon Brush-furred Rat) – F. Dieterlen Lophuromys sikapusi Rusty-bellied Brush-furred Rat – F. Dieterlen Lophuromys woosnami Woosnam’s Brush-furred Rat – F. Dieterlen Genus Uranomys Rudd’s Brush-furred Mouse – D. C. D. Happold Uranomys ruddi Rudd’s Brush-furred Mouse – D. C. D. Happold Subfamily GERBILLINAE Gerbils and Jirds – M. D. Carleton & G. G. Musser

248 249 250 251 252 253 254 255 257 258 259 260

Genus Ammodillus Ammodile – D. C. D. Happold Ammodillus imbellis Ammodile (Walo) – D. C. D. Happold

262

Genus Desmodilliscus Brauer’s Dwarf Gerbil – L. Granjon Desmodilliscus braueri Brauer’s Dwarf Gerbil (Pouched Gerbil) – L. Granjon

264

Genus Desmodillus Cape Short-tailed Gerbil – J. A. J. Nel Desmodillus auricularis Cape Short-tailed Gerbil (Namaqua Gerbil) – J. A. J. Nel

266

Genus Gerbilliscus Gerbils – L. Granjon & E. R. Dempster Gerbilliscus afra Cape Gerbil – E. R. Dempster Gerbilliscus boehmi Boehm’s Gerbil – D. C. D. Happold Gerbilliscus brantsii Highveld Gerbil – E. R. Dempster Gerbilliscus gambianus Gambian Gerbil – J.-M. Duplantier & L. Granjon Gerbilliscus guineae Guinea Gerbil – L. Granjon & J.-M. Duplantier Gerbilliscus inclusus Gorongoza Gerbil – E. R. Dempster Gerbilliscus kempi Kemp’s Gerbil (Northern Savanna Gerbil) – D. C. D. Happold Gerbilliscus leucogaster Bushveld Gerbil – E. R. Dempster Gerbilliscus nigricaudus Black-tailed Gerbil – D. C. D. Happold Gerbilliscus phillipsi Phillips’s Gerbil – D. C. D. Happold Gerbilliscus robustus Fringe-tailed Gerbil – D. C. D. Happold Gerbilliscus validus Savanna Gerbil (Southern Savanna Gerbil) – D. C. D. Happold

268 270 272 273

263

264

267

274 276 277 278 279 281 283 284 285

Genus Gerbillurus Hairy-footed Gerbils – E. R. Dempster Gerbillurus paeba Pygmy Hairy-footed Gerbil (Paeba Hairyfooted Gerbil) – M. Perrin & E. R. Dempster Gerbillurus setzeri Setzer’s Hairy-footed Gerbil – M. Perrin & E. R. Dempster Gerbillurus tytonis Dune Hairy-footed Gerbil – M. Perrin & E. R. Dempster Gerbillurus vallinus Brush-tailed Hairy-footed Gerbil – M. Perrin & E. R. Dempster

287

Genus Gerbillus Gerbils – L. Granjon Gerbillus acticola Berbera Gerbil – D. C. D. Happold Gerbillus amoenus Charming Gerbil (Pleasant Gerbil) – D. C. D. Happold Gerbillus andersoni Anderson’s Gerbil – D. C. D. Happold Gerbillus bottai Botta’s Gerbil – D. C. D. Happold Gerbillus brockmani Brockman’s Gerbil – D. C. D. Happold Gerbillus campestris North African Gerbil – D. C. D. Happold Gerbillus cosensi Cosens’s Gerbil – D. C. D. Happold Gerbillus dasyurus Wagner’s Gerbil – D. C. D. Happold Gerbillus dunni Dunn’s Gerbil – D. C. D. Happold Gerbillus floweri Flower’s Gerbil – L. Granjon Gerbillus gerbillus Lesser Egyptian Gerbil (Small Egyptian Gerbil) – L. Granjon Gerbillus harwoodi Harwood’s Gerbil – D. C. D. Happold Gerbillus henleyi Henley’s Gerbil – L. Granjon Gerbillus hesperinus Moroccan Gerbil – D. C. D. Happold Gerbillus hoogstraali Hoogstraal’s Gerbil – D. C. D. Happold Gerbillus juliani Julian’s Gerbil – D. C. D. Happold Gerbillus latastei Lataste’s Gerbil – D. C. D. Happold Gerbillus lowei Lowe’s Gerbil – D. C. D. Happold Gerbillus mackilligini Mackilligin’s Gerbil – D. C. D. Happold Gerbillus maghrebi Maghreb Gerbil – S. Aulagnier Gerbillus nancillus Sudan Gerbil – L. Granjon Gerbillus nanus Dwarf Gerbil (Baluchistan Gerbil) – L. Granjon Gerbillus nigeriae Nigerian Gerbil – B. Sicard Gerbillus occiduus Occidental Gerbil – L. Granjon Gerbillus percivali Percival’s Gerbil – D. C. D. Happold Gerbillus perpallidus Pale Gerbil – D. C. D. Happold Gerbillus pulvinatus Cushioned Gerbil (Rhoad’s Gerbil) – D. C. D. Happold Gerbillus pusillus Least Gerbil – D. C. D. Happold Gerbillus pyramidum Greater Egyptian Gerbil – L. Granjon Gerbillus rosalinda Rosalind’s Gerbil (Rosalinda Gerbil) – D. C. D. Happold Gerbillus rupicola Rock Gerbil – L. Granjon Gerbillus simoni Simon’s Gerbil – D. C. D. Happold Gerbillus somalicus Somalian Gerbil – D. C. D. Happold Gerbillus stigmonyx Khartoum Gerbil – D. C. D. Happold Gerbillus tarabuli Tarabul’s Gerbil (Libyan Gerbil) – L. Granjon Gerbillus watersi Waters’s Gerbil – D. C. D. Happold

295 297

288 290 291 293

298 299 300 301 302 304 304 306 307 308 309 310 311 312 313 314 315 316 317 318 318 320 321 322 323 324 325 325 327 328 328 330 330 331 332

Genus Meriones Jirds – E. Fichet-Calvet 333 Meriones crassus Sundevall’s Jird (Silky Jird) – E. Fichet-Calvet 335 Meriones libycus Libyan Jird – E. Fichet-Calvet 336

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Contents

Meriones shawi Shaw’s Jird – E. Fichet-Calvet

338

Genus Microdillus Peel’s Pygmy Gerbil – D. C. D. Happold Microdillus peeli Peel’s Pygmy Gerbil (Somali Pygmy Gerbil) – D. C. D. Happold

339

Genus Pachyuromys Fat-tailed Jird – D. C. D. Happold Pachyuromys duprasi Fat-tailed Jird – D. C. D. Happold

341 341

Genus Psammomys Sand Rats – E. Fichet-Calvet Psammomys obesus Fat Sand Rat – E. Fichet-Calvet Psammomys vexillaris Pale Sand Rat (Lesser Sand Rat) – E. Fichet-Calvet 

343 344

Genus Sekeetamys Bushy-tailed Jird – C. Denys Sekeetamys calurus Bushy-tailed Jird – C. Denys

347 347

340

346

Genus Taterillus Taterils (Gerbils) – L. Granjon & G. Dobigny 349 Taterillus arenarius Sand Tateril (Robbins’s Tateril) – L. Granjon & G. Dobigny 350 Taterillus congicus Congo Tateril – L. Granjon & G. Dobigny 351 Taterillus emini Emin’s Tateril – L. Granjon & G. Dobigny 352 Taterillus gracilis Slender Tateril (Gracile Tateril) – L. Granjon & G. Dobigny 353 Taterillus lacustris Lake Chad Tateril – L. Granjon & G. Dobigny 354 Taterillus petteri Petter’s Tateril – Bruno Sicard 355 Taterillus pygargus Senegal Tateril (Cuvier’s Tateril) – L. Granjon & G. Dobigny 356 Taterillus tranieri Tranier’s Tateril – G. Dobigny & L. Granjon 357 Subfamily LEIMACOMYINAE Büttner’s Forest Mouse – G. G. Musser & M. D. Carleton Genus Leimacomys Büttner’s Forest Mouse – F. Dieterlen Leimacomys buettneri Büttner’s Forest Mouse (Büttner’s Togo Mouse) – F. Dieterlen Subfamily MURINAE Rats and Mice – G. G. Musser & M. D. Carleton

358 359 359 361

Genus Aethomys Veld Rats – C. T. Chimimba, A. V. Linzey & M. H. Kesner 362 Aethomys bocagei Bocage’s Veld Rat (Bocage’s Aethomys) – C. T. Chimimba, A. V. Linzey & M. H. Kesner 364 Aethomys chrysophilus Red Veld Rat (Red Aethomys) – A. V. Linzey, M. H. Kesner & C. T. Chimimba 365 Aethomys granti Grant’s Veld Rat (Grant’s Aethomys) – C. T. Chimimba, A. V. Linzey & M. H. Kesner 367 Aethomys hindei Hinde’s Veld Rat (Hinde’s Aethomys) – A.V. Linzey, C. T. Chimimba & M. H. Kesner 368 Aethomys ineptus Tete Veld Rat (Tete Aethomys) – A. V. Linzey, M. H. Kesner & C. T. Chimimba 369 Aethomys kaiseri Kaiser’s Veld Rat (Kaiser’s Aethomys) – A.V. Linzey, C. T. Chimimba & M. H. Kesner 370 Aethomys namaquensis Namaqua Veld Rat (Namaqua Aethomys) – M. H. Kesner, A. V. Linzey & C. T. Chimimba 371

Aethomys nyikae Nyika Veld Rat (Nyika Aethomys) – A. V. Linzey, C. T. Chimimba & M. H. Kesner Aethomys silindensis Selinda Veld Rat (Selinda Aethomys) – C. T. Chimimba, A. V. Linzey & M. H. Kesner Aethomys stannarius West African Veld Rat (West African Aethomys) – A. V. Linzey, C. T. Chimimba & M. H. Kesner Aethomys thomasi Thomas’s Veld Rat (Thomas’s Aethomys) – C. T. Chimimba, A. V. Linzey & M. H. Kesner

373 374 375 376

Genus Apodemus Field Mice – C. Denys Apodemus sylvaticus Long-tailed Field Mouse (Wood Mouse) – C. Denys

377

Genus Arvicanthis Grass Rats – L. Granjon & J.-F. Ducroz Arvicanthis abyssinicus Ethiopian Grass Rat (Abyssinian Grass Rat) – Afework Bekele Arvicanthis ansorgei Ansorge’s Grass Rat (Sudanian Arvicanthis) – L. Granjon & J.-F. Ducroz Arvicanthis blicki Blick’s Grass Rat – Afework Bekele Arvicanthis nairobae Nairobi Grass Rat – S. Takata Arvicanthis neumanni Neumann’s Grass Rat (Somali Grass Rat) – Afework Bekele Arvicanthis niloticus Nile Grass Rat (Unstriped Grass Rat) – L. Granjon, Afework Bekele & J.-F. Ducroz Arvicanthis rufinus Rufous Grass Rat (Guinean Grass Rat) – L. Granjon & J.-F. Ducroz

379

Genus Colomys African Water Rat – F. Dieterlen Colomys goslingi African Water Rat (Velvet Mouse, African Wading Rat) – F. Dieterlen

389

Genus Dasymys Shaggy Rats – M. D. Carleton Dasymys foxi Fox’s Shaggy Rat – M. D. Carleton Dasymys incomtus Common Shaggy Rat – N. Pillay Dasymys montanus Montane Shaggy Rat – M. D. Carleton Dasymys nudipes Angolan Shaggy Rat – M. D. Carleton Dasymys rufulus Rufous Shaggy Rat – J.-M. Duplantier

392 394 395 396 397 399

Genus Dephomys Defua Rat – D. C. D. Happold Dephomys defua Defua Rat – D. C. D. Happold

400 400

Genus Desmomys Scrub Rats – D. C. D Happold Desmomys harringtoni Harrington’s Scrub Rat (Harrington’s Desmomys) – Afework Bekele Desmomys yaldeni Yalden’s Scrub Rat (Yalden’s Desmomys) – D. W.Yalden

402

Genus Grammomys Thicket Rats – F. Dieterlen Grammomys aridulus Jebel Marra Thicket Rat (Arid Woodland Grammomys) – F. Dieterlen Grammomys buntingi Bunting’s Thicket Rat (Bunting’s Grammomys) – D. C. D. Happold Grammomys caniceps Grey-headed Thicket Rat (Greyheaded Grammomys) – R. Hutterer Grammomys cometes Mozambique Thicket Rat (Mozambique Grammomys) – F. Dieterlen

404

378

381 382 383 384 386 387 388

390

402 403

406 407 408 409 11

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Grammomys dolichurus Woodland Thicket Rat (Common Grammomys) – D. C. D. Happold Grammomys dryas Albertine Rift Thicket Rat (Albertine Rift Grammomys) – S. O. Bober & J. C. Kerbis Peterhans Grammomys gigas Giant Thicket Rat (Mount Kenya Grammomys) – F. Dieterlen Grammomys ibeanus East African Thicket Rat (East African Grammomys) – F. Dieterlen Grammomys kuru Shining Thicket Rat (Eastern Rainforest Grammomys) – D. C. D. Happold Grammomys macmillani Macmillan’s Thicket Rat (Macmillan’s Grammomys) – F. Dieterlen Grammomys minnae Ethiopian Thicket Rat (Ethiopian Grammomys) – D. W.Yalden Genus Heimyscus African Smoky Mouse – J. C. Ray & J. R. Malcolm Heimyscus fumosus African Smoky Mouse – J. C. Ray & J. R. Malcolm

410 411 412 413 414 416 417 418 418

Genus Hybomys Forest Mice – M. D. Carleton Hybomys badius Cameroon Forest Mouse (Cameroon Highland Hybomys) – R. Hutterer Hybomys basilii Basilio’s Forest Mouse (Bioko Hybomys) – R. Hutterer Hybomys lunaris Rwenzori Forest Mouse (Rwenzori Hybomys) – F. Dieterlen Hybomys planifrons Liberian Forest Mouse (Liberian Forest Hybomys) – M. D. Carleton Hybomys trivirgatus Three-striped Forest Mouse (West African Hybomys) – M. D. Carleton & D. C. D. Happold Hybomys univittatus One-striped Forest Mouse (Peters’s Hybomys) – J. C. Ray

420

Genus Hylomyscus Wood Mice – F. Dieterlen Hylomyscus aeta Beaded Wood Mouse (Beaded Hylomyscus) – J. C. Ray & J. R. Malcolm Hylomyscus alleni Allen’s Wood Mouse (Allen’s Hylomyscus) – J. C. Ray Hylomyscus baeri Baer’s Wood Mouse (Baer’s Hylomyscus) – D. C. D. Happold Hylomyscus carillus Angolan Wood Mouse (Angolan Hylomyscus) – D. C. D. Happold Hylomyscus denniae Montane Wood Mouse (Montane Hylomyscus) – F. Dieterlen Hylomyscus grandis Large Wood Mouse (Mount Oku Hylomyscus) – R. Hutterer Hylomyscus parvus Lesser Wood Mouse (Lesser Hylomyscus) – J.-M. Duplantier Hylomyscus stella Stella Wood Mouse (Stella Hylomyscus) – F. Dieterlen

429

Genus Lamottemys Mount Oku Rat – R. Hutterer Lamottemys okuensis Mount Oku Rat (Mount Oku Lamottemys) – R. Hutterer

439

Genus Lemniscomys Grass Mice – M. D. Carleton Lemniscomys barbarus Barbary Grass Mouse (Barbary Lemniscomys) – D. C. D. Happold Lemniscomys bellieri Bellier’s Grass Mouse (Bellier’s Lemniscomys) – E. Van der Straeten Lemniscomys griselda Griselda’s Grass Mouse (Griselda’s Lemniscomys) – A. Monadjem Lemniscomys hoogstraali Hoogstraal’s Grass Mouse (Hoogstraal’s Lemniscomys) – F. Dieterlen Lemniscomys linulus Senegal Grass Mouse (Senegal Lemniscomys) – E. Van der Straeten Lemniscomys macculus Buffoon Grass Mouse (Buffoon Lemniscomys) – F. Dieterlen Lemniscomys mittendorfi Mittendorf’s Grass Mouse (Mittendorf’s Lemniscomys) – E. Van der Straeten Lemniscomys rosalia Single-striped Grass Mouse (Singlestriped Lemniscomys) – A. Monadjem Lemniscomys roseveari Rosevear’s Grass Mouse (Rosevear’s Lemniscomys) – E. Van der Straeten Lemniscomys striatus Striated Grass Mouse (Striated Lemniscomys) – D. C. D. Happold Lemniscomys zebra Zebra Grass Mouse (Heuglin’s Lemniscomys) – D. C. D. Happold & F. Dieterlen

441

Genus Malacomys Swamp Rats – D. C. D. Happold Malacomys cansdalei Cansdale’s Swamp Rat (Cansdale’s Malacomys) – D. C. D. Happold Malacomys edwardsi Edwards’s Swamp Rat (Edwards’s Malacomys) – D. C. D. Happold Malacomys longipes Long-footed Swamp Rat (Common Malacomys) – D. C. D. Happold

455

Genus Mastomys Multimammate Mice – H. Leirs Mastomys awashensis Awash Multimammate Mouse (Awash Mastomys) – L. A. Lavrenchenko & H. Leirs Mastomys coucha Southern African Multimammate Mouse (Southern African Mastomys) – H. Leirs Mastomys erythroleucus Guinea Multimammate Mouse (Reddish-white Mastomys) – H. Leirs Mastomys huberti Hubert’s Multimammate Mouse (Hubert’s Mastomys) – H. Leirs Mastomys kollmannspergeri Kollmannsperger’s Multimammate Mouse (Kollmannsperger’s Mastomys) – H. Leirs Mastomys natalensis Natal Multimammate Mouse (Natal Mastomys) – H. Leirs Mastomys pernanus Dwarf Multimammate Mouse (Dwarf Mastomys) – H. Leirs Mastomys shortridgei Shortridge’s Multimammate Mouse (Shortridge’s Mastomys) – H. Leirs

460

Genus Muriculus Ethiopian Striped Mouse – D. W.Yalden Muriculus imberbis Ethiopian Striped Mouse – D. W.Yalden

472

443 444 445 446 447 448 449 449 451 452 454

422 423 423 424

456 457 458

425 427

430 431 433 433 434 436

462 463 464 465 467 468 470 471

437 438

440

Genus Mus Old World Mice and Pygmy Mice – D. C. D. Happold & F. Veyrunes Mus baoulei Baoule Pygmy Mouse – L. Granjon

472 473 476

12

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Mus bufo Toad Pygmy Mouse (Rwenzori Mouse) – F. Dieterlen Mus callewaerti Callewaert’s Pygmy Mouse – D. C. D. Happold Mus goundae Gounda River Pygmy Mouse – F. Petter Mus haussa Hausa Pygmy Mouse – F. Petter Mus indutus Desert Pygmy Mouse – A. Monadjem Mus mahomet Mahomet Pygmy Mouse – D. W.Yalden Mus mattheyi Matthey’s Pygmy Mouse – F. Petter Mus minutoides Tiny Pygmy Mouse – A. Monadjem Mus musculoides West African Pygmy Mouse – D. C. D. Happold Mus musculus House Mouse – D. C. D. Happold Mus neavei Neave’s Pygmy Mouse – F. Petter Mus orangiae Orange Pygmy Mouse – A. Monadjem Mus oubanguii Oubangui Pygmy Mouse – F. Petter Mus setulosus Peters’s Pygmy Mouse – D. C. D. Happold Mus setzeri Setzer’s Pygmy Mouse – A. Monadjem Mus sorella Sorella Pygmy Mouse (Thomas’s Pygmy Mouse) – F. Petter Mus spretus Algerian Mouse (Western Mediterranean Mouse) – D. C. D. Happold Mus tenellus Delicate Pygmy Mouse – F. Petter Mus triton Grey-bellied Pygmy Mouse – F. Dieterlen & D. C. D. Happold

477 478 479 480 481 482 483 484 486 487 489 490 491 492 493 494 495 496 497

Genus Mylomys Mill Rats (Three-toed Grass Rats) – F. Dieterlen 499 Mylomys dybowskii Dybowski’s Mill Rat (Dybowski’s Three-toed Grass Rat, Common Mylomys) – F. Dieterlen 500 Mylomys rex King Mill Rat (Ethiopian Three-toed Grass 501 Rat, Ethiopian Mylomys) – D. W.Yalden Genus Myomyscus Meadow Mice – D. C. D. Happold Myomyscus angolensis Angolan Meadow Mouse (Angolan Myomyscus) – H. Leirs Myomyscus brockmani Brockman’s Meadow Mouse (Smoky Meadow Mouse, Brockman’s Myomyscus) – D. C. D. Happold Myomyscus verreauxii Verreaux’s Meadow Mouse (Whitefooted Mouse) – D. C. D. Happold

502

Genus Nesokia Bandicoot Rats – D. C. D. Happold Nesokia indica Short-tailed Bandicoot Rat (Egyptian Pest Rat) – D. C. D. Happold

506

503 504 505

506

Genus Nilopegamys Ethiopian Water Rat – D. C. D. Happold 508 Nilopegamys plumbeus Ethiopian Water Rat – D. C. D. Happold 508 Genus Oenomys Rufous-nosed Rats – F. Dieterlen Oenomys hypoxanthus Common Rufous-nosed Rat (Common Oenomys) – F. Dieterlen Oenomys ornatus West African Rufous-nosed Rat (West African Oenomys) – F. Dieterlen

509

Genus Pelomys Creek Rats – F. Dieterlen

513

510 512

Pelomys campanae  Angolan Creek Rat (Angolan Pelomys) – F. Dieterlen Pelomys fallax  East African Creek Rat (East African Pelomys) – F. Dieterlen Pelomys hopkinsi  Hopkins’s Creek Rat (Hopkins’s Pelomys) – F. Dieterlen Pelomys isseli  Ssese Islands Creek Rat (Lake Victoria Pelomys) – F. Dieterlen Pelomys minor  Least Creek Rat (Least Pelomys) – F. Dieterlen Genus Praomys Soft-furred Mice – D. C. D. Happold Praomys daltoni  Dalton’s Soft-furred Mouse (Dalton’s Praomys) – L. Granjon Praomys degraaffi  De Graaff’s Soft-furred Mouse (De Graaff’s Praomys) – J. C. Kerbis Peterhans & S. O. Bober Praomys delectorum  Delicate Soft-furred Mouse (East African Praomys) – D. C. D. Happold Praomys derooi  De Roo’s Soft-furred Mouse (De Roo’s Praomys) – L. Granjon Praomys hartwigi  Hartwig’s Soft-furred Mouse (Hartwig’s Praomys) – R. Hutterer Praomys jacksoni  Jackson’s Soft-furred Mouse (Jackson’s Praomys) – F. Dieterlen Praomys lukolelae  Lukolela Soft-furred Mouse (Lukolela Praomys) – D. C. D. Happold Praomys minor  Least Soft-furred Mouse (Least Praomys) – E. Van der Straeten Praomys misonnei  Misonne’s Soft-furred Mouse (Misonne’s Praomys) – F. Dieterlen Praomys morio  Cameroon Soft-furred Mouse (Cameroon Praomys) – D. C. D. Happold Praomys mutoni  Riverine Soft-furred Mouse (Masako Softfurred Mouse, Riverine Praomys) – E. Van der Straeten Praomys obscurus  Obscure Soft-furred Mouse (Gotel Mountain Praomys) – R. Hutterer Praomys petteri  Petter’s Soft-furred Mouse (Petter’s Praomys) – E. Van der Straeten Praomys rostratus  West African Soft-furred Mouse (West African Praomys) – E. Van der Straeten Praomys tullbergi  Tullberg’s Soft-furred Mouse (Tullberg’s Praomys) – D. C. D. Happold Praomys verschureni  Verschuren’s Soft-furred Mouse (Verschuren’s Praomys) – F. Dieterlen Genus Rattus Rats – D. C. D. Happold Rattus norvegicus  Brown Rat (Norway Rat) – D. C. D. Happold Rattus rattus  Black Rat – D. C. D. Happold Genus Rhabdomys Four-striped Grass Mouse – D. C. D. Happold Rhabdomys pumilio  Four-striped Grass Mouse – D. C. D. Happold Genus Stenocephalemys Ethiopian Rats – D. W.Yalden Stenocephalemys albipes  White-footed Ethiopian Rat (White-footed Stenocephalemys) – Afework Bekele

514 515 517 518 519 519 522 523 524 526 527 527 529 530 531 532 533 534 535 536 537 538 539 540 541 544 545 547 549 13

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Stenocephalemys albocaudata  White-tailed Ethiopian Rat (White-tailed Stenocephalemys) – D. W.Yalden Stenocephalemys griseicauda  Grey-tailed Ethiopian Rat (Grey-tailed Stenocephalemys) – D. W.Yalden Stenocephalemys ruppi  Rupp’s Ethiopian Rat (Gughe Highlands Rat, Rupp’s Stenocephalemys) – D. W.Yalden

550 552

554 554

Genus Thallomys Acacia Rats (Tree Rats) – M. Perrin Thallomys loringi  Loring’s Acacia Rat (Loring’s Thallomys) – M. D. Carleton Thallomys nigricauda  Black-tailed Acacia Rat (Black-tailed Thallomys) – J. A. J. Nel Thallomys paedulcus  Sundevall’s Acacia Rat (Acacia Thallomys) – M. Perrin Thallomys shortridgei  Shortridge’s Acacia Rat (Shortridge’s Thallomys) – J. A. J. Nel

556

GENUS Thamnomys Thicket Rats – F. Dieterlen Thamnomys kempi  Kemp’s Thicket Rat (Kemp’s Thamnomys) – F. Dieterlen Thamnomys schoutedeni  Schouteden’s Thicket Rat (Schouteden’s Thallomys) – F. Dieterlen Thamnomys venustus  Charming Thicket Rat (Thomas’s Thamnomys) – F. Dieterlen

563

GENUS Zelotomys Broad-headed Mice – J. A. J. Nel Zelotomys hildegardeae  Hildegarde’s Broad-headed Mouse (Hildegarde’s Zelotomys) – J. A. J. Nel Zelotomys woosnami  Woosnam’s Broad-headed Mouse (Woosnam’s Zelotomys) – J. A. J. Nel

567

GENUS Otomys Vlei Rats – P. J. Taylor Otomys anchietae  Anchieta’s Vlei Rat (Angolan Vlei Rat) – P. J. Taylor Otomys angoniensis  Angoni Vlei Rat – P. J. Taylor Otomys barbouri  Barbour’s Vlei Rat – V. Clausnitzer Otomys burtoni  Burton’s Vlei Rat – P. J. Taylor Otomys cuanzensis  Cuanza Vlei Rat – P. J. Taylor Otomys denti  Dent’s Vlei Rat – P. J. Taylor Otomys irroratus  Southern African Vlei Rat – P. J. Taylor Otomys lacustris  Lake Vlei Rat – V. Clausnitzer Otomys laminatus  Laminate Vlei Rat (KwaZulu Vlei Rat) – P. J. Taylor Otomys occidentalis  Western Vlei Rat – P. J. Taylor Otomys saundersiae  Saunders’s Vlei Rat – P. J. Taylor Otomys sloggetti  Sloggett’s Vlei Rat – P. J. Taylor Otomys tropicalis  Tropical Vlei Rat (East African Vlei Rat) – P. J. Taylor Otomys typus  Ethiopian Vlei Rat – D. W.Yalden

594

GENUS Parotomys Whistling Rats – T. P. Jackson Parotomys brantsii  Brants’s Whistling Rat – T. P. Jackson Parotomys littledalei  Littledale’s Whistling Rat – T. P. Jackson

596 597 600

Family ANOMALURIDAE Anomalures – J. Kingdon

602

553

Genus Stochomys Target Rat – D. C. D. Happold Stochomys longicaudatus  Target Rat – D. C. D. Happold

Subfamily OTOMYINAE Vlei Rats and Whistling Rats – M. D. Carleton

Otomys unisulcatus  Karoo Vlei Rat (Bush Karoo Rat) – T. P. Jackson

558 559 561

GENUS Anomalurus Anomalures (Scaly-tailed Squirrels) – J. Kingdon Anomalurus beecrofti  Beecroft’s Anomalure (Beecroft’s Scaly-tailed Squirrel) – J. Kingdon Anomalurus derbianus  Lord Derby’s Anomalure (Lord Derby’s Scaly-tailed Squirrel) – J. C. Ray Anomalurus pelii  Pel’s Anomalure (Pel’s Scaly-tailed Squirrel) – J. Kingdon Anomalurus pusillus  Lesser Anomalure (Dwarf Scaly-tailed Squirrel) – J. Kingdon

603 604 606 608 610

562

564 565

GENUS Idiurus Pygmy Anomalures – R. W. Thorington, Jr Idiurus macrotis  Long-eared Pygmy Anomalure (Longeared Scaly-tailed Flying Squirrel) – B. J. Stafford & R. W. Thorington, Jr Idiurus zenkeri  Zenker’s PygmyAnomalure (Pygmy Scaly-tailed Flying Squirrel) – B. J. Stafford & R. W. Thorington, Jr

611 612 614

566

568

GENUS Zenkerella Cameroon Anomalure – D. C. D. Happold Zenkerella insignis  Cameroon Anomalure (Cameroon Scaly-tail) – B. J. Stafford & R. W. Thorington, Jr

569

Family PEDETIDAE Springhares – T. M. Butynski

618

GENUS Pedetes Springhares – T. M. Butynski Pedetes capensis  Southern African Springhare – T. M. Butynski Pedetes surdaster  East African Springhare – T. M. Butynski & J. Kalina

618

571 574 576 577 579 580 581 582 583 585 586 587 588 589 591 592

Family CTENODACTYLIDAE Gundis and Pectinator – P. Gouat

615 616

619 624 628

GENUS Ctenodactylus Gundis – P. Gouat Ctenodactylus gundi  Atlas Gundi (Common Gundi) – P. Gouat Ctenodactylus vali  Thomas’s Gundi – P. Gouat

629

GENUS Felovia Felou Gundi – P. Gouat Felovia vae  Felou Gundi – P. Gouat

634 635

GENUS Massoutiera Mzab Gundi – P. Gouat Massoutiera mzabi  Mzab Gundi – P. Gouat

636 636

GENUS Pectinator Speke’s Pectinator – P. Gouat Pectinator spekei  Speke’s Pectinator – P. Gouat

638 638

630 632

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Family BATHYERGIDAE Mole-rats – J. U. M. Jarvis

641

GENUS Bathyergus Dune Mole-rats – J. U. M. Jarvis 644 Bathyergus janetta  Namaqua Dune Mole-rat – J. U. M. Jarvis 644 Bathyergus suillus  Cape Dune Mole-rat – J. U. M. Jarvis 646 GENUS Cryptomys Mole-rats – N. C. Bennett Cryptomys anselli  Ansell’s Mole-rat – N. C. Bennett & H. Burda Cryptomys bocagei  Bocage’s Mole-rat – N. C. Bennett Cryptomys damarensis  Damaraland Mole-rat – N. C. Bennett Cryptomys darlingi  Darling’s Mole-rat (Mashona Mole-rat) – N. C. Bennett Cryptomys foxi  Fox’s Mole-rat – N. C. Bennett Cryptomys hottentotus  Common Mole-rat (Hottentot Mole-rat) – N. C. Bennett Cryptomys kafuensis  Kafue Mole-rat – N. C. Bennett & H. Burda Cryptomys mechowi  Giant Mole-rat – N. C. Bennett & H. Burda Cryptomys ochraceocinereus  Ochre Mole-rat – N. C. Bennett Cryptomys zechi  Togo Mole-rat – N. C. Bennett

648

GENUS Georychus Cape Mole-rat – N. C. Bennett Georychus capensis  Cape Mole-rat (Blesmol) – N. C. Bennett

662

GENUS Heliophobius Silvery Mole-rat – J. U. M. Jarvis Heliophobius argenteocinereus  Silvery Mole-rat – J. U. M. Jarvis

664

GENUS Heterocephalus Naked Mole-rat – J. U. M. Jarvis Heterocephalus glaber  Naked Mole-rat – J. U. M. Jarvis

667 668

Family HYSTRICIDAE Porcupines – D. C. D. Happold

671

649 650 651 653 654

Family MYOCASTORIDAE Coypu – D. C. D. Happold

691

Genus Myocastor Coypu691 Myocastor coypus  Coypu (Nutria) – D. C. D. Happold 691 Order LAGOMORPHA Hares, Rock-hares, Rabbits and Pikas – D. C. D. Happold

693

Family LEPORIDAE Hares, Rock-hares and Rabbits – D. C. D. Happold

694 696

Genus Lepus Hares – D. C. D. Happold Lepus capensis  Cape Hare – D. C. D. Happold Lepus fagani  Ethiopian Hare – D. C. D. Happold Lepus habessinicus  Abyssinian Hare – D. C. D. Happold Lepus saxatilis  Scrub Hare – D. C. D. Happold Lepus starcki  Ethiopian Highland Hare (Starck’s Hare) – D. C. D. Happold Lepus victoriae  African Savanna Hare – D. C. D. Happold

698 699 701 702 703

Genus Oryctolagus European Rabbit – D. C. D. Happold Oryctolagus cuniculus  European Rabbit – D. C. D. Happold

708 708

Genus Poelagus Bunyoro Rabbit – D. C. D. Happold Poelagus marjorita  Bunyoro Rabbit (Uganda Grass Hare) – D. C. D. Happold

710

712

676

Genus Pronolagus Rock-hares – D. C. D. Happold Pronolagus crassicaudatus  Natal Red Rock-hare – D. C. D. Happold Pronolagus randensis  Jameson’s Red Rock-hare – D. C. D. Happold Pronolagus rupestris  Smith’s Red Rock-hare – D. C. D. Happold Pronolagus saundersiae  Hewitt’s Red Rock-hare – D. C. D. Happold

678

Appendix: New Taxa 2005–2010

718

Glossary

719

Bibliography

731

Authors of Volume III

774

Indexes French names German names English names Scientific names

777 778 780 782

696

658 659 660 661

663

705 706

665

674

Family PETROMURIDAE Noki (Dassie Rat) – C. G. Coetzee

680

Genus Petromus Noki (Dassie Rat) – C. G. Coetzee Petromus typicus  Noki (Dassie Rat) – C. G. Coetzee

681 681

Family THRYONOMYIDAE Cane Rats – D. C. D. Happold 685 Genus Thryonomys Cane Rats – D. C. D. Happold Thryonomys gregorianus  Lesser Cane Rat (Lesser Grass Cutter) – D. C. D. Happold

688

Genus Bunolagus Riverine Rabbit – D. C. D. Happold Bunolagus monticularis  Riverine Rabbit – D. C. D. Happold

655

Genus Atherurus Brush-tailed Porcupines – D. C. D. Happold 672 Atherurus africanus  African Brush-tailed Porcupine – D. C. D. Happold 672 Genus Hystrix Crested Porcupines – D. C. D. Happold Hystrix africaeaustralis  Cape Crested Porcupine (Cape Porcupine) – D. C. D. Happold Hystrix cristata  North African Crested Porcupine (Crested Porcupine) – D. C. D. Happold

Thryonomys swinderianus  Greater Cane Rat (Cutting-grass, Grass Cutter) – D. C. D. Happold

686 687

710

713 714 715 717

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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.

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.

above left:

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

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Acknowledgements for Volume III D. C. D. Happold

I owe a great debt of gratitude to the many people who have helped and encouraged me with my research on African small mammals for nearly 50 years and hence have contributed enormously to this volume. My interest in rodents (and other small mammals) began when I spent three months in 1959 at the ‘Institut pour la Recherche Scientifique d’Afrique Centrale’ in the then Belgian Congo; here I met Urs Rahm whose enthusiasm for small mammals has been an inspiration ever since. When I was beginning my research, the following colleagues were a marvellous source of inspiration: Frank Ansell †, Mike Delany, Fritz Dieterlen, Harry Hoogstraal †, Dieter Kock, Waldo Meester †, Xavier Misonne †, Francis Petter †, Don Rosevear †, Hank Setzer † and Reay Smithers †. Since then, other colleagues and friends have given advice, sent books and publications, and encouraged me in my studies. I also acknowledge the many collectors who, from the nineteenth century onwards, collected specimens, often in remote parts of Africa and often under appallingly difficult conditions, and sent them back to museums in Britain, Europe and North America where they still provide invaluable information. I am also most grateful to my colleagues (many of whom are authors of profiles in this volume) who have given very generously of their time and knowledge to make this volume possible. [† Deceased] This volume owes a great deal to the museums that house specimens of African small mammals. For many years, I have visited museums whenever the opportunity has occurred, and I am most grateful to the curators and their assistants who have helped me during my visits: David Harrison, Paul Bates and Malcolm Pearch (Harrison Zoological Institute, Sevenoaks, England), Francis Petter and Jacques Cuisin (Museum National d’Histoire Naturelle, Paris, France), Lydia Kigo (National Museum of Kenya, Nairobi, Kenya), Wim Wendelen (Royal Museum of Central Africa, Tervuren, Belgium), Judith Eger (Royal Ontario Museum, Toronto, Canada), Dieter Kock (Senckenberg Museum, Frankfurt, Germany), Teresa Kearney and Christian Chimimba (Transvaal Museum, Pretoria, South Africa) and Rainer Hutterer (Museum Alexander Koenig, Bonn, Germany). I owe a particular debt of gratitude to the Natural History Museum, London, England (formerly British Museum of Natural History), especially Gordon Corbet, Robert Hayman †, John Edwards Hill †, Paula Jenkins, Jean Ingles, Daphne Hills and Louise Tomsett who have allowed me (since 1964) to examine the collections and have assisted me in numerous ways. Particular thanks are due to Michael and Mona Ensor and Sara Churchfield in London, the Ostermeyer and Schlöder families in Bonn, our friends in Corniche Verte in Brussels, and Mike

and Anne-Marie Swift in Nairobi, who provided welcoming homes and sustenance after busy days in museums. I wish to thank the various universities where I have been employed, for their financial (and other) support for my studies on African mammals: Australian National University, Canberra, Australia; university of Ibadan, Nigeria; University of Khartoum, Sudan; and University of Malawi, Zomba, Malawi. The Australian Research Grants Committee also provided funds for my more recent work in Malawi. Very special thanks are due to my wife Meredith who has assisted me as research assistant, collaborator, co-author and reviewer; in fact, a lot of our research would not have been accomplished without her enthusiasm and encouragement. This volume would not have been possible without the contributions and invaluable collaboration from the authors themselves. Many authors had other commitments and found it extremely hard to find the time for writing profiles. Great credit is due to such authors, and to those who wrote numerous profiles, for their dedication and perseverance. My thanks are also due to the many reviewers (listed here in alphabetical order) who reviewed profiles and gave specialist advice for this volume: Corne Anderson, Dick Ashford, Paul Bates, Afework Bekele, Nigel Bennett, Mike Carleton, Christian Chimimba, Viola Clausnitzer, Neels Coetzee, Norbert Cordeiro, Marco Corti, José Crawford-Cabral, Fritz Dieterlen, Colleen Downs, Jean-Marc Duplantier, Louise Emmons, Chris G. Faulkes, Trish Fleming, Laurent Granjon, Mike J. Griffin, Peter Grubb, Barbara Herzig-Straschil, Tim Jackson, Jenny Jarvis, Graham Kerley, Michael Knight, Dieter Kock, Leonid Lavrenchenko, Emile Lecompte, Herwig Leirs, Jay Malcolm, Sarita Maree, Andrea Mess, Ara Monadjem, Guy Musser, Jan Nel, Dean Peinke, Mike Perrin, Francis Petter, Victor Rambau, Galen Rathbun, Brian Robbins, Lynn Robbins, Carsten Schradin, Anje Schunke, Claudio Sillero-Zubiri, William Stanley, Radim Sumbera, Peter Taylor, Michel Thèvenot, Michel Tranier, Erik Van der Straeten, Frederic Veyrunes, Derek Yalden and Truman Young. Anke Hoffman kindly provided most of the German names for the small mammals in this volume. Finally, I wish to express my thanks to the coordinating team of the Global Mammal Assessment, an initiative of the International Union for the Conservation of Nature (IUCN) and The Zoological Society of London who organized the workshops referred to in the General Acknowledgements. Also, I am most grateful to Elaine Leek who did a magnificent job copy-editing the text of this volume, and to Zoe Cokeliss who digitized all the distribution maps of the small mammals supplied by the authors. 17

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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 serves 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, seldom seen, 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 karyotypic and molecular/genetic characters of species. 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 previously unpublished information. 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

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The continent of Africa

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 because 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, etc.). Thus, 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, tribes (e.g. in Bovidae), subgenera (e.g. in Procolobus) and species-groups, or ‘superspecies’ (e.g. in Cercopithecus). 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, which, 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. Maps showing the political boundaries of Africa (Figure 1a), the Province of South Africa (Figure 1b) and the physical features of Africa with the major rivers and lakes (Figure 1c) are provided, as well as a list of the 47 countries including 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 six 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, 9,000 mi²], Swaziland [17,400 km², 6,700 mi²] and The Gambia [11,300 km², 4,400 mi²]). The human population density 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 divided 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. Thirteen biotic zones are recognized, two of which may be divided into smaller categories. The biotic zones in which each species of mammal has been recorded are listed in each profile for several reasons. They indicate the environmental conditions in which the species lives and they provide data with which the geographic distribution can 19

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An Introduction and Guide

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

20

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The continent of Africa

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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An Introduction and Guide

The rodents, hares and rabbits of Africa

1

This volume, Volume III, is devoted to the orders Rodentia and Lagomorpha. The order Rodentia contains more species (395 spp.) than any other order of African mammals and comprises about 36% of African mammals; in contrast, the order Lagomorpha contains relatively few species (13 spp.). These two orders, together with two orders (Afrosoricida and Macroscelidea) in Volume I and three orders (Erinaceomorpha, Soricomorpha and Chiroptera) in Volume IV, constitute the ‘small mammals of Africa’ and collectively comprise 74.1% (827 of 1116) of all African mammalian species. Many of these small species have not been studied in detail because of their rarity and small geographic ranges. However, some of the more common and widespread species are quite well known, especially with respect to their importance within their communities and because of their importance as ‘bushmeat’ and as hosts for diseases of humans and livestock. The profiles for Volume III were submitted to the editor between 2001 and 2005. It has not been possible to revise profiles since then except for a few details such as the completion of citations for papers previously given as ‘in press’, alterations (if necessary) to the category of threat in the conservation section of species profiles, and where there have been changes in taxonomy and distribution. An Appendix has been added listing new taxa of African rodents described during the period 2005–2010 and which have not been profiled in this volume.

2

3 6a

4

5

7

5

6

6a

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

8 6

6b

6c 10 9

11a 11b

12 11c 13

Figure 2. The biotic zones of Africa.

be explained and predicted. Furthermore, the number of biotic zones exploited by a species indicates its level of habitat tolerance and the extent to which it is vulnerable to loss of a particular type of habitat. The Rainforest Biotic Zone (Figure 3) and the South-West Arid Biotic Zone are divided into regions and sub-regions that reflect the different biogeographical distributions of species, each region/ sub-region having a community of mammals and other animals that is different to any other. Details of the Biotic Zones of Africa, and the regions and sub-regions of the Rainforest Biotic Zone and the SouthWest Arid Biotic Zone, are given in Volume I of Mammals of Africa.

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 and other information is as follows: Scientific name (genus and species)  The currently accepted name of the species.

ge Ni r

WEST CENTRAL Ben

Niger Delta

Gabon

Eastern Nigeria ga

a San

Gabon

Ubangi

Ghanaian

Ivory Coast

ue

Western Nigeria Cross

ra

Western

lta Vo

Sassand

Liberian WESTERN

EAST CENTRAL

i

go

n Co

0

500

1000

1500

2000 km

South Central

laba Lua

am Lom

SOUTH CENTRAL

East Central

Figure 3. The Rainforest Biotic Zone showing the regions, sub-regions and refugia. Regions are indicated in capital letters and colours: Western region – green; West Central region – brown; East Central region – purple; South Central region – blue. Sub-regions are indicated in lower case letters. Refugia are indicated in lower case italics (after Happold 1996 and references therein; see also Happold & Lock, Volume I, Mammals of Africa).

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Species profiles

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. In Volume III, French names were either provided by authors, or taken from Gunther (2002); most of the German names were provided by Anke Hoffman.

Description  This section, together with the illustrations and relevant tables, provides sufficient information to identify the species as well as describing characters that are relevant to the habits and lifestyle of 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 characters 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 are given. The characters described in this section are common to all subspecies (if any) of the species (see also Geographic Variation). It was not possible – or desirable – to describe the same suite of characters for every species. Instead, an appropriate selection was made for each family and/or genus, and therefore the same suite of characters is described for all members of the relevant taxon. Consequently the descriptions of related species are comparable and compatible. Characters that are diagnostic to the genus are not necessarily repeated in a species profile; hence higher taxa profiles should also be consulted. The number and arrangement of nipples in adult females is noted wherever this feature varies between the taxa being discussed (see Glossary). The Tables allow easy comparison between taxa within a genus or family.

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 the one it is placed in now. The scientific name is followed by the publication in which the species was described, and the type locality (i.e. where the type specimen [or type series] was obtained). Most of this information is taken from Wilson & Reeder (2005). Taxonomy  This section contains information about previous scientific names of the species, and problems and controversies (if any) associated with its nomenclature and relationships with other species. For some species, there is considerable information about these topics; for others, there may be nothing. Synonyms are listed in alphabetical order (without the taxonomic authority for each unless essential for clarity) and the number of subspecies (if any) is presented; most of this information is 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. In late 2006, a revised edition of the Atlas of Mammalian Chromosomes was published (O’Brien et al. 2006), but it has not been possible to incorporate the findings of that important work here.

postorbital patch forehead

crown

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 the characters that distinguish it from other subspecies of the species. 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

auditory meatus

mid-dorsal stripe

ear-fold

inner side-stripe outer side-stripe

postauricular patch

supraorbital patch eye-ring

2 rump

vibrissa preorbital patch

1

3 4 5

dorsal region plantar pads (in black)

neck

nose mussle lip (upper)

flank

chin suborbital patch cheek throat

ventral region

chest subauricular patch

hindlimb

sole of left hindfoot

shoulder axilary region tail

forelimb forefoot

claw

oot

hindf

digit

External characters of a hypothetical rodent. No species has this particular combination of characters.

pencil

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An Introduction and Guide

External characters of a hypothetical lagomorph.

tip of ear inner surface of ear outer surface of ear

inner margin of ear

outer margin of ear

crown eye cheek

eye-ring

nuchal patch

nasal region

lip (upper)

dorsal region

neck chin gular patch (on throat)

p rum

flank

chest ventral region

tail

forelimb hindlimb forefoot

hindfoot sole of hindfoot

refer to profiles of the similar species in question). In some instances, similar species that are allopatric in distribution are also included. Distribution  The first sentence, ‘Endemic to Africa’, if given, indicates that this is an African species and does not occur in the wild on any other continent; if a species also occurs outside Africa (and hence is not endemic), this is noted at the end of this section. The next sentence gives the Biotic Zone (or Zones) in which the species has been recorded because this information indicates the sorts of environments exploited by that species, its tolerance of different environmental conditions, and the extent to which it is likely to be threatened by habitat change. Also, it is the basis for predictions of its distribution outside the currently known limits. Finally, the countries (or parts of countries) where the species has been recorded are listed; sometimes other data (such as altitudinal range and habitat) are also included. As a general rule, descriptions of the geographic range for species with very restricted distributions are more precise in terms of information given (including, for example, geographic coordinates) than for more widespread species, where a more generalized statement is adequate. A distribution map (see below) augments the information given here. Habitat  This section provides a description of the habitat, or 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 variations in habitat characteristics. Abundance  This section attempts to indicate the comparative abundance of the species in its habitat. For many species, quantitative data are unavailable but the species can be assessed as ‘abundant’,

‘common’, ‘uncommon’, ‘rare’, ‘rarely seen but frequently heard’, ‘rarely collected’ etc. For some species, abundance may be expressed as estimates of density (e.g. number/ha or number/km2), or relative abundance within the community (e.g. ‘comprised 40% of small mammals captured’, ‘the second most numerous species’). For the better-known or rare species, actual numbers of individuals for the species may be given. 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 that show how the species uniquely interacts with its environment, with conspecifics, and with other animals. This section may also describe species-specific adaptations for feeding, locomotion, burrowing, domiciles, 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 emphasized. Foraging and Food  The first sentence briefly describes the diet of the species (e.g. insectivorous, carnivorous, granivorous, etc.). This may be followed by the methods of collecting food (foraging), size of home-range and daily distance moved, and descriptions of feeding behaviour.The diet, if well-known, is then described in one or more of the following ways: a list of the taxa of animals or plants consumed, a quantitative measure based on direct observations, or by a qualitative or quantitative analysis of the stomach contents or faeces. Social and Reproductive Behaviour  Topics in this section may include social organizations (e.g. solitary, social, or colonial), group

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Species profiles

size and composition; agonistic and amicable behaviour, territoriality and home-range (including quantitative data), courtship and mating, parental behaviour, and parent–young interactions, cooperative breeding, social 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 chronologies, if known, give data for both individuals and local populations. 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, known parasites (usually ectoparasites only) and some diseases are listed. Additional information is given if the species is a host to diseases that affect humans and domestic stock, and if it is utilized as food for humans (‘bushmeat’). Remarks  This subheading subsumes five of the above subheadings (Adaptations, Foraging and Food, Social and Reproductive Behaviour, Reproduction and Population Structure, and Predators, Parasites and Diseases) in those instances where there is little or no information available.

Conservation  The conservation status of the species (i.e. its IUCN Category) is taken from the ‘Red List of Threatened Species’ prepared by the International Union for Conservation of Nature (IUCN). The IUCN Red List Categories follow the definitions given in the IUCN Red List Categories and Criteria Version 3.1 (see www.iucnredlist.org) and are listed in Table 4. For those species that are classified as threatened (i.e. ‘Vulnerable’, ‘Endangered’ and ‘Critically Endangered’), readers may obtain detailed reasons (the criteria) for the classification on the IUCN Red List website. The status of some species has been changed in recent years because of improved knowledge, changes in taxonomy, or the impact of threatening processes or conservation action; detailed reasons for the present status, and past status, are given on the IUCN Red List website. If a species is listed on an Appendix I or Appendix II under CITES (Convention on International Trade in Endangered Species; www.cites.org), 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 abbreviation and definition for each measurement is 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. For most measurements, data for males and females are combined but where there is sexual dimorphism, measurements for males and females are given separately. Where possible, measurements also detail 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

Table 4.  Definitions for the IUCN Red List categories (from IUCN – Red List Categories, Version 3.1). 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)

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An Introduction and Guide

and measured are given in Table 5. Most museum records have been provided by the author of the profile; others – when an author did not have the measurements or did not have the opportunity to visit museums – were provided by the editor. Key References  A select list of references, which provides more general information on the species. Each reference is given in full in the Bibliography. 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  For selected taxa (mainly families and genera) tables provide details of the main characteristics of these taxa and can be used as an aid to identification. The tables were prepared by the editor.

Higher taxon profiles The profiles for orders, families and genera are less structured than for the species. 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

Table 5.  Acronyms for museum and private collections. Acronym

Museum name

ACBG

Museo di Anatomia Comparata, Università di Roma ‘La Sapienza’, Rome, Italy. Amatole Museum, King William’s Town, South Africa [formerly Kaffrarian Museum]. American Museum of Natural History, New York, USA. Natural History Museum, London, UK [formerly British Museum (Natural History)]. Carnegie Museum of Natural History, Pittsburgh, USA. Durban Natural Science Museum, Durban, South Africa. Field Museum of Natural History, Chicago, USA. Happold Collection, Canberra, Australia (private collection). Harrison Zoological Museum, Sevenoaks, Kent, UK. Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium. Los Angeles County Museum, Los Angeles, USA. McGregor Museum, Kimberley, South Africa. Museum National d’Histoire Naturelle, Paris, France. Museo Zoologico de ‘la Specola’, Università di Firenze, Italy. National Museums of Kenya, Nairobi, Kenya. National Museum of Namibia, Windhoek, Namibia. Naturhistorisches Museum, Wien (Vienna), Austria. Royal Museum for Central Africa, Tervuren, Belgium. South African Museum, Cape Town, South Africa. Senckenberg Museum, Frankfurt, Germany. Staatliches Museum für Naturkunde, Stuttgart, Germany. Transvaal Museum, Pretoria, South Africa. United States National Museum of Natural History, Smithsonian Institution, Washington, USA. Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany. Zoological Museum, Moscow University, Moscow, Russia.

AM AMNH BMNH CM DM FMNH HC HZM IRSN LACM MM MNHN MZUF NMK NMN NMW RMCA SAM SMF SMNS TM USNM ZFMK ZMMU

taxon has only one lower taxon. Higher taxa profiles provide the characteristics common to all members of that taxon. Some of these characteristics may not be repeated in lower taxon profiles (unless essential for identification) so readers are encouraged to consult also the next higher taxon profile, e.g. the species profile for Lophuromys sikapusi should be consulted in association with the genus Lophuromys profile.

Distribution maps Each species profile, with very few exceptions, contains a panAfrican map showing the geographic range of the species. Most maps were provided by the author of the profile and were compiled from literature records and museum specimens; some maps were provided by the editor(s) when it was not possible for the author to do so. 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(s). When presented, different colour shading denotes subspecies. • × = isolated locations considered to be separate from the main geographic range(s). Some locations indicated by × may include two or more closely spaced locations. • ? = locality of uncertain validity; relevant information usually in text. • red arrow = present on 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 RODENTIA

Order RODENTIA – Rodents Rodentia Bowdich, 1821.

Sciuridae (10 genera, 37 species)* Gliridae (2 genera, 16 species) Dipodidae (2 genera, 3 species) Spalacidae (2 genera, 3 species) Nesomyidae (12 genera, 34 species)

Cricetidae (2 genera, 2 species) Muridae (50 genera, 264 species)** Anomaluridae (3 genera, 7 species) Pedetidae (1 genus, 2 species) Ctenodactylidae (4 genera, 5 species) Bathyergidae (5 genera, 15 species) Hystricidae (2 genera, 3 species) Petromuridae (1 genus, 1 species) Thryonomyidae (1 genus, 2 species) Myocastoridae (1 genus, 1 species)†

Squirrels Dormice Jerboas Mole-rats, African Root-rats Pouched Rats and Mice, Swamp Mouse, Climbing Mice, Large-eared Mice, Fat Mice, White-tailed Rat, Rock Mice Voles, Maned Rat Gerbils and Jirds, Old World Rats and Mice, Vlei Rats Anomalures

p. 38 p. 102 p. 135 p. 143

p. 602

Springhares Gundis

p. 618 p. 628

Mole-rats Porcupines Noki (Dassie Rat) Cane Rats Coypu

p. 641 p. 671 p. 680 p. 685 p. 691

p. 153

p. 210 p. 216

* Including Sciurus carolinensis introduced recently into South Africa. ** Including Mus musculus, Rattus norvegicus and Rattus rattus introduced in recent historical times. † Introduced recently into Kenya.

Distribution and diversity Rodentia is the largest order of Mammalia, encompassing 2,277 of 5,422 living mammal species or approximately 42% of worldwide mammalian biodiversity (Wilson & Reeder 2005); of these 2,277 rodent species, 395 species (in 15 families and 98 genera) occur in Africa (see above and Table 1). Rodents are indigenous to every continent except Antarctica and inhabit most small to large landbridge and oceanic islands (originally absent from Iceland, New Zealand and some oceanic and high Arctic islands). The order’s global presence has become nearly ubiquitous as certain commensal species (see subfamily Murinae profile) have expanded their ranges in concert with human peregrinations during the Holocene. Large islands, island archipelagos and those continents with a geologic history of periodic isolation, have afforded ideal geographic settings for in situ radiations of major clades of Rodentia. Nowhere, with the arguable exception of South America, is this cladogenetic exuberance of the order so spectacularly showcased as in Africa, where 14 of the 33 recognized rodent families (sensu Wilson & Reeder 2005) are indigenous; a 15th family, Myocastoridae, has been introduced into Africa (Table 6). Seven of these 15 families are endemic to the continent (Table 6); at the generic and specific levels, the extent of endemism is substantially greater and highlights the contribution of African landscapes to the phylogenetic diversification of these rodent families (Table 1). Most non-endemic genera and species involve North African taxa that range into the nearby Arabian Peninsula and

Table 6. The 15 families of recent African Rodentia, with the number of constituent genera and species. Percentage endemism is based on the geographic limits of the African continent as defined for this volume and on the taxa recognized herein. Family Sciuridae Gliridae Dipodidae Spalacidae Nesomyidae Cricetidae Muridae Anomaluridae Pedetidae Ctenodactylidae Bathyergidae Hystricidae Petromuridae Thryonomyidae Myocastoridae Totals % endemics a

Endemic No No No No Yesa No No Yes Yes Yes Yes No Yes Yes Nob 15 47%

No. of genera 10 2 2 2 12 2 50 3 1 4 5 2 1 1 1 98

No. of endemic genera 9 1 0 1 12 1 41 3 1 4 5 0 1 1 0 80 82%

No. of species 37 16 3 3 34 2 264 7 2 5 15 3 1 2 1 395

No. of endemic species 36 16 1 2 34 2 250 7 2 5 15 2 1 2 0 375 94.9%

Excluding the Malagasy subfamily Nesomyinae. b Introduced family.

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Order RODENTIA 2000

Verheyen (16)............................. Setzer (34).............. Roberts (190)............................ Allen (21).......................... Osgood (39)..................... Thomas (387).................................. Heller (65).......... Dollman (79)........ Wroughton (67)........ De Winton (41)........ Lataste (19).........

Number of Taxa

1500

1000

500

Figure 4. The number of taxa of African rodents. Green line: cumulative number of species-group taxa (species and subspecies) described from Linnaeus (ca. 1750) to the year 2003 (total = 1729); principal authors and the span of their scientific contributions (dotted lines) are indicated, with the number of African taxa they described in parentheses. Red line: number of species considered scientifically valid (line fitted visually); red dots indicate species numbers recognised by Murray (1866), Trouessart (1897–98), Allen (1939), Ellerman (1940–41), Corbet & Hill (1980, 1991), Wilson & Reeder (1993, 2005). (From original by M. D. Carleton.)

Heuglin (22)............. Peters (34)....................... Smith (29).............. Gray (22)............................ Temminck (13)........................ Cuvier (13).................. Lichtenstein (11)..................................... Linnaeus (6) ............

0

1750

1800

1850

1900

1950

2000

Year

Middle East; generic and specific endemism is nearly 100% within sub-Saharan zones, where taxa tend to be more narrowly restricted in distribution and limited to distinct sub-regions (e.g. Coe & Skinner 1993, Happold 1996). The magnitude of species endemism within Africa, albeit nearly 95% (Table 6), surely remains an underestimate, for the number of rodent species considered to be valid has continued to increase in the last quarter-century (Figure 4). Knowledge of the diversity of African rodents initially grew slowly in the post-Linnaean discovery phase (ca. 1800–1860s). However, an exceptionally fertile period of natural history exploration and taxonomic description commenced in the late 1800s, during which the subspecies emerged as a standard rank to formally identify intra-specific variation. Oldfield Thomas of the British Museum of Natural History was a prodigious worker and influential systematist throughout this eruptive phase of taxonomic description (ca. 1880–1930s). By the conclusion of this era, more than 600 species of African rodents were recognized (Allen 1939). During the middle 1900s (ca. 1940–1970s), the number of species considered to be valid dipped to less than 300 (Corbet & Hill 1980), a decrease driven by widespread acceptance of the biological species concept and its taxonomic application as intergrading geographic races (subspecies). New subspecies continued to be named at a high rate, and many species described during the preceding eruptive phase were cursorily demoted to subspecies. The last subspecies of African Rodentia were named in 1983. The revived appreciation of greater species richness, as witnessed in the past two decades, has been impelled by new field discoveries, by reinvigorated museum-based revision and by improved methodologies for evaluating and taxonomically interpreting interpopulation differences, be they morphological, chromosomal, or molecular. Thus, the approximately 400 species covered herein are a tenuous summary of an animated study in progress, whose outcome over the next 10–15 years is likely to stabilize the number of African rodents at about 500 species (Figure 4).

Coupled with its species richness, the Rodentia is remarkable for its morphological diversity, and essentially every ecological, behavioural, trophic and locomotory adaptive theme that has evolved within the order is represented among the African taxa (see familial and generic accounts).

Morphological definition and incisor function Notwithstanding the protean morphological variety exhibited by rodents, the order is well defined in combination by several derived characters or functionally related character complexes. The upper and lower jaws each bear a single pair of enlarged and ever-growing chisellike incisors that ontogenetically represent retained deciduous second incisors (i 2/2 – Luckett 1985); the incisor enamel is mainly limited to the anterior surface and is two-layered, the inner layer of which is composed of hydroxyapatite crystal prisms arranged in characteristic decussating patterns (Martin 1993, 1999). The dentition consists of a maximum of 22 teeth (incisors 1/1, canines 0/0, premolars 2/1, molars 3 /3) as documented in the earliest of fossil species currently recognized as rodents. The upper and lower molars are quadritubercular, the uppers possessing a longitudinal enamel connection (mure) between the protocone and hypocone and the lowers lacking a paraconid (Meng et al. 2003, Marivaux et al. 2004).The glenoid fossa is elongate, lacking pre- and postglenoid processes, and enables the free anterior–posterior excursion of the mandible into effective occlusion for gnawing (incisors) or chewing (cheekteeth). A conspicuous diastema is present between the anterior incisors and posterior cheekteeth due to loss of the first and last incisors, canines and anterior premolars; the upper diastema is significantly longer than the lower. With formation of a diastema, fur-covered labial infoldings occur behind the incisors to partially segregate gnawing from chewing activity. The masticatory muscles

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Order RODENTIA

are complexly subdivided, the masseteric series being massive in size and volume relative to the temporalis; a discrete superficial masseter is always present. In primitive rodents, all masseter muscles originate from the zygomatic arch, and the infraorbital foramen is small, serving as passage only for nerves and blood vessels (protrogomorphous). In derived rodents, different masseteric bundles extend anteriorly either onto an expansive zygomatic plate (sciuromorphous), through a greatly enlarged infraorbital foramen onto the lateral rostrum (hystricomorphous), or onto both in combination (myomorphous). The latter three zygomasseteric morphologies are represented among living African Rodentia (Figure 5) and have figured prominently as characters in its subordinal classification (see below). The auditory bulla is composed entirely of the ectotympanic bone. The literature on rodent morphology is extensive: see Tullberg (1899), Ellerman (1940) and Carleton (1984) for additional morphological characterization of Rodentia; see Hartenberger (1985), Luckett (1985), Luckett & Hartenberger (1993), Landry (1999) and Meng et al. (2003) for phylogenetic commentary on the preceding traits and others as ordinal synapomorphies, particularly as compared with Lagomorpha. The enlarged incisor teeth, reduced to a single pair each at the front of the upper and lower jaws, represent a key evolutionary innovation (sensu Brooks & McLennan 1991) that plausibly fostered the remarkable success of the order, which typically constitutes the most species-rich clade of Mammalia found in the fossil record since its initial radiation in the early to middle Eocene (40–55 mya). That success owes much to the intrinsic biomechanical properties of the continually erupting incisors and their consequent effectiveness as a multipurpose tool. The differential hardness of the enamel and dentine, together with microstructural specializations of the enamel layers, allow formation of an acute edge. Hydroxyapatite crystal prisms of the inner enamel layer are arranged in intricately interwoven patterns (pauciserial in early rodents, multiserial and uniserial in derived lineages – Martin 1993); these further reinforce the working bevel of the incisors and inhibit propagation of cracks along the relatively brittle and thin enamel sheath during conditions of intense activity and when strong forces are concentrated on the narrow tips (von Koenigswald 1985, Martin 1993). Ultimately, the keen sharpness and chisel shape of the incisors are achieved by deliberate honing motions of the lower incisors upon the upper incisors, not as a coincidental result of wear (von Koenigswald 1985, Druzinsky 1995). Indeed, some researchers consider the chisel shape of the rodent lower incisor and its intentional maintenance as a cardinal morphological-behavioural synapomorphy of the order (Landry 1999, Meng et al. 2003). The adaptive role of the incisors is usually described as ‘gnawing’, but that term inadequately captures the range and multiplicity of their uses. Biting and cutting, piercing and stabbing, holding and seizing, tearing and slicing, gouging and scraping (digging), chipping and prising, interdental sharpening and chattering, gathering and carrying of foodstuffs or nesting material, carrying of young by parents and teat-clinging by neonates, are other functions observed within species and across the order (Lawrence 1941, Landry 1970, Offermans & De Vree 1990, Druzinsky 1995, van der Merwe & Botha 1998). Most of the functions of the incisors involve feeding and food habits and reflect the diverse trophic niches and dietary flexibility that are hallmarks of Rodentia (Landry 1970); other functions relate to aspects of communication, behaviour, ecology and reproduction. The exceptional mobility of the mandible, so integral to a rodent’s

livelihood, was evolutionarily realized through extensive reorientation of the jaw muscles and concomitant remodelling of the head skeleton. Although these adaptations originally appeared to protrude the mandible for incisor occlusion and to strengthen incisal cutting (Meng et al. 2003), they coincidentally preadapted the masticatory apparatus for subsequent evolution of molar hypsodonty, planar occlusal surfaces and propalinal chewing suited to triturating high-fibre plant material, notably grasses (Landry 1999, Mess et al. 2001). Expanding reliance upon herbivorous resources has characterized several rodent lineages that independently radiated in concert with the development of open environments in the Oligocene and proliferation of grasslands in the Miocene (Jacobs et al. 1999, Mess et al. 2001).

(a) Sciuromorpha

(b) Myomorpha

(c) Hystricomorpha

Figure 5. Orientation of jaw musculature in the three suborders of African Rodentia. Stippled arrow = masseter lateralis muscle (= middle masseter). Black arrow = masseter medialis muscle (= deep masseter). Clear arrow = muscle attached to inner surface of zygomatic arch. (a) Sciuromorpha: masseter lateralis originates on outer side of skull in front of orbit (zygomatic plate) and inserts on posterior part of mandible, and masseter medialis originates on the inner surface of the zygomatic arch and inserts on the middle part of the mandible. Infraorbital foramen is very small; no part of the masseter muscle passes through the foramen. (b) Myomorpha: masseter lateralis is similar to that of Sciuromorpha but the masseter medialis has pushed up through the orbit and through the enlarged infraorbital foramen so its origin is on the rostrum. (c) Hystricomorpha: masseter lateralis retains the primitive position originating on the inner surface of the zygomatic arch and inserts on the mandible, and the masseter medialis has its origin more anteriorly on the mandible and passes through the enlarged infraorbital foramen and inserts on the rostrum. Other jaw muscles (superficial masseter muscle and temporalis muscles), which are similar in all suborders, are not shown. The area of muscle attachment to the underlying bone is more extensive than the arrows indicate (after Romer 1966).

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Phylogeny and classification Such morphological traits have long encouraged the view that living rodents are derived from a common ancestor, as conveyed in numerous classifications of Rodentia from Brandt (1855) to Landry (1999). Taxonomically broad, cladistic studies of morphological characters have sustained the historical view of Rodentia as a monophyletic group (Novacek 1985, Novacek & Wyss 1986, Luckett & Hartenberger 1993, Shoshani & McKenna 1998, Meng et al. 2003). Although some early gene-sequencing studies implied, or concluded, that hystricognath rodents (Cavia and relatives) represent a separate mammalian order and that Rodentia as conventionally classified is polyphyletic (Graur et al. 1991, 1992, D’Erchia et al. 1996), the ensuing wave of molecular research, based on wider sampling of taxa (particularly the in-group Rodentia) and genes, has decisively solidified the case for rodent monophyly (Huchon et al. 1999, 2002, Robinson-Rechavi et al. 2000, Adkins et al. 2001, 2003, Debry & Sagel 2001, Lin et al. 2002, Armine-Madsen et al. 2003, Debry 2003, Waddell & Shelley 2003, Reyes et al. 2004). Recognition of major lines of descent within Rodentia and their formal classification, usually at the rank of suborders, historically has provoked more contention than the question of monophyly of the order.

The two prevalent classificatory schemes stem from Brandt (1855) and Tullberg (1899): Brandt’s tri-subordinal division emphasized zygomasseteric morphology (sciuromorphy, hystricomorphy and myomorphy); and Tullberg’s dual subordinal arrangement integrated mandibular conformation (sciurognathy versus hystricognathy) with zygomasseteric criteria (Table 7). Subsequently, most systematists have adopted or elaborated either Brandt’s (Alston 1876, Simpson 1945, Wood 1965) or Tullberg’s subordinal taxonomies (Ellerman 1940, Chaline & Mein 1979, Landry 1999). Other classifications have recognized anywhere from five to 16 major phyletic divisions within Rodentia, whether or not called a suborder per se (Thomas 1897, Miller & Gidley 1918,Wood 1955,Thaler 1966, Hartenberger 1985, 1998, McKenna & Bell 1997, Carleton & Musser 2005). In spite of these classificatory disagreements at the level of suborder, certain phyletic affinities among the 33 recognized families have consistently emerged based on morphological and gene-sequencing evidence published over the past decade. Although ranks between family and order are not considered in this volume, the following four kinship-groups are apparent among the 14 rodent families that occur in Africa. Morphological characters, evidence for monophyly, and fossil history of individual families are covered in the respective family introduction.

Table 7. Taxonomic placement and presumed higher-level relationships among 14 African families as interpreted in past influential classifications of Rodentia.a Alston (1876)

Tullberg (1899)

Simpson (1945)

McKenna & Bell (1997)

Sciuromopha Sciuridae Anomaluridae

Sciurognathi Sciuromorphi Sciuridae

Sciuromorpha Sciuridae Sciuromorpha i.s. Anomaluroidea Anomaluridae Pedetidae Myomorpha Gliroidea Gliridae Dipodoidea Dipodidae Muroidea Spalacidae Cricetidae [Nesomyidae] Muridae

Sciuromorpha Sciuridae

Myomorpha Gliridae Dipodidae [Pedetidae] Spalacidae [Bathyergidae] Muridae [Cricetidae] [Nesomyidae]

Hystricomorpha Hystricidae Octodontidaeb [Ctenodactylidae] [Petromuridae] [Thryonomidae]

Myomorphi Myoidea Gliridae Dipodidae Spalacidae Nesomyidae Cricetidae Muridae Anomaluroidea Anomaluridae Pedetidae Ctenodactyloidea Ctenodactylidae Hystricognathi Hystricomorphi Hystricidae Petromuridae Thryonomyidae Bathyergomorphi Bathyergidae

Hystricomorpha Hystricidae Petromuridae Thryonomyidae Hystricomorpha i.s. Bathyergidae Suborder i.s. Ctenodactylidae

Myomorpha Myodonta Dipodidae Muridae [Spalacidae] [Cricetidae] [Nesomyidae] Gliromorpha Gliridae Anomaluromorpha Anomaluridae Pedetidae Sciuravida Ctenodactylidae Hystricognatha Hystricidae Petromuridae Thryonomyidae Bathyergidae

i.s. = incertae sedis. Those African families in brackets were not specifically recognized in a given classification, but their genera belong to the family listed above. b In early classifications, such as Alston (1876) and Thomas (1897), Petromus, Thryonomys and the ctenodactylid genera were grouped with the Octodontidae of South America. Subsequent research, beginning with Tullberg (1899), has indicated that these African families constitute distinct families endemic to Africa and that they are only distantly related to the octodont rodents of South America. a

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(1) Sciuridae + Gliridae (= Sciuriomorpha)

Sciurid rodents possess a sciurognathus mandible and sciuromorphous masseteric condition and have composed the core of Brandt’s (1855) Sciuromorpha, as reflected in classifications generated from the late 1800s to the present (e.g. Alston 1876, Thomas 1897, Simpson 1945, McKenna & Bell 1997). Union of Sciuridae with Gliridae (dormice) in an expanded clade, however, recently has garnered additional evidence. The zygomasseteric structure of extant glirids (excepting African Graphiurus) was widely interpreted as myomorphous, and so influenced systematists to associate dormice with Myomorpha (Alston 1876, Thomas 1897, Ellerman 1940, Chaline & Mein 1979, Wahlert et al. 1993). However, the ‘myomorphy’ observed in most living dormice is now understood to be convergent to the zygomasseteric configuration in Myomorpha proper and has been termed ‘pseudomyomorphy’ (Vianey-Liaud 1985, Maier et al. 2002). The deep and rich fossil record of Gliridae implies their derivation in the very early Eocene from protrogomorphous ischyromyids (VianeyLiaud 1985, 1994, Hartenberger 1994, 1998), the same fossil group also known to be ancestral to Sciuridae (and North American Aplodontidae). In agreement with the palaeontological interpretation, analyses of both mitochondrial and nuclear genes disclose Gliridae as sister-group to Sciuridae or to a sciurid-aplodontid clade (Kramerov et al. 1999, Huchon et al. 1999, 2002, Adkins et al. 2001, 2003, Debry & Sagel 2001, Montgelard et al. 2002, Reyes et al. 2004).

(2) Dipodidea [Dipodidae] + Muroidea [Spalacidae–Nesomyidae–Cricetidae– Muridae] (= Myomorpha)

Common ancestry of the Dipodoidea and Muroidea has long been recognized, notably as first depicted by Tullberg (1899). Their close phyletic union has been sustained by substantial palaeontological data (Wilson 1949, Emry 1981, Walsh 1997, Emry et al. 1998) and morphological systems, such as molar occlusal patterns and loss of the lower fourth premolar (Flynn et al. 1985, Marivaux et al. 2004), cephalic arterial plans (Bugge 1985) and foetal membrane traits (Luckett 1985). Although extant dipodoids (hystricomorphous) and muroids (myomorphous) differ in their zygomasseteric construction, extinct cricetids of the late Eocene and early Oligocene possessed a hystricomorphous infraorbital condition (Flynn et al. 1985, VianeyLiaud 1985); fossils with intermediate zygomasseteric structures, along with plausible myological transformations (Klingener 1964), endorse the derivation of myomorphy from an hystricomorphous ancestor. Such a conclusion is further implicit in molecular studies of broad taxonomic scope (Nedbal et al. 1996,Adkins et al. 2001, 2003, Debry & Sagel 2001, Michaux et al. 2001, Jansa & Weksler 2004). Schaub (1958) formally identified the cognate relationship of Dipodoidea and Muroidea as the infraorder Myodonta, within the suborder Myomorpha, a classification also recognized by McKenna & Bell (1997).

(3) Anomaluridae + Pedetidae (= Anomaluromorpha)

Scaly-tailed squirrels (Anomaluridae) and springhares (Pedetidae) are characterized by a hystricomorphous skull and sciurognathus mandible, an unusual combination of traits that has confused their clear-cut assignment to the traditional suborders. Some systematists have treated one or both families as incertae sedis within Rodentia (e.g. Simpson 1945, Wood 1965, Chaline & Mein 1979, Hartenberger

1998). Others have arranged the two groups in the same superfamily or suborder (Winge 1887, Tullberg 1899, McKenna & Bell 1997), and this kinship has drawn support based on arterial patterns (Bugge 1974b, George 1981c), middle ear anatomy (Lavocat & Parent 1985, Meng 1990) and mitochondrial genes (Montgelard et al. 2001, 2002). Some investigators dispute any special relationship between Anomaluridae and Pedetidae (e.g. Jaeger 1988a, Hartenberger 1998, Landry 1999), especially given fundamental microstructural differences in their incisor enamel (Martin 1993, 1995). These authors would independently derive Anomaluridae from certain Eocene families (Martin 1993, Hartenberger 1998) or ally it as a sister-group with Sciuridae (Landry 1999), and tentatively affiliate Pedetidae with the Ctenodactylidae– Hystricognathi clade (George 1985b, Martin 1995, Landry 1999).The phylogenetic issues surrounding Anomaluridae and Pedetidae and their suprafamilial classification continue to invite resolution.

(4) Ctenodactylidae + Hystricognathi [Bathyergidae, Hystricidae, Petromuridae, Thryonomyidae] (= Hystricomorpha)

The African gundis, Ctenodactylidae, represent another group whose phylogenetic placement has perplexed systematists. The family was early placed in Hystricomorpha (Alston 1876, Thomas 1897), but later arranged within Sciurognathi (Ellerman 1940, Chaline & Mein 1979), isolated (among living families) in its own suborder, Sciuravida (McKenna & Bell 1997), or left as a suborder or infraorder incertae sedis (Simpson 1945, Wood 1955, 1965). A jointly impressive body of research supports association of ctenodactylids with Hystricomorpha sensu lato. Such corroborative studies broadly draw upon morphology, both individual character systems and multicharacter cladistic analyses (Landry 1957, 1999, Bugge 1985, George 1985b, Lavocat & Parent 1985, Meng 1990, Martin 1994, 1995, Bryant & McKenna 1995), palaeontology (Flynn et al. 1986, Marivaux et al. 2002, 2004, Marivaux & Welcomme 2003) and DNA sequences (Huchon et al. 2000, 2002, Adkins et al. 2001, 2003, Huchon & Douzery 2001, Montgelard et al. 2002, Jenkins et al. 2005). Since Tullberg’s (1899) seminal monograph, the four African families that possess hystricomorphous jaw musculature and a hystricognathus mandible (Bathyergidae, Hystricidae, Petromuridae, Thryonomyidae) have been viewed as closely related to one another and to those in the New World. Subsequent research has ratified his arrangement in the broad sense (see summaries in Luckett & Hartenberger 1985, Landry 1999, Jenkins et al. 2005). Whether the Palaeotropical families (Phiomorpha sensu Lavocat 1971) and Neotropical families (Caviomorpha sensu Wood & Patterson 1959) represent monophyletic radiations within Hystricognathi remains uncertain, although some recent investigations support such an interpretation (Nedbal et al. 1994, Huchon & Douzery 2001, Marivaux et al. 2004). Rodents have long been accepted as most closely related to hares and rabbits. The two groups were initially arranged as the suborders Simplicidentata (rodents) and Duplicidentata (hares and rabbits) in the order Rodentia or Glires (e.g. Alston 1876, Thomas 1897, Gregory 1910) and later placed in separate orders, Rodentia and Lagomorpha, within the cohort Glires (Gidley 1912, Simpson 1945, Landry 1999). Surprisingly, a persuasive palaeontological foundation for their cognate relationship did not emerge until the late 1900s (Li et al. 1987, Li & Ting 1993, Dashzeveg et al. 1998).With increased knowledge of these extinct groups (Mimotonidae, Eurymylidae and others), relationships 31

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among basal Glires and their classification have become correspondingly complex (see Meng et al. 2003). Notwithstanding the phylogenetic intricacies involving extinct gliriform mammals, common ancestry of the orders Lagomorpha and Rodentia among living mammals is decisively sustained in modern phylogenetic studies (for morphological evidence, see Novacek & Wyss 1986, Novacek et al. 1988, Luckett & Hartenberger 1993, Shoshani & McKenna 1998, Martin 1999, Meng & Wyss 2001, Meng et al. 2003; for molecular evidence, see Eizirik et al. 2001, Murphy et al. 2001, Huchon et al. 2002, Lin et al. 2002, Armine-Madsen et al. 2003, Waddell & Shelley 2003). The clade Rodentia–Lagomorpha is distantly related to Primates, Dermoptera (colugos) and Scandentia (tree shrews), a phyletic association of five orders now recognized as the superorder Euarchontoglires (Murphy et al. 2001, Scally et al. 2001, Huchon et al. 2002, Lin et al. 2002, Helgen 2003, Waddell & Shelley 2003, Reyes et al. 2004).

Geologic age and continental areas of origin The oldest rodents represent several ischyromyid genera (Sciuromorpha) from the late Palaeocene of North America, and these or related forms are documented in the early Eocene of Asia and Europe (McKenna & Bell 1997, Hartenberger 1998). The explosive diversification of Rodentia into principal lineages (suborders) occurred during the early to middle Eocene, and examples of most modern families are encountered by the middle to late Oligocene (Wood 1959, Hartenberger 1996, 1998). In Africa, the earliest rodents so far recorded appear in the early Eocene, including forms (Zegdoumyidae) considered to be ancestral to living Anomaluridae (Vianey-Liaud et al. 1994), and in the late Eocene–early Oligocene, including forms (Phiomyidae) related to African Hystricognathi, in particular Thryonomyidae (Wood 1968, Lavocat 1978, Jaeger et al. 1985).These Eocene and Oligocene records originate from sites in the Sahara and along the Mediterranean rim of North-West Africa. The first geologic appearances of most African rodent families, including the largest groups (Sciuridae, Gliridae, Nesomyidae, Muridae, Bathyergidae), date from the middle to late Miocene, the majority as recorded from sub-Saharan countries (Table 8). Based on the oldest fossils so far discovered and their cladistically basal stature within Glires or Rodentia, Asia is presently accepted as the continental area of origin for the order (Bryant & McKenna 1995, Dawson & Beard 1996, Hartenberger 1996, Beard 1998, Meng & Wyss 2001). African rodent groups are generally thought to be derived from southern Asian progenitors, some time during the middle to late Miocene, and they presumably reached Africa by immigration through the Arabian Peninsula into East Africa and along the Mediterranean into North Africa. The timing and direction of faunal interchange are based on intercontinental comparisons of earliest fossil dates (Lavocat 1978, Jacobs 1985, Wang 1994, Winkler 1994, Flynn & Jacobs 1999) and on cladistic biogeography as inferred from phylogenetic studies (Huchon & Douzery 2001, Mercer & Roth 2003). Such a biogeographic origin and dispersion chronology convincingly account for certain families that occur in the Sahara and North Africa. For example, the ctenodactylids persist as relicts from a much older but now extinct radiation in Asia (Flynn et al. 1986, Wang 1994, Dashzeveg & Meng 1998), and the few African dipodids

Table 8. Earliest geologic occurrence of extant families and subfamilies of African Rodentia. Family/Subfamily

Epoch

Sciuridae1 Gliridae2

E. Miocene (Vulcanisciurus, Kenya) M.–L. Miocene (Graphiurus, South Africa; Microdryomys, Morocco; Otaviglis, Namibia) M. Miocene (Protalactaga, Morocco) Recent (Spalax, Libya)

Dipodidae3 Spalacidae4 Spalacinae Tachyoryctinae? Subfamily? Nesomyidae5 Cricetomyinae Dendromurinae Mystromyinae Petromyscinae Cricetidae6 Arvicolinae Lophiomyinae Muridae7 Deomyinae Gerbillinae

Murinae Otomyinae Anomaluridae8 Pedetidae9 Ctenodactylidae10 Bathyergidae11 Hystricidae12 Petromuridae13 Thryonomyidae14

L. Miocene (Nakalimys, Kenya) M.–L. Miocene (Harasibomys, Namibia; Pronakalimys, Kenya) E. Miocene? (Protarsomys, Kenya) L. Miocene (Saccostomus, Ethiopia, Namibia) M.–L. Miocene (Dendromus, Ethiopia, Namibia; Mabokomys, Kenya; Ternania, Kenya) E. Pliocene (Mystromys, South Africa) L. Miocene (Harimyscus, Namibia) M. Miocene (Democricetodon, Kenya) M. Pleistocene (Microtus, Libya) L. Miocene (Lophiomys, Morocco) L. Miocene (Preacomys, Ethiopia, Namibia) M.–L. Miocene (Mioharimys, Namibia; Myocricetodon, Kenya, Morocco; Protatera, Morocco) M.–L. Miocene (Aethomys, Namibia; Leakeymys, Kenya; Paraethomys, Algeria; Saidomys, Kenya) L. Miocene (Euryotomys, South Africa) L. Eocene (Nementchamys, Algeria) E. Miocene (Megapedetes, Kenya; Parapedetes, Namibia) M. Miocene (Africanomys, Morocco, Tunisia) E. Miocene (Proheliophobius, Kenya) L. Miocene (Hystrix, Kenya) E. Pleistocene (Petromus, South Africa) L. Eocene–E. Oligocene (Guadeamus, Egypt; Paraphiomys, Egypt)

Epoch: E. = Early; M. = Middle; L. = Late. References: 1Lavocat 1973. 2Lavocat 1978, Denys 1990b, Mein et al. 2000a. 3Jaeger 1977b. 4Flynn & Sabatier 1984, Tong & Jaeger 1993, Mein et al. 2000a. 5Lavocat 1973, Denys 1991, Tong & Jaeger 1993, Winkler 1998, Mein et al. 2000b, 2004, Geraads 2001. 6Tong & Jaeger 1993, McKenna & Bell 1997, Geraads 1998. 7Jaeger 1977b, Tong & Jaeger 1993, Sènègas & Avery 1998, Mein et al. 2000b, 2004, Winkler 2001. 8Jaeger et al. 1985, Vianey-Liaud et al. 1994. 9Lavocat, 1978. 10Robinson & Black 1973, Jaeger 1977b,Winkler 1994. 11Winkler, 2001. 12Lavocat 1973. 13Lavocat 1978. 14Wood 1968, Lavocat 1978.

(3 spp.) represent outliers of the core diversity of Dipodidae, both as fossils and extant species, in Asia (Klingener 1984, Wang & Dawson 1994, Wang & Qiu 2000). Other taxa plausibly represent much younger (Pliocene–Pleistocene) incursions to North Africa, presumably following a circum-Mediterranean route (Cricetidae [Arvicolinae], Spalacidae [Spalacinae], Gliridae [Eliomys]). The timing and direction of continental exchange are not so clearcut for many groups whose distribution and diversity are centred in sub-Saharan Africa, although the conventional explanation emphasizes migration from Asia into Africa (Jacobs 1985, Winkler 1994, Flynn

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& Jacobs 1999). For example, all African sciurids belong to an extensive radiation (Xerinae: Xerini and Protoxerini) that is notably abundant in sub-Saharan biomes (Mercer & Roth 2003, Steppan et al. 2004); Holarctic ground squirrels and chipmunks constitute a derived clade (Marmotini) within this subfamily. The other extant genus of African Gliridae, Graphiurus, is modestly speciose within sub-Saharan forests and woodlands and is the sole member of a distinctive subfamily (Graphiurinae) that represents the earliest phyletic branch within the family (Wahlert et al. 1993, Montgelard et al. 2003), which also occurs in Europe and Asia (Hartenberger 1994). A related graphiurine (Otaviglis) has been recently described from the late Miocene of Namibia, and Graphiurus itself is recorded from the latest Miocene of South Africa (Table 8); according to molecular estimates, differentiation of the graphiurine lineage dates to 40–50 mya (Montgelard et al. 2003). Living members of Nesomyidae are similarly known only from the sub-Saharan region (some fossil Dendromurinae documented from North Africa and Arabian Peninsula – see subfamily profile), and the several subfamilies compose a clade separate from, perhaps basal to, the radiation of Cricetidae and Muridae (Michaux et al. 2001, Jansa & Weksler 2004). The Miocene diversity of the family, including earliest records of extant genera, has been revealed by significant studies published since the middle 1990s (Table 8). Three of the five subfamilies of Muridae (Deomyinae, Leimacomyinae, Otomyinae) are concentrated within the sub-Saharan region (Acomys, a deomyine, also occurs in North Africa and the Middle East). The murid subfamily Gerbillinae has a broader distribution throughout Africa and into the Middle East and central Asia, but early-branching clades of Gerbillinae (Ammodillini, Taterillini) predominantly lie within the sub-Saharan region (Pavlinov et al. 1990). The Murinae, too, has an extensive distribution within Africa, across Eurasia and into Indomalaya and Australia; generic and specific endemism within sub-Saharan biomes is pronounced and includes a few large, well-defined clades (Watts & Baverstock 1995a, Ducroz et al. 2001, Lecompte et al. 2002b). A molecular-clock estimate places the divergence between Deomyinae, Gerbillinae and Murinae in the early Miocene (Michaux et al. 2001). Bathyergidae (African mole-rats) is another family endemic to the sub-Saharan region and contains five strongly differentiated genera (Honeycutt et al. 1991). Molecular studies divulge startlingly older dates for phyletic divergence among bathyergids (middle Eocene, ca. 40 mya – Huchon & Douzery 2001) than presently reflected in palaeontological data (Table 8) and point to an autochthonous East African origin of the family (Faulkes et al. 2004). Although Asia, or to a lesser extent Europe, seems the probable source-area for Africa’s ancient (Eocene) rodent stocks, the foregoing examples implicate the continent as an important secondary centre for the origin (Oligocene–early Miocene) of major groups. The substantial taxonomic differentiation at the generic, subfamily and family levels, Miocene appearances of extant genera, and deep cladistic patterns all indicate a longer phase of phylogenetic differentiation of Rodentia within sub-Saharan Africa than can be presently documented by the fossil record.The rarity (early Miocene) or absence (Oligocene) of critical-aged, productive fossil sites within the region offers, at best, a precarious foundation to argue continental origins and intercontinental exchanges for many African rodent families. Palaeontological investigation continues, of course, and it is instructive that some of the more exciting discoveries of

earlier appearances have emerged from contributions of the past decade. Future phylogeographic studies that include ample species sampling of genera, subfamilies and families with both African and Asian geographic ranges (e.g. Hystricidae, Spalacidae, Muridae [Gerbillinae, Murinae]) would measurably improve our understanding of continental origin and directions of faunal interchange.

Biology The immense variation in the biological characteristics of African rodents reflects the large number of species, their evolutionary diversity, and the very varied environments where they are found. This section provides a necessarily brief overview of the diversity of some aspects of the biology of the order. More detailed information may be found in each genus and species profile. General reviews are given in Delany & Happold (1979), Delany (1972, 1986) and Happold (2001); and the role of rodents in African ecosystems is discussed by Dieterlen (1989) and Happold (1996) for rainforests, Happold (1984) for deserts, and Happold (1983) for savannas. The amount of biological information on African rodents varies greatly depending on the species; for some, nothing is known because of their rarity and limited geographical distribution; for many, a limited number of observations are available; and for only a few species (those which are widespread, unusual, or of economic importance) there is substantial information. In spite of the impressive diversity of the order, few species are biologically well understood, and even the best-known species are not so well known as many of Africa’s iconic large mammals that belong to far less speciose orders (Proboscidea, Primates, Carnivora, Perissodactyla and Ruminantia). African rodents vary greatly in size; the majority weigh 20–80 g, but the smallest weighs only 3 g (Mus haussa), and the largest weigh ca. 3.25 kg (Pedetes spp.) and ca. 20 kg (Hystrix spp.). Most African rodents are typically ‘mouse-like’ or ‘rat-like’ but there are many that are very different; for example, jerboas (Allactaga, Jaculus) and springhares (Pedetes) have extremely long hindlimbs and jump like kangaroos; anomalures (Anomalurus, Idiurus) have large winglike membranes so they can glide from tree to tree; and mole-rats (Bathyergus, Georhychus, Heliophobius) have blunt muzzles, cylindrical bodies, large short spatulate forelimbs, rudimentary (or non-existent) eyes, and huge incisor teeth (for excavating burrows). Because of the large number of species representing families of distant evolutionary origins, there are some interesting examples of ecological and morphological convergence: species of Dipodidae and Pedetidae are bipedal and saltatorial; species of Bathyergidae and Spalacidae are permanently subterranean; species of Muridae, Gliridae and Sciuridae all include some species that are arboreal; and species of Dipodidae and Gerbillinae have the ability to produce very concentrated urine and hence can live in very dry habitats devoid of free water. Likewise, convergence in form and biology occurs in distantly related taxa that inhabit similar but well separated habitats; for example the following genera occur, respectively, in the Sahara Arid BZ and the South-West Arid BZ: Gerbillus spp. and Gerbillurus spp., Atlantoxerus and Xerus spp., Ctenodactylus spp. and Petromus typica, and Acomys spp. and Petromyscus spp. The rich morphological variety exhibited by African rodents presupposes the remarkable diversity recorded in their life histories, habits and ecological niches. 33

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Order RODENTIA

Rodents are found throughout the African continent, in every biotic zone, and in almost every habitat from the driest of deserts to the wettest of rainforests. Most species are terrestrial, living on the surface of the ground; many of them rest in burrows or under rocks when inactive. Other species may be described as scansorial because they also clamber about in low vegetation close to the ground. Arboreal species (several genera of squirrels, Graphiurus, Grammomys, Thallomys) spend all (or most) of their time in large shrubs and trees, and rarely descend to the ground. Some small species are adapted for climbing on tall grasses (Dendromus, Dendroprionomys, Prionomys) and have opposable fingers and prehensile tails for holding on to slender grass stems. A number of genera (Bathyergus, Georhychus, Heliophobius, Heterocephalus) are subterranean and live permanently underground. Only a few species typically live in wet swampy habitats and are capable of swimming; such forms (Deomys ferrugineus, Colomys goslingi and Malacomys spp.) are characterized by relatively long limbs and large hindfeet. Most species are very habitat-specific, being found in only one or two habitats. In contrast, a few species (e.g. Mastomys natalensis, Cricetomys gambianus, Thryonomys swinderianus, Hystrix spp.) are widespread and habitat generalists. The geographical range of many species is relatively well known from museum specimens and trapping records (see maps in each species profile). However, such geographical ranges do not provide any indication of the abundance of a species within its range. Abundance may be expressed either as the numbers of individuals captured using a standardized trapping method, or as a proportion (usually expressed as a percentage) that a species contributes to the community in which it lives, or as the density of individuals/unit area (such as the number of individuals of a species per hectare, or the total number of individuals of all species per hectare). Such measures may vary greatly between habitats and locations, and according to the time of year. Typical examples of proportional representation are: individuals of Dasymys rufulus comprise 14.6% of the community of trappable small mammals at Mount Nimba (Guinea), 9.2% in the savannas at Bolo (Côte d’Ivoire) and 7.5% in the savannas at Lamto (Côte d’Ivoire). Dephomys defua contributes 10% in four different rainforests in Ghana, Heimyscus fumosus contributes 4.6–20.5% (mean 11.5%) and forms the second or third commonest species of nine syntopic species in Gabon, and Praomys daltoni comprises 0–78% (mean 50%) at 11 localities in Nigeria. Density is a particularly useful measure of abundance because it provides actual numbers; for example, the density of Hybomys univittatus in forests of the Central African Republic was 3.02/ha (in lightly logged forest) and 4.14/ha (in heavily logged forest) (Malcolm & Ray 2000); the density of Lophuromys flavopunctatus in DR Congo varied from 9.8/ha (in savannas) to 19.6/ha (in dry seasonal swamps) (Misonne 1963); and the density of Aethomys namaquensis in different countries ranged from 2/ha in succulent Karoo, South Africa, to 5.4/ha in Namibia, 4–13/ha in Mozambique and 5.2–27.3/ ha in Zimbabwe. Density also varies seasonally: e.g. the density of Arvicanthis abyssinicus in Ethiopia was 65–75/ha during the wet season and 130–250/ha during the dry season (Müller 1977), and in Praomys tullbergi in Nigerian rainforest, density ranged between 9.5/ha and 27.9/ha according to season over a period of three years (Happold 1977). Seasonal fluctuations in numbers tend to be smallest in species that live in rainforest, and largest in species that live in conditions where rainfall is low and unpredictable. Inter-annual fluctuations are typically rather large in dry localities: e.g. densities of Taterillus

pygargus in the sahel savanna of Senegal varied from 0/ha after the drought of 1972 to 180/ha after the good wet season of 1975 (Poulet 1978). For just a few species, live-trapping of individual animals over a long period of time has provided detailed information on population dynamics, home-ranges, movements and longevity (e.g. Praomys tullbergi, Acomys spinosissimus, Mastomys natalensis). Such relatively wellknown species are the exception among African rodents. Many more have been rarely encountered and have very restricted geographical ranges, and hence very little is known about the biology of such species. Whether this ‘rarity’ is real or due to inappropriate methods of study is uncertain; whatever the reason, obtaining information on such species provides a challenge for ecologists in the future. The examples given above refer only to single species; other measures concern all the species that comprise a community; these include ‘species richness’ (= the number of species in the community) and density (= the total number of individuals of all species known to be alive [KTBA]) in the community. These community measures are especially useful when assessing trophic interactions between species, the overall impact of the community on its environment, and the interactions between different species in the community, especially in respect of reproductive strategy and seasonal changes in demography. For example, the species richness in a rainforest in SW Gabon was 11 species of rodents (plus ten species of shrews) (Nicolas & Colyn 2003; Nicolas et al. 2005). In a similar rainforest in Nigeria, species richness was seven species of rodents (and two species of shrews) and density, which varied according to season, was 14.3–25/ha (Happold 1977). (Equivalent data on numbers are unavailable for Gabon.) Similar variations are recorded in savanna habitats: in Brachystegia savanna in Malawi, species richness was seven species of rodents (and one species of sengi [Macroscelidae]) and density was 3–21/ha according to season (Happold & Happold 1991); equivalent data for thornbush savanna in Malawi was seven species of rodents (and one species of shrew) and 1–33/ha (Happold & Happold 1990). In Brachystegia savanna in Zimbabwe, species richness was seven species of rodents (and one species of shrew) and density was 2–10/ha (Linzey & Kesner 1997a). Habitat has a very strong influence on community structure; for example, species richness and density of small rodents during one year in Zimbabwe varied greatly in five neighbouring habitats as follows: riverine grasslands – 7 spp. 1–36/ha; miombo woodland – 7 spp., 2–10/ha; mopane woodland – 5 spp., 1.6–19/ha; talus – 4 spp., 8–25/ha; and thicket – 0.8–2/ha (Linzey & Kesner 1997a) (see also below). The reasons for such variation between habitats are uncertain and need further investigation; but the fact that such variations do occur implies differing levels of resources, differing levels of exploitation by rodents of these resources, and differing numbers and diversity of rodent predators in each habitat. Rodents show a wide range of reproductive methods and reproductive strategies. The strategy of any particular species is determined partly by its phylogeny, but also by the environment and location. Species vary greatly in their fecundity: squirrels, anomalurids and Pedetes have only 1–2 young per litter; many species have 3–6 young per litter; some have up to 10/litter; and the most fecund species (e.g. Mastomys natalensis, Heterocephalus glaber) may have as many as ca. 20/litter. Litter-size within a species may vary geographically, seasonally and in relation to the size/weight of the female; for example in Saccostomus campestris mean embryo number varies from 4.8 (South Africa) to 7 (Botswana) (see profile for

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references); in Gerbilliscus nigricaudus mean litter size is 4.2 in the dry season and 5.5 in the wet season (Neal 1982); and in Acomys spinosissimus mean litter-size is 2.9 for smaller females and 3.6 for larger females (Hanney 1965). Pregnancy and lactation is adaptively timed to ensure a high survival rate for the young. Because small rodents have relatively short gestations (ca. 25–40 days), they can be very responsive to changes in climate. Hence the start of the wet season (in tropical climates), or the end of ‘winter’ (in temperate Africa) initiates the reproductive process, and the young are born and weaned while the favourable conditions last. Within this reproductive period, many species show a ‘peak’ of reproductive activity (as measured by the percentage of adult females that are pregnant). In Beamys hindei, for example, pregnancy rate ranged from 20% to 60% during the nine months of reproductive activity (Fitzgibbon et al. 1995), and in Hylomyscus denniae in E DR Congo, pregnancy rate ranged from 80% (dry season/early wet season) to 9% (wet season) and then to 47% (late wet season/early dry season) (F. Dieterlen unpubl.). Larger rodents, with longer gestations (e.g. 100–120 days in Atherurus africanus, 112 days in Hystrix cristata, ca. 80 days in Pedetes capensis) are unable to respond so quickly to changing environmental conditions. Because of the short gestation of many species of small rodents, a female may have several litters during the time when conditions are favourable for reproduction. A strategy that includes short gestation, rapid growth of young to independence and a postpartum oestrus (so the female is able to conceive again immediately after parturition) is especially important to species where favourable conditions are short-lived and/or unpredictable. A corollary of small size is a short life-span (often only a few months) and hence it is necessary for small rodents to attain sexual maturity rapidly, to have large litters, and to produce several litters in quick succession in order to maximize their lifetime reproductive success. For Acomys wilsoni in Kenya (mean embryo number: 2.2), estimates suggest 7.9 litters/female/year and an annual production of 17.4 young/female/year (Neal 1983). Population structure within a species is a consequence of the characteristics of its reproduction and environment quality. Again, there are few long-term studies that document the changing structure of rodent populations. In Praomys tullbergi, a rainforest species that reproduces in most months of the year (with two peaks of reproductive activity) and has comparatively small litters, the population structure remains more or less constant throughout the year; young, subadults and adults are represented in all months (with the ratio of each changing slightly month by month) (Happold 1977). In contrast, the savanna-dwelling Mastomys natalensis reproduces only during the wet season and early dry season and has comparatively large litters; the population consists mostly of subadults at the end of the dry season, mostly adults and many young during the wet season and early dry season, and mostly subadults again (after most of the reproductive adults have died) in the mid- and late dry season. These examples (which probably represent the two extremes of population structure) also illustrate another aspect of demography in small rodents: species that occupy relatively constant and predictable environments throughout the year experience only small population fluctuations (e.g. the maximum number of Praomys tullbergi in Nigerian rainforest is only 2–3 times the minimum number; Happold 1977); whereas those in environments that are less predictable and have large seasonal variations in climate and resources exhibit larger fluctuations (e.g. the maximum number of Mastomys natalensis in

thornbush savanna is up to 33 times the minimum population size; Happold & Happold 1991). Because most species of rodents are small, only a limited volume of food can be ingested, and hence that food needs to be highly nutritious. The majority of species of African rodents feed on plants, and may be broadly categorized as herbivores (those feeding on grasses, leaves and/or herbs), granivores (seeds, nuts) or frugivores (fruit and flowers). Others are omnivores, feeding on plants and animals (usually arthropods), and a very few are wholly insectivores. For those species that have been investigated, the diet tends to be rather precise; preferred food items (i.e. those that are most nutritious in relation to their bulk) are selected, and most species alter their diet seasonally. For example, squirrels are primarily granivorous and frugivorous, although some species (e.g. Funisciurus lemniscatus, F. pyrropus, Myosciurus pumilio) are omnivorous because they also eat significant amounts of insects (mainly termites). Most glirids are omnivorous. Most species of Muridae are herbivorous or omnivorous, feeding on a variety of grasses, herbs, seeds and fruits; many are opportunistic and although basically herbivorous, will include a small amount of arthropod food in their diet when other foods are unavailable during some seasons of the year. A few species are wholly herbivorous (Arvicanthis spp., Otomys spp., Ctenodactylus spp.). Others are consistently omnivorous, with a high percentage of arthropods and worms in their diet (e.g. Acomys spp., Lophuromys spp., Uranomys ruddi, Heimyscus fumatus, Hybomys spp., Malacomys spp.). Only a few species are known to be wholly insectivorous (Deomys ferrugineus, Colomys goslingi). Subterranean species (species of Bathyergidae and Spalacidae) feed on a variety of underground corms, bulbs, tubers, fibrous roots and rhizomes. Perhaps because of their tropical and subtropical distributions, it is unusual for African rodents to store food, although there are some exceptions: some squirrels collect nuts, which are placed in caches; Beamys hindei, Cricetomys spp. and Saccostomus spp. carry seeds in their cheek pouches and store them in their burrows for later consumption; and some subterranean species store geophytes. The foregoing generalizations on food habits underscore the necessity of thorough, year-round, trophic studies of African rodents, especially for species in the families Nesomyidae and Muridae. The behaviour of African rodents has not been studied as extensively as that for the larger mammals.Their small size, nocturnal activity and lack of visibility make it very difficult to pursue behavioural studies on rodents (and other small mammals). However, there are several notable studies on diurnal species that are easily visible during the daytime (see, for examples, profiles on several species of Sciuridae, Rhabdomys pumilio, Thallomys spp., Pedetes capensis, Ctenodactylus gundi, Thryonomys spp. and Hystrix spp.). Additional studies (which provide information on sociality, agonistic behaviour, nesting behaviour and maternal behaviour) have been made on several species in captivity. Although most species tend to be asocial and/or solitary except for times during reproductive activity, some species form loose aggregations and social groups (e.g. squirrels, gundis, porcupines). All species of rodents that have been studied show many forms of intra- and inter-specific communication; these include vocal (e.g. calls, squeaks, whistles), visual (e.g. amicable or aggressive displays, postures, tail-flicking, etc.), non-vocal auditory (foot-drumming, tooth-chattering) and olfactory (scent-marking) signals.Territoriality is well-developed in some species and territories are maintained by visual and olfactory signals, and also (in some species) by fighting and chasing. Maternal behaviour has been well-studied (usually in 35

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captivity) in some species; most mothers are solitary when raising their young but there are interesting exceptions such as some species of Acomys (Muridae: Deomyinae) in which there is social care of the young by the several mothers. One of the most studied species, Heterocephalus glaber (Bathyergidae), has a social system resembling that of eusocial insects. Within any family, there may be some species that are asocial and territorial, and others that are highly social (see, e.g. Bathyergidae, Sciuridae, Muridae: Otomyinae). Rodents play an important role in ecological processes in Africa due to the many species present, their collectively widespread distributions and the sometimes large population numbers within particular habitats. Because the vast majority of species are herbivores and granivores, they assume a major role in recycling nutrients. Granivorous species transport seeds away from the parent plant and bury them, thus increasing seed dispersal. By their normal activities, rodents change the environment in which they live – a process known as ‘ecological engineering’; familiar examples include burrowing whereby the soil profile is altered and new soil is brought to the surface where it can act as ‘seed-beds’ for plants; defecating at regular latrines, which provides small areas of increased nutrients which promotes plant growth; and feeding on grasses and herbs, which promotes plant growth (in the same way as do larger herbivorous mammals). Although the influence of one individual rodent may seem trivial (compared with that of a mega-herbivore), the very large numbers of rodents in many ecosystems mean that their total influence is not inconsiderable. The only quantitative data available showed that in the tall grass plains of the Serengeti (where there are many large herbivores), small mammals utilized 69 kg/ha (= 1.2% of the total) of the annual grass biomass compared with 1122 kg/ha (=18.8% of the total) for the large mammals (Sinclair

1975). At times, rodents may be detrimental to human endeavours; some species (e.g. Mastomys spp., Gerbilliscus spp.) destroy seeds and crops in agricultural fields, and also feed on stored grains in houses and granaries. The role of rodents in African savanna and rainforest ecosystems has been reviewed by Happold (1983, 1996). Rodents are important prey for many small and medium-sized carnivores such as mongooses, genets, civets, small canids, small felids, owls (especially Tyto alba and T. capensis), hawks and snakes (see entries under ‘Predators, Parasites and Diseases’ in each species profile). For example, Giant Root-rats Tachyoryctes macrocephalus form 47% and Lophuromys melanonyx form 40% of the food of Ethiopian Wolves Canis simensis (Sillero-Zubiri & Gottelli 1995); at least 14 species of rodents form nearly 30% of the prey of seven species of small carnivores in the rainforests of the Central African Republic (the other significant prey was shrews, arthropods and herpetofauna) (Ray 1998); and small rodents formed almost all of the food of African Grass-owls Tyto capensis on Zomba Plateau, Malawi, where four species (Otomys angoniensis, Dasymys incomtus, Pelomys fallax and Dendromus nyikae) comprised the majority of the prey (Happold & Happold 1986). Humans are also important predators of some species such as Cricetomys emini, C. gambianus and Thryonomys spp. Some species of rodents are responsible directly (or indirectly through parasites that they carry) for the transmission of diseases of humans; well-known examples are some species of Gerbilliscus (formerly Tatera) and Gerbillus (plague), Meriones spp. and Psammomys obesus (leishmaniasis), Rattus rattus and R. norvegicus (plague) and Mastomys erythroleucus and M. natalensis (Lassa fever and other diseases). Control measures may be necessary when population numbers of these species become too large close to human environments.

Table 9. African families of the order Rodentia. Family

Length of taila

Tailb

Pelage texture

Cheekteeth

Jaw musculaturec

HB (typical size or range) (mm)d

Ecology and form

Anomaluridae

L

H

Soft

4

Hy

63–75, 280–380

Gliridae Petromuridae Sciuridae

L L L

H H H

Soft Soft Soft

4 4

/4 /4 4 /4, 5/4

Hy, My Hy Sc

mostly 60–100 140–220 110–126, 180–240

Dipodidae

L

T

Soft

3

Hy

98–118, 137–160

Pedetidae

L

T

Soft

3

Hy

350–430

Nesomyidae Cricetidae Muridae

L L L (some S)

P P P

3

/3 /3 3 /3

My My My

50–63, 270–300 100–124, 180–300 44–52, 135–190

Ctenodactylidae Bathyergidae Hystricidae Thryonomyidae Myocastoridae Spalacidae

S S S S S S or A

H P P P P P or NA

Soft Soft Soft (some coarse) Soft Soft Spines Coarse Coarse Soft

3

Hy Hy Hy Hy Hy My

125–230 90–190 650–850 400–790 470–570 160–200, 225–315

Arboreal; patagia present (except in Zenkerella) Arboreal; patagia absent Terrestrial, rupicolous Arboreal; patagia absent; one species 60–70 mm Terrestrial; elongated hindlimbs Terrestrial, elongated hindlimbs Terrestrial, some scansorial Terrestrial Terrestrial, some scansorial, some rupicolous Terrestrial, rupicolous Subterranean Terrestrial Terrestrial Aquatic; terrestrial Subterranean

/4

/3 /3

3

/3 (4/4) /4 (up to 6/6) 4 /4 4 /4 4 /4 3 /3 4

a

S = tail short 50% of HB; A = tail absent (or not visible externally). P = naked or with small bristles; H = covered with long hairs; T = well-developed tuft of hair at tip; NA = not applicable. c Hy = hystricomorphous; My = myomorphous; Sc = sciuriomorphous. d Approximate and typical sizes. Where there is wide variation within a family, the smallest and largest species are indicated. b

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Order RODENTIA

Systematic arrangement of Rodentia in Mammals of Africa

Size categories

The 15 families of African rodents (14 indigenous and the introduced Myocastoridae) are treated in accordance with the phylogenetic relationships reviewed above and as presented by Carleton & Musser (2005) in the Third Edition of Mammal Species of the World (Wilson & Reeder 2005).The families, in the sequence given in this volume (Table 6, see also Table 9), are Sciuridae, Gliridae, Dipodidae, Spalacidae, Nesomyidae, Cricetidae, Muridae, Anomaluridae, Pedetidae, Ctenodactylidae, Bathyergidae, Hystricidae, Petromuridae, Thryonomyidae and Myocastoridae. Four families (Spalacidae, Nesomyidae, Cricetidae, Muridae) are further divided into subfamilies, which are arranged alphabetically, as are the genera within each subfamily and the species within each genus.

In the Descriptions, an indication of the size of each species is recorded comparatively to other species in the family. The size categories, based on mean head and body length (HB), are: Family

Size categories (mean head and body length) Very small: 50–84 mm Small: 85–119 mm Medium-sized: 120–154 mm Large: 155–189 mm Very large: 190–350 mm Extremely large: >350 mm Small: 70–99 mm Medium: 100–129 mm Large: 130–159 mm Very small: 70–119 mm Small: 120–169 mm Medium sized: 170–219 mm Large: 220–269 mm Very large: 270–421 mm Small: 200 mm

Dipodidae, Spalacidae, Nesomyidae, Cricetidae, Muridae, Pedetidae, Ctenodactylidae, Hystricidae, Petromuridae, Thryonomidae, Myocastoridae Gliridae

Sciuridae, Anomaluridae 43

20

7

38 17

27 36

Bathyergidae

18

16 24

44 16

27

34

13

30

19

40

20 42 30 13

18

34

41

40 29

36 15

24 25

11

21

19

8 43 35

31

39 3 14

37

22 33

4 3

22

The length of the tail, expressed as a percentage of the length of HB, is described as very short (161%). The names of the bones of the skull are shown in Figure 6, and the nomenclature of the cusps on the cheekteeth of a murine rodent are shown in Figure 7.

28

Michael D. Carleton & D. C. D. Happold ANTERIOR

23

15

2

12

27 36

2

protocone

28 24

5

5 30

32

26

6 10

4

LINGUAL (inside)

M1

4

6 7

9

8

3

1

8

hypocone

1

Figure 6. The bones and terminology of the skull of a ‘typical’ murid rodent (Praomys tullbergi). 1 Angular process, 2 Anterior palatal foramen, 3 Auditory bulla, 4 Auditory meatus, 5 Basioccipital, 6 Basisphenoid, 7 Braincase, 8 Cheekteeth (molar row), 9 Condylar process, 10 Coronoid process, 11 Diastema, 12 Foramen magnum, 13 Frontal, 14 Glenoid fossa, 15 Incisor, 16 Infraorbital foramen, 17 Interorbital constriction, 18 Interparietal, 19 Jugal, 20 Lachrymal, 21 Masseteric knob, 22 Mastoid, 23 Mastoid process, 24 Maxilla, 25 Maxillary process (zygomatic arch), 26 Mesopterygoid fossa, 27 Nasal, 28 Occipital condyle, 29 Occipital, 30 Orbit, 31 Palate, 32 Palatine, 33 Paraoccipital process, 34 Parietal, 35 Posterior palatal foramen, 36 Premaxilla, 37 Pterygoid process, 38 Rostrum, 39 Squamosal process (zygomatic arch), 40 Squamosal, 41 Supraoccipital crest (if present), 42 Supraorbital ridge (if present), 43 Zygomatic arch (Zygoma), 44 Zygomatic plate.

M2

M3

4 7

LABIAL (outside)

metacone

3

1 15

9

paracone

5

6

8 9 1 3 4 5+6

7

8 9

POSTERIOR Figure 7. A schematic view of the upper left cheek teeth (M1, M2, M3) of a murine rodent showing the nomenclature of the cusps. Cusps labelled as 1, 2, etc. (= t1, t2, etc.). Inner (lingual) side on left, outer (labial) side on right. Cusps = black dots. Lines = lophs or ridges joining cusps. t2 on M2 and M3 never or rarely present. t3 of M3 very small or absent. t5 and t6 of M3 usually joined. Wear on occusal surface (with age and use) obliterates the cusp pattern.

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Family SCIURIDAE

Family SCIURIDAE SQUIRRELS

Sciuridae Fischer de Waldheim, 1817. Adversaria zoologica, p. 408. Allosciurus (1 species) Atlantoxerus (1 species) Epixerus (1 species) Funisciurus (10 species) Heliosciurus (6 species) Myosciurus (1 species) Paraxerus (11 species) Protoxerus (1 species) Sciurus (1 species) (introduced) Xerus (4 species)

Slender-tailed Squirrel Barbary Ground Squirrel Splendid-tailed Squirrel Rope Squirrels Sun Squirrels Pygmy Squirrel Bush Squirrels Giant Squirrel Grey Squirrel Ground Squirrels

p. 40 p. 42 p. 44 p. 46 p. 61 p. 70 p. 72 p. 89 p. 92 p. 93

The Sciuridae is a polygeneric family of rodents distributed throughout most of the Americas, Eurasia and Africa, in habitats ranging from alpine to tropical and from desert to rainforest. There are 51 genera and 277 species in the family (Thorington & Hoffmann 2005). The Sciuridae is divided into five subfamilies, with all African squirrels included within the subfamily Xerinae, which also includes all the North American and northern Eurasian ground squirrels. The squirrels are a very successful group, widespread on all continents except Australia and Antarctica. The present fauna of South America includes three genera and 19 species, North America has eight genera and 83 species, Europe has four genera and six species, and Asia has 22 genera and 92 species. Africa has ten genera and 37 species of squirrels, including the introduced Grey Squirrrel Sciurus carolinensis (subfamily Sciurinae), in South Africa. The anomalures (or scaly-tailed flying squirrels) of Africa belong to a different family of rodents, the Anomaluridae. Squirrels are small to large rodents, usually with a long well-haired tail and a relatively short rostrum. Some species are rather plain with a uniform reddish or brownish colouration, others are very colourful. Most squirrels have long tails, with long lateral hairs and shorter dorsal and ventral hairs; the tail may be slender or bushy and in some species has noticeable rings or bands of colour along its length. Longitudinal lateral side-stripes are common among terrestrial squirrels and some genera of arboreal squirrels. There are four digits on the forefoot (Digit 1 small or vestigial) and five long digits on the hindfoot. The skull is sciuromorphous, with the anterior deep masseter muscle originating on the zygomatic plate and (except in Paraxerus and Funisciurus; Thorington & Darrow 1996) the lateral surface of the rostrum. No part of the masseter muscle passes through the infraorbital foramen. Postorbital process present. Supraorbital ridge with small foramen (or notch) at anterior end in some genera.The masseteric tubercle, a bony knob-like projection on the maxilla bone anterior to the cheekteeth, is present in some genera and a useful diagnostic character. Dental formula is I /1, C 0/0, P 1/1 or 2/1, M 3/3 = 20 or 22.There are four upper cheekteeth (one premolar (= P4) and three molars) in Epixerus, most Heliosciurus, Myosciurus and Protoxerus, and five upper cheekteeth (two premolars (P3, which is often very small, and P4) and three molars in Allosciurus, Atlantoxerus, Funisciurus, Paraxerus and Xerus. Size categories of species in the family (based on mean head and body length) are given in the order Rodentia profile. Squirrels are terrestrial and arboreal, and one group of arboreal squirrels (the non-African Pteromyini) consists of species that are also able to glide. Most terrestrial species live in underground burrows, and

most arboreal squirrels make nests in tree hollows or on tree branches. Many squirrels feed extensively on hard seeds, but diets vary greatly from soft fruit or leaves to fungi, insects or other animal matter. Many non-African species of squirrels (e.g. Marmots, Chipmunks) hibernate, some for seven months each year. A few species aestivate during the hottest months of the year. Dwarfism has evolved independently three or four times in different groups of arboreal squirrels, in Africa, Asia and South America (Mercer & Roth 2003, Thorington et al. 1997). On all three continents, these diminutive squirrels have evolved a form of foraging known as bark-gleaning.The African Pygmy Squirrel Myosciurus is one of the smallest species, averaging 16 g in weight. The largest terrestrial squirrels are the Marmots Marmota, some of which may reach 9 kg. In contrast, the largest African ground squirrels, Xerus, do not exceed 1 kg. The largest tree squirrels are the Oriental Giant Squirrels Ratufa, with some individuals exceeding 2 kg. The largest African tree squirrels, Protoxerus, average 700 g. Most squirrels are solitary, with each individual having its own home-range. Some species are territorial. Colonial community structure and complex social behaviour occur in a number of terrestrial squirrels, especially the Prairie Dogs Cynomys, Marmots Marmota and Ground Squirrels Spermophilus of North America. Some African Ground Squirrels are also colonial (Atlantoxerus, Xerus inauris and Xerus princeps) but most African arboreal squirrels are not, and are seen singly, in pairs, or in small groups. Squirrels exhibit a variety of reproductive strategies, with average litter-sizes ranging from slightly more than one to more than nine. The largest litter-sizes are found among the North American ground squirrels, which have only a single litter per year; e.g. in Spermophilus, the average litter-size ranges from 3.3 to 9.3, and in Marmota from 3.4 to 4.2 (Heaney 1984). The largest of these may defer breeding until two or three years of age and produce litters every other year. North American tree squirrels have smaller litter-sizes (averaging 1.9–4.5 young), and commonly have more than one litter per year. In contrast, African Ground Squirrels have 2–4 young/litter (Moore 1961), and African tree squirrels have only 1–2 young/litter (Emmons 1979a). African tree squirrels are born at a more advanced stage of development and are ready to leave the nest at an earlier age than North American tree squirrels: e.g. the Paraxerus leave at 3–4 weeks of age, and young of Tamiasciurus and Sciurus at 5–9 weeks (Ferron 1984). Emmons (1979a) attributes this to a greater risk of nest predation in the African tropics. The earliest fossil squirrel is Douglassia of the late Eocene of North America (Emry & Thorington 1982). Squirrels (Palaeosciurus) appear in the European fauna in the early Oligocene (Vianey-Liaud 1974), immediately after the time known as the ‘grande coupure’, when many Asian species entered the European fauna. The African Ground Squirrels are first known from Heteroxerus in the late Oligocene of Europe. The genus Getuloxerus, perhaps synonymous with Atlantoxerus (the extant Barbary Ground Squirrel), is found in Spain and North Africa in the late Miocene (Black 1972a). Squirrels are first found in sub-Saharan Africa in the Miocene (Lavocat 1973). Volcanisciurus of the early Miocene is not referable to any modern tribe. Kubwaxerus of the upper Miocene is closely related to the extant genera Epixerus and Protoxerus and is the earliest known member of the Protoxerini

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Family SCIURIDAE

Table 10. Genera in the family Sciuridae. Genera

Posterior end of bony palate

Masseteric tubercle

Side-stripe on each flank

Mean HB* (mm)

Number of cheekteeth (upper/ lower jaw)

Mean GLS* (mm)

Notes

Allosciurus (1 sp.)

In line with posterior end of M3

Small

None

240

5

59

Atlantoxerus (1 sp.) Epixerus (1 sp.)

Considerably posterior to M3 Posterior to M3

Prominent

1 (+ single middorsal stripe) None

194

5

46

Terrestrial/arboreal. West African rainforests (Liberia/ Sierra Leone to Ghana) Terrestrial. NW Africa

284

4

69

Funisciurus (9 spp.)

Approximately in line with posterior end of M3

Not prominent

1 (sometimes obscure)a

152–211

5

38.8– 52.7

Heliosciurus (6 spp.)

Approximately in line with posterior end of M3

Prominent

None

196–237

4

42.7– 54.2

Myosciurus (1 sp.) Paraxerus (11 spp.)

Approximately in line with posterior end of M3 Approximately in line with posterior end of M3

Absent

None

66

4

21.3

Prominent

None, 1 or 2c

102–230

5

30.6– 55.6

Protoxerus (1 sp.) Xerus (4 spp.)

Approximately in line with posterior end of M3 Considerably posterior to M3

Not prominent Prominent

None

297

5

67.8

1d

212–259

5

50.7– 60.8

Not prominent

/4

/4 /4

/4

/4 b

/4 /4

/4 or 4/4 /4

Terrestrial and arboreal. Rainforests of West and central Africa Arboreal and tangles. Rainforests of West and central Africa; some spp. extend to relict forests in savanna Arboreal. Widespread. Rainforest and savanna of sub-Saharan Africa Arboreal. Rainforests SE Nigeria to Gabon Rainforest and tangles, woodland savanna. West, central and East Africa Arboreal. Rainforest of West and central Africa Terrestrial. Savanna and semi-arid; sub-Saharan Africa

* Where range of values given, smaller value refers to smallest species in genus and larger value to largest species in genus. a Except F. isabella, with two black stripes on each flank. b Except H. ruwenzorii – 5/4 with small P3 present. c Plus single mid-dorsal stripe in two species with two stripes on each flank (P. boehmi, P.alexandri). d Except X. rutilis.

(Cifelli et al. 1986). It is also the largest known African squirrel and is estimated to have weighed 1.4 kg. It had the thickest incisors of any known squirrel and probably, like Epixerus, subsisted on very thick nuts. Paraxerus from the Pliocene of East Africa is the earliest known of the Paraxerus–Funisciurus group of squirrels. Because of recent molecular studies (Mercer & Roth 2003, Steppan et al. 2004), the family Sciuridae is now divided into five subfamilies (Thorington & Hoffmann 2005); only one of these – the Xerinae – is represented in Africa. Within this subfamily there are three tribes: the Xerini, the Protoxerini and the Marmotini, the first two tribes containing all the African squirrels.The Xerini includes the two genera of African Ground Squirrels, Xerus and Atlantoxerus. The Protoxerini includes seven genera: the Sun Squirrels Heliosciurus, the African Giant Squirrel Protoxerus, the Slender-tailed Squirrel Allosciurus, the Splendidtailed Squirrel Epixerus, the African Bush Squirrels Paraxerus, the Rope Squirrels Funisciurus and the African Pygmy Squirrel Myosciurus. These tribal assignments, and the presence of Xerini in the fossil record of Europe, suggest that the Xerini invaded Africa from Eurasia. The Protoxerini are endemic to Africa and may have evolved there from an unidentified ancestor. The nine genera of endemic African squirrels (Table 10) are listed here alphabetically by genus; additionally there is a tenth genus, Sciurus, which has been introduced into South Africa. Allosciurus: Large unstriped arboreal squirrel of West Africa, with long slender non-bushy tail. Skull: rostrum curved down anteriorly,

supraorbital notch present, cheekteeth 5/4. Atlantoxerus: Ground Squirrel of North Africa.Three pale longitudinal stripes on back. Skull: bony palate extends well behind third molar, cheekteeth 5/4, masseteric tubercles prominent. (1 sp.) Epixerus: Large unstriped, terrestrial rainforest squirrel. Bushy tail with ventral V-shaped black and white pattern. Skull: bony palate ends in line with posterior end of third molar, cheekteeth 4/4, masseteric tubercles not prominent, skull sutures still prominent in adult between parietal bones and surrounding interparietal bone. (1 sp.) Funisciurus: Mostly small forest squirrels, often with a side-stripe on each side of body. Skull: bony palate ends in line with posterior end of third molar, cheekteeth 5/4, cheekteeth flat-crowned, fossa for origin of anterior deep masseter muscle does not extend onto the rostrum. (10 spp.) Heliosciurus: Medium to large arboreal forest squirrels of sub-Saharan Africa. Skull: bony palate ends in line with posterior end of third molar, cheekteeth 4/4, third premolar absent in all but one species. Prominent masseteric tubercles, fossa for origin of anterior deep masseter muscle extends onto the rostrum. (6 spp.) Myosciurus: Pygmy tree squirrel. Length of head and body less than 7.5 cm. Skull: bony palate ends in line with posterior end of third molar, cheekteeth 4/4. skull length less than 25 mm, body mass less than 20 g. (1 sp.) Paraxerus: Diverse group of forest and bush squirrels, some species with side-stripes, some without side-stripes. Skull: bony palate 39

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ends in line with posterior end of third molar, cheekteeth 5/4, fossa for origin of anterior deep masseter muscle does not extend onto the rostrum, mandibular cheekteeth cuspidate, upper cheekteeth flat-crowned. (11 spp.) Protoxerus: One of the largest arboreal forest squirrels of subSaharan Africa. Tail not bushy. Skull: bony palate ends in line with posterior end of third molar, cheekteeth 4/4, masseteric tubercles not prominent, fossa for origin of anterior deep masseter muscle extends onto the rostrum, skull sutures fused in adult between parietal bones and surrounding interparietal bone. (1 sp.)

Sciurus: Exotic species found only in parts of South Africa. No stripes on back. Skull: bony palate extends well behind third molar; cheekteeth 5 /4, masseteric tubercles not prominent, fossa for origin of anterior deep masseter muscle does not extend onto the rostrum. (1 sp.) Xerus: Ground squirrels of sub-Saharan Africa. Bristly fur; most species have a white side-stripe on each side of body. Skull: bony palate extends well behind third molar, cheekteeth 5/4, masseteric tubercles prominent. (4 spp.) Richard W. Thorington, Jr

GENUS Allosciurus Slender-tailed Squirrel Allosciurus Conisbee, 1953. A list of Names proposed for Genera and Subgenera of Recent Mammmals: 6.

Monotypic genus. A large squirrel confined to the Rainforest BZ (Western Region). The characters of the genus include comparatively large size (mean HB 240 mm), back and flanks without longitudinal side-stripes, and long slender hairy tail (mean 298 mm), which is longer than HB (cf. Protoxerus). Females have four pairs of nipples; presence of baculum in !! not known. The skull is characterized by the curved-down rostrum (in lateral view), supraorbital notch (on supraorbital ridge at lateral side of frontal bone) open above the orbit (not a foramen), infraorbital foramina large and oval, masseteric knob small, five upper cheekteeth and auditory bullae well developed, extending ventrally well below the line of the cheekteeth. The single species, Allosciurus aubinnii, is frequently considered as belonging to Protoxerus, but its many unique features indicate that it should be placed in its own genus (see Thomas 1909a, Ellerman 1940, 1941, Rosevear 1969, Grubb et al. 1998; contra Moore 1959, Amtmann 1966,Thorington & Hoffmann 2005). Allosciurus differs from Protoxerus and Epixerus (and from Heliosiurus where indicated) in a number of important ways: (1) skull with unusual shape, relatively high, strongly convex in nasofrontal region with down-curved rostrum (cf. not downcurved in Protoxerus, Epixerus and Heliosciurus); (2) cheekteeth always 5/4

(cf. 4/4 in Heliosciurus, though 5/4 in Heliosciurus ruwenzorii and rarely in some Epixerus specimens); (3) skull length 65 mm); (4) supraorbital notch present (cf. closed to form supraorbital foramen – and in Heliosciurus) (Figure 8); (5) no sharp boundary between hair of flanks and underparts (cf. sharp boundary between hairs of flanks and relatively hairless underparts); (6) tail dark-coloured, with long black tips to hairs (cf. tail marked with dark and pale bands – and in Heliosciurus); (7) colouration dark (cf. relieved by light speckling and/ or reddish tones – and in Heliosciurus). Allosciurus forms a monophyletic clade with Heliosciurus, Protoxerus and Epixerus (Moore 1959).

Allosciurus aubinnii.

Figure 8. Skull and mandible of Allosciurus aubinnii (BMNH 45.331).

Peter Grubb

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Allosciurus aubinnii

Allosciurus aubinnii SLENDER-TAILED SQUIRREL (AUBINN’S SQUIRREL) Fr. Écureuil d’Aubinn; Ger. Dünnschwanzhörnchen Allosciurus aubinnii (Gray, 1873). Ann. Mag. Nat. Hist., ser. 4, 12: 65. Fanti, Ashanti Province, Ghana.

Taxonomy Originally described in the genus Myrsilus. Rosevear (1969) considers there to be sufficiently important differences in the skull and pelage for this species to be placed in its own genus, Allosciurus, separate from Protoxerus. Grubb et al. (1998) also place the species in Allosciurus (see genus profile). In contrast to Protoxerus, Allosciurus has no trace of red or rufous in the pelage, the tail lacks any white rings or banding, and the rostrum is downcurved. Hoffmann et al. (1993) and Thorington & Hoffman (2005) retain the species in the genus Protoxerus, but invoke Allosciurus as a subgenus. Synonyms: salae. Two subspecies. Chromosome number: not known. Description Large dark slender squirrel without any other special colouring or marking. Dorsal pelage, head and limbs, dark brown to brownish-black, slightly speckled with yellow or buff; hairs brownish-black with 2–3 narrow, pale yellow bands. Ventral pelage similar but paler than dorsal pelage. When viewed from side, head has rounded profile due to down-curved rostrum. Ears short and rounded with dense hairs on outer surface. Tail long (ca. 125% of HB); slender and not bushy; black with intermixture of yellow or buff towards tip; hairs long (ca. 35 mm), mostly black at base, and with wide pale bands towards tip. Skull heavily built but less so than in P. stangeri and Epixerus ebii; rostrum down-curved, supraorbital ridge with open notch; cheekteeth 5/4; posterior end of bony palate in line with posterior end of upper M3; masseteric tubercle not prominent (see also genus profile). Nipples: 1 + 1 + 1 + 1 = 8. Geographic Variation P. a. aubinnii: Ghana, Côte d’Ivoire. Pelage as described above. P. a. salae: Liberia (and probably Sierra Leone – Grubb et al. 1998). Very dark form; hairs along mid-dorsal line without bands; hairs of tail without bands except at tip. Similar Species Epixerus ebii. Pelage contains rufous colouration; tail long and bushy, flecked with white. Protoxerus stangeri. Grizzled white pelage on head and chest; large bushy tail liberally flecked with white. Distribution Endemic to Africa. Rainforest BZ (Western Region). Recorded from Sierra Leone (near the Liberian border), Liberia, Côte d’Ivoire and Ghana. Habitat Moist primary rainforests, often associated with Raphia palms (G. S. Cansdale in Rosevear 1969). Maybe partly a groundliving squirrel (G. S. Cansdale in Grubb et al. 1998).

Allosciurus aubinnii

Abundance Usually rare or uncommon, but may be as common as P. stangeri in some moist mature rainforests (Booth 1960). Remarks One stomach contained the husks of Raphia palms (G. S. Cansdale in Rosevear 1969). Probably solitary. Conservation

IUCN Category: Data Deficient.

Measurements Allosciurus aubinnii HB: 240 (230–250) mm T: 298 (270–310) mm HF: 58 (55–61) mm E: 20 (18–21) mm WT: n. d. GLS: 59.3 (59–60) mm GWS: n. d. P3–M3: 11.1 (10.8–11.4) mm Ghana (for P. a. aubinnii) (Rosevear 1969) Sample size not given Key References

Grubb et al. 1998; Rosevear 1969.

Chad E. Schennum & Richard W. Thorington, Jr

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Family SCIURIDAE

GENUS Atlantoxerus Barbary Ground Squirrel Atlantoxerus Forsyth Major, 1893. Proc. Zool. Soc. Lond. 1893: 139. Type species: Sciurus getulus Linnaeus, 1758.

The genus Atlantoxerus was originally a subgenus of Xerus, and later raised to full generic rank by Thomas (1909c). Closely related to Xerus, it presently comprises only one species, which occurs in NW Africa, originally described as Sciurus getulus. The single species in the genus mainly inhabits rocky areas in semi-arid and arid regions where they rely on natural shelters or self-dug burrows to escape the extremes of climate. Placed in the Tribe Xerini (together with Xerus, the only other genus of ground squirrels). Barbary Ground Squirrels are medium-large squirrels with a striking bushy tail, relatively large eyes and small ears (without hair brush). The head tends to be blunt and round, with powerful muscles. The nostrils are narrow and naked, and can be closed by the tip of the muzzle. In contrast to other Palaearctic ground squirrels, the skull is not angular, has a strong sagittal crest, and the palate extends posteriorly to the cheekteeth. The hindfeet have naked soles and slender digits. The claws, adapted to digging, are long and only slightly curved. Most features are similar to those of the genus Xerus, but the fur is not as bristly, although it is short and stiff. Dental formula: I 1/1/, C 0/0, P 2/1, M 3/3 = 22. The skull is characterized by two upper premolars (P3 is minute, sometimes absent) on each side of the upper jaw, each upper incisor frequently has traces of a groove, and molar surfaces are slightly concave with transversal ridges (M1 and M2 are lacking mesostyle, lower molars without anteroconid), palate long (ca. 62% of GLS) with the posterior end of bony palate well posterior to M3 (as in Xerus and unlike other sciurids); masseteric tubercle very prominent (Figure 9). The baculum is long and narrow, ends in a spatulate enlargement with a small crest above and a larger one, bent to the left, below (Cabrera 1932).

The genus Atlantoxerus was first identified with certainty from the early Miocene of China (Li & Qiu 1980, Qiu et al. 1999) and middle Miocene of Arabia (Sen & Thomas 1979) Later, it differentiated and spread quickly over southern Europe (Spain; Pelaez-Campomanes 2001), North Africa (Morocco; Lavocat 1961) and eastern Asia (China; Wu 1988). In the upper Miocene, at least four species are known from Spain (De Bruijn & Mein 1968), France (Aguilar et al. 1995), Algeria (Ameur 1988), Morocco (Geraads 1998) and Rhodes (De Bruijn et al. 1970). In Morocco, several specimens related to A. getulus have been reported from the lower and middle Pleistocene (Jaeger 1975, Michel 1990). However, the origin of A. getulus (the only extant species of the genus) remains unknown; it does not seem to be a descendant of any known extinct species of the genus that were present in the Maghreb at earlier periods. Jaeger (1977a) suggested a close relationship with the Spanish fossil-form A. androveri (De Bruijn & Mein 1968). Additional fossils from the Pliocene are needed to clarify the evolution of the genus, as well as genetical studies on the tribe Xerini to elucidate the relationships of the various genera. Stéphane Aulagnier

Atlantoxerus getulus. Figure 9. Skull and mandible of Atlantoxerus getulus (BMNH 22.5.30.20).

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Atlantoxerus getulus

Atlantoxerus getulus BARBARY GROUND SQUIRREL Fr. Ecureuil fouisseur de Barbarie; Ger. Nordafrikanischer Erdhörnchen (Atlashörnchen) Atlantoxerus getulus (Linnaeus, 1758). Syst. Nat., 10th edn., 1: 64. ‘in Africa’. Restricted by Thomas (1911) to ‘Barbary’, and by Cabrera (1932) to Agadir, Morocco.

Taxonomy Originally described in the genus Sciurus. Synonyms: praetextus, trivittatus. Subspecies: none. Chromosome number: 2n = 38, aFN = 68. Description Medium-sized ground-dwelling squirrel with a short pale greyish-brown dorsal pelage and three pale stripes along the body. Pelage short (ca. 3–5 mm) and slightly coarse. Dorsal pelage greyish-brown; hairs pale brown or pale buff terminally, sometimes with black tip. Long black guard hairs, especially on midback and rump. Single white side-stripe on each flank from behind shoulders to mid-rump (not to base of tail), bordered on each side by darker background colouration. Single less obvious pale stripe on mid-dorsal line, shorter in length than white side-stripes on flanks. Elongated patch of whitish hair on lower flank between limbs.Ventral pelage sparse; hairs white, sometimes grey at base. Head is rounded and similar in colour to back. Ears small, eyes with narrow pale eyering. Limbs moderately long, similar in colour to body. Forefoot with four long digits, each with sharp claw; Digit 1 reduced to small tubercle without claw. Hindfoot with five digits; Digit 1 short; all with long curved claws. Tail of moderate length (ca. 80% of HB), hairs long (ca. 25–30 mm), creamy-white with two wide black bands equidistant along length of hair giving a black–white banded pattern; tip white. Young differ from adults by having longer and smoother pelage. Skull: cheekteeth 5/4; P3 very small and peg-like, sometimes absent; posterior end of bony palate well posterior to M3; masseteric tubercle prominent. Nipples: 1 + 1 + 1 + 1 = 8. Geographic Variation None recorded. Similar Species Xerus erythropus. Dorsal pelage brown, harsh, marked with a single short stripe on each side; muzzle longer with a projecting nose; distribution overlaps only in the Souss plain (Morocco) (Blanc & Petter 1959). Distribution Endemic to NW Africa. Mediterranean Coastal BZ. Recorded in Morocco and a small area of NW Algeria (Ksours Mts); known from the coastal zone up to 4165 m, mainly in the Middle and Grand Atlas south to Agadir, the Anti-Atlas, and the northern edge of the western Sahara south to Sequiat el Hamra. Introduced (extralimitally) to Fuenteventura (Canary Is.) in 1965. Habitat Stony regions and open habitats, from mountain slopes to desert. Common in open country with scattered trees and bushes of Juniper Juniperus spp., Thuya Tetraclinis articulata and Argan Argania spinosa, but avoiding areas devoid of vegetation and forests. Also present in various agricultural habitats, favouring stone walls, which provide refuge. Not recorded in irrigated fields. Abundance Abundant on the lower slopes and valleys. Very common up to 2000 m in the Grand Atlas; but then becoming less

Atlantoxerus getulus

common with increasing altitude. Population numbers also decrease dramatically in the eastern part of the range. Maybe locally numerous in some desert habitats. Fluctuations of population numbers are suspected; for example, they were very numerous in 1952 and rare in 1971 in the same region (Saint Girons 1974). Adaptations Diurnal and terrestrial. Exhibits two periods of activity: in the morning from 07:00h to 11:00h with a peak of activity at about at 09:00h, and in the afternoon from 14:00h to 18:00h with a peak at 15:00h. Optimal temperature for activity is 24 °C. The activity rhythm observed in the Toubkal massif was similar to that at Ademine forest and Jbel Bani at lower altitudes (Petter & Saint Girons 1965). Shelters in burrows during the night and during the heat of the day. Burrows are excavated under rocks, or among stones in screes that have been consolidated by vegetation. Requires permanent water in the southern part of its distribution. At high altitudes, daily activity is reduced greatly during winter, even in the absence of snow cover. True hibernation has not been recorded. Foraging and Food Mainly herbivorous, sometimes omnivorous. Forages on acorns, nuts and seeds, including those of Argan, but also fruits of wild olive-trees Oliva europaea, Juniper and Thuya (Cabrera 1932), and fruits and seeds of Pistachio trees Pistacia atlantica. The diet may also include grasses and roots (Carpentier 1932), and also domestic crops, where fallen seeds and stems are eaten (Brosset 1960). Occasionally feeds on insects (Cabrera 1932), and on rubbish in small villages. 43

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Family SCIURIDAE

Social and Reproductive Behaviour The simplest family unit is composed of a single " with her young. Two "" with their litters may share a common shelter, at least when the young are able to move around the shelter independently. Pairs of "" are common before the reproductive season, and sharing of a common natal nest is suspected. These family units may form loose colonies. More complex social groups, including at least three lactating "" with their young and an adult ! have been observed (Gouat & Yahyaoui 2001). Group composition differs greatly (especially during the reproductive season), even between sites that are geographically close together. During the mating period !! usually stand on a prominent site during the day and call for "". Several !! can court one ", chasing each other. Records of matings are very rare, and matings are suspected to occur in burrows. Reproduction and Population Structure Reproductive season varies geographically. Females pregnant in E Morocco in April (Brosset 1960), by which time young have already been born in Saharan Atlas (Cabrera 1932) and young have been weaned in Western Sahara (Valverde 1957). At higher altitudes (Middle Atlas, Grand Atlas, Anti-Atlas), young born in July. Two litters/year may be possible in the Tazenakht region (Anti-Atlas). Litter-size: up to four; in Canary Is., litters of up to nine have been recorded.Young weigh 6–9.5 g at birth, and they leave the burrow when 5–6 weeks old. Captive animals have lived for five years.

Predators, Parasites and Diseases Main predators are diurnal raptors (occasionally owls), and also foxes and genets. Many parasites have been collected (Dollfus 1953, Hastriter & Tipton 1975); the louse Neohematopinus pectinifer is reported to carry spirochetes, causing relapsing fever in humans (Blanc et al. 1933). Conservation IUCN Category: Least Concern. Widespread within its range, and not threatened. Measurements Atlantoxerus getulus HB: 194 (165–230) mm, n = 18 T: 158 (90–190) mm, n = 18 HF: 46.5 (41–52) mm, n = 18 E: 15.5 (12–18) mm, n = 16 WT: 250 g, n = 1 GLS: 45.9 (38.4–50.0) mm, n = 13 GWS: 27.9 (23.6–30.8) mm, n = 13 P4–M3: 8.8 (7.2–9.7) mm, n = 12 Morocco (MNHN) Key References Gouat & Yahyaoui 2001; Petter & Saint Girons 1965; Saint Girons 1953, 1974. Stéphane Aulagnier, Patrick Gouat & Michel Thévenot

GENUS Epixerus Western Palm Squirrel Epixerus Thomas, 1909. Ann. Mag. Nat. Hist., ser. 8, 3: 472. Type species: Sciurus wilsoni Du Chaillu, 1860 (= Sciurus ebii Temminck, 1853).

Epixerus contains a single species distributed in the Rainforest BZ of West and Equatorial Africa. The genus is distinguished by its large size, long hindfoot (usually over 55 mm) and tapered bushy tail with strongly banded hairs that produce a splendid pattern of chevrons when spread and viewed from below. Skull characteristics are large size (>65 mm), palate extends well posteriorly to the posterior end of M3, ratio of palatal length to interorbital breadth is 56.0% to 65% (Amtmann 1966), masseteric tubercle absent or not prominent, and supraorbital ridge has a foramen (Figure 10). Dental formula: I 1/1, C 0/0, P 1/1, M 3/3 = 20. Additional characters are given in the species profile. The number of species in the genus is debatable. Two species are sometimes recognized (e.g. Verheyen 1959, Hoffman et al. 1993): E. wilsoni (du Chaillu, 1860), with a subspecies E. w. mayumbicus, and E. ebii (Temminck 1853), with two subspecies E. e. ebii and E. e. jonesi. Kuhn (1964) revised the genus and, on the basis of a large geographic sample, recognized a single species, E. ebii (see also Amtmann 1966, Thorington & Hoffman 2005). Here, a single species, E. ebii, is recognized. Louise H. Emmons Figure 10. Skull and mandible of Epixerus ebii (BMNH 98.3.19.12).

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Epixerus ebii

Epixerus ebii WESTERN PALM SQUIRREL Fr. Écureuil des palmier; Ger. Westliches Palmenhörnchen Epixerus ebii (Temminck, 1853). Esquisses zoologiques sur la côte de Guiné, p. 129. ‘Les grandes fôrets de la Guiné’. Ghana.

Taxonomy Originally described in the genus Sciurus (see also genus profile). The form wilsoni was formerly considered as a distinct species (e.g. by Perret & Aellen 1956, Verheyen 1959 and Rosevear 1969), but is here considered as a subspecies. Synonyms: jonesi, mayumbicus, wilsoni. Subspecies: three. Chromosome number: not known. Description Very large, reddish, slender-bodied squirrel with long legs and long, bushy, grey-frosted tail. Pelage short. Dorsal pelage dull brownish-red, finely grizzled with yellow and black. No side-stripes on flanks.Ventral pelage sparse with some whitish hairs on chest; yellow ventral skin sharply delineated from lateral pelage. Head large and broad, with large eyes; ears relatively large, naked, extending above crown. Bulging cheeks (due to large highly developed masseter muscles). Limbs and feet long and slender, similar in colour to dorsal pelage. Tail long (ca. 95% of HB), bushy, tapered at tip, frosted grey, with indistinct alternating bands of blackish and pale grey. Alternating bands of red, black and grey on the tail hairs create a pattern of chevrons on the underside of the tail when hairs are spread and viewed from below (not normally visible in the field).Tail carried horizontally, straight out behind the body, or hanging, not curled up against back. Skull: cheekteeth 4/4; posterior end of bony palate posterior to M3; masseteric tubercle not prominent. Nipples: 1 + 1 + 1 + 1 = 8; one specimen with five pairs = 10 was reported in Liberia (Kuhn 1964). Geographic Variation E. e. jonesi: Liberia and Sierra Leone. Head and dorsal pelage rufous. E. e. ebii: Ghana and Côte d’Ivoire. Head rufous, dorsal pelage browner.

Epixerus ebii.

E. e. wilsoni: Cameroon, Gabon, Equatorial Guinea and Congo. Head and dorsal pelage reddish-brown. Similar Species Protoxerus stangeri. Larger, greyer, with stout body, long back, short robust legs; head grizzled grey; short ears that do not extend above crown; arboreal. Distribution Endemic to Africa. Rainforest BZ (Western and West Central Regions). Recorded from Liberia and Sierra Leone to Ghana; S Cameroon, Gabon, Equatorial Guinea and Congo. Habitat Evergreen rainforests. Undergrowth in forest, including Raphia palms. In Sierra Leone, recorded from montane forest at 1020 m (Grubb et al. 1998). Abundance Rare in collections and rarely seen; but in Gabon it is more common than usually supposed. Its terrestrial habits, extreme wariness and tendency to escape silently on the ground at the slightest disturbance, allow it to exist undetected by Western naturalists (who often mistake it for a Forest Giant Squirrel Protoxerus stangeri). None the less, local peoples who hunt game with nets know this species well and have specific names for it. The very short activity period (see below) may contribute to the perceived rareness of these squirrels.

Epixerus ebii

Adaptations Diurnal. A cursorial, fast-running terrestrial species with elongated, slender limbs and feet, and a slender lower abdomen; 45

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sometimes assumes a digitigrade posture with forelimbs bearing weight only on the tips of the digits. The masseter muscles are highly developed and the incisors are robust, consistent with a diet of kernels from hard nuts. A prominent glandular pocket in the inside corner of the lips may be used in scent marking (Emmons 1975, 1980). Western Palm Squirrels make dens in tree hollows and favour those hollows with small entrances (mean height: 8.1 m [1.5–20 m]; n = 12 den sites; Emmons 1975).The nest itself is undescribed.They leave the nest at dawn; in Gabon they have a short activity period (mean 7.69 h) and return to the nest in the late afternoon or well before dusk (mean entry time 14:50h; range: 12:30–17:03h, n = 16; Emmons 1980). Climbing to the nest is their only arboreal activity. Foraging and Food Feeds terrestrially on fallen seeds and fruit, especially on thick-walled nuts (98% dry mass, n = 4 stomach contents; Emmons 1980). A few insects are also consumed. To feed on nuts it jumps up from the ground onto a low feeding perch (0.5– 1.5 m), probably to scan for predators while deafened by gnawing. In Gabon, feeds mostly on nuts of Panda oleosa, a large, elephantdispersed fruit. The presence of this squirrel in an area can be readily ascertained by the presence of large middens of shells from opened Panda nuts beneath low perches: the seeds are extracted by either splitting along the lines of dehiscence (old nuts) or by the extraordinary feat of ‘sawing’ nuts in half across the middle (fresh nuts). No other animal opens Panda nuts by this method. Western Palm Squirrels hoard Panda nuts by burying them in the ground at distances of 15–20 m or more from the parent tree (Emmons 1975), thereby assisting the dispersal of the seeds. Social and Reproductive Behaviour In 50 observations, squirrels were solitary 80% of the time and seen with 1–3 others in the remainder of sightings (Emmons 1980). Two adults feeding under the same Panda tree tolerated each other and showed no aggression. Two adult !! followed by radio-tracking in Gabon had home-ranges of 21.1 ha and 21.7 ha, and a single subadult " had a home-range of 13.9 ha. Movement is rapid: !! travel at a mean rate of 115 m/h and "" at 144 m/h (means of all hours when active). The social organization is unknown, but in one apparent ‘mating chase’ several calling !! pursued a presumably oestrus " (Emmons 1980).

Vocalizations are described in detail in Emmons (1978). The low intensity alarm call consists of soft, short (about 400 ms), rapid, machinegun-like bursts of chattering of the incisor teeth. High intensity alarm is indicated by a staccatto series of short, broadfrequency pulses. When alarmed, a squirrel may climb to a low vantage point on a horizontal or vertical support, with the long bushy tail hanging straight downward. At the start of each high intensity alarm call, the base of the tail is wagged rapidly and vigorously from side to side a few times, the tail-tip following passively, whipping loosely back and forth below the body. Squirrels calling from the ground hold the tail horizontally, with no wagging display. Reproduction and Population Structure litter-size is 2 (n = 1, Emmons 1979a). Predators, Parasites and Diseases local people in hunting nets.

The only published

Sometimes captured by

Conservation IUCN Category: Least Concern. Likely to be threatened only by loss of mature rainforest. Homeranges are large so that the density of squirrels is always relatively low. Measurements Epixerus ebii HB: 284 (275–300) mm, n = 7 T: 272 (250–310) mm, n = 5 HF: 68 (65–70) mm, n = 5 E: 20–21 mm, n = ?* WT: 594 (540–650) g, n = 7 GLS: 69.4 (68.6–69.8) mm, n = 4 GWS: 36.9 (35.9–37.7) mm, n = 3 P4–M3: 10.8 (10.3–11.2) mm, n = 7 Gabon (Emmons 1975, L. Emmons unpubl.) *Rosevear 1969 Key References Emmons 1975, 1978, 1980; Kuhn 1964. Louise H. Emmons

GENUS Funisciurus Rope Squirrels Funisciurus Trouessart, 1880. Le Naturaliste 2 (37): 293. Type species: Sciurus isabella Gray, 1862.

A genus of small- to medium-sized squirrels with ten species endemic to the Rainforest BZ and parts of the adjacent Rainforest–Savanna Mosaics (Table 11).The greatest diversity is in west-central Africa, with five species in Cameroon. Most species are arboreal; some species live in the canopy in natural rainforest, and others live in tangled vegetation close to the ground and in secondary growth. A few species enter plantations and gardens. The vernacular name ‘Rope Squirrel’ refers to the ability of these squirrels to clamber and climb on rope-sized branches and lianas. An alternative vernacular name ‘African Striped Squirrel’ refers to the lateral side-stripe(s) on each flank.

The genus is characterized by relatively small size (mean HB usually 130–200 mm), long well-haired tail slightly shorter than the head and body, and (in most species) a longitudinal side-stripe along each flank. The species of the genus are the smallest of African squirrels, except for the single species of Myosciurus (HB 61–74 mm). The head is small, with small ears held close to the skull; the limbs are short, and the long tail is covered with long hairs that are held erect to give a bushy appearance. The tail is commonly held upwards and backwards, close to the back with the tip curling away from the shoulders; it is constantly flicked backwards and forwards as a means of

46

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Funisciurus substriatus.

Funisciurus pyrropus.

Funisciurus congicus.

Funisciurus squirrels.

communication and when the squirrel is agitated. Females have either two pairs of nipples (most species) or four pairs (F. leucogenys). Rope Squirrels produce a range of vocal noises, including clicks, squeaks and chatters. The skull is characterized by the dental formula I 1/1, C 0/0, P 2/1, 3 M /3 = 22; orange ungrooved incisors, five (not four) cheekteeth in

each upper ramus (cf. Heliosciurus); small masseteric tubercle; fossa for origin of anterior deep masseter muscle does not extend onto the rostrum; supraorbital foramen absent; and the posterior end of the bony palate is approximately in line with the posterior end of M3 (in this latter respect it is similar to several other genera of squirrels but different from Atlantoxerus, Xerus and Epixerus where it 47

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Figure 11. Skull and mandible of Funisciurus anerythrus (BMNH 0.2.5.33).

is posterior to M3). Cusps of cheekteeth are worn down early in life, leaving flattened surfaces (not concave with prominent ridges as in Heliosciurus) (Figure 11). The genus exhibits considerable diversity in morphology, especially in colour and in the form of the lateral side-stripe. Species in the genus range from small black-striped squirrels to somewhat larger squirrels with successive reduction of the stripes on each side. The gradual reduction in number and size of the side-stripes is illustrated in the following sequence: (a) F. isabella, F. lemniscatus (smaller species; two black stripes); (b) F. congicus (smaller species; stripe pattern slightly modified to a blackish-brown side-stripe bordered by a cream stripe); (c) F. bayonii, F. substriatus (relatively small species; stripes reduced to a pale lateral side-stripe); (d) F. pyrrhopus, F. anerythrus (larger species, reddish pigment in many subspecies; pale side- stripe, ofen rather obscure); (e) F. leucogenys (largest; stripes reduced to line of whitish spots); and (f) F. carruthersi (relatively large, no side-stripe). See Rosevear (1969) for a full discussion of the genus. The genus is placed in the subfamily Xerinae and tribe Protoxerini (which also includes all the other African sciurids except Atlantoxerus and Xerus – the Ground Squirrels – which are placed in the tribe Xerini) (Thorington & Hoffman 2005). Species in the genus are distinguished by colour, number and character of the lateral side-stripes, colouration of body and limbs, and body and skull size (Table 11). Peter Grubb

Table 11. Species in the genus Funisciurus. Arranged in order of increasing number of side-stripes on each flank. Notes

Species

Side-stripes on each flank

Red colouration on pelage

General colouration

HB mean (mm)

HF mean (mm)

GLS mean (mm)

F. carruthersi

Absent

None

Olive-green

203

48

48

F. anerythrus

One, pale

None

Brown

182

44

47

F. bayonii

One, pale, usually indistinct

None

Dull olive, speckled black

ca. 180

42

42

None

Yellowish-brown

160

40

33

None

Greenish-yellow

ca. 172

43

44

Forelimbs and hindlimbs

Grey-black

211

46

52

Head and cheeks

Grey-brown

209

51

51

None

Grizzled brown

35–36

41

Rainforests of central Africa

None

Brown

43

Rainforests of central Africa

None

Olive-brown

42

Gabon

F. congicus

F. substriatus F. pyrropus F. leucogenys F. isabella F. lemniscatus F. duchaillui

One, wide white bordered ventrally by brownish-black One, pale white bordered ventrally by dark stripe One, white One, whitish-yellow broken into spots Two, black, separated by buffy-brown Two, black separated by yellow Two, black

158 !! 152 "" 164 !! 160 "" 197

36 !! 35 "" 45

Montane forests, Rwenzori region Rainforests, Nigeria to W Uganda NE Angola, SW DR Congo south of Kasai and Congo rivers. Rare Rainforests of DR Congo (south of river), Angola, Namibia Guinea and Sudan Savanna, Burkina to Benin Rainforests, Senegal to DR Congo and Rwanda Rainforests, Ghana to Central African Republic

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Funisciurus anerythrus

Funisciurus anerythrus THOMAS’S ROPE SQUIRREL (REDLESS TREE SQUIRREL) Fr. Funisciure de Thomas; Ger. Rotloses Baumhörnchen Funisciurus anerythrus (Thomas, 1890). Proc. Zool. Soc. Lond. 1890: 447, pl. 40. Buguera, south of Lake Albert, Uganda.

Taxonomy Originally described as a subspecies of Sciurus pyrropus in 1890 and later raised to specific rank.There has been great confusion about the taxonomic placement of the named subspecific forms of F. anerythrus and F. pyrropus; several subspecies have ping-ponged back and forth between the two species. In Gabon, where the local forms of the two species are sympatric, they are unequivocally distinguished by their physical characters and their entirely distinctive calls, as well as by major ecological differences in habitat, diet and nest sites (Emmons 1975, 1980).These distinctions, described below, should be considered when assigning West African forms to one of these two species. Synonyms: bandarum, mystax, niapu, ochrogaster, raptorum. Subspecies: four. Chromosome number: 2n = 38, FN = 62 (Benin, Civitelli et al. 1996). Description Medium-sized plain brown squirrel with a pale sidestripe from shoulder to rump. Dorsal pelage grizzled brown flecked with buff; hairs banded. Single pale buff side-stripe from shoulder to hindquarters, bordered below with dark brown. Ventral pelage thickly haired, dull grey to white or orange. Head similar to dorsal pelage. Eye bordered above and below with pale buff line. Ears small. Fore- and hindlimbs dull buff; digits with well-developed claws. Tail medium (ca. 85% of HB – relatively short for a squirrel), hairs long, banded reddish at base, black distally, white at tip.Tail appears reddish medially when viewed from below, and from above it is dark with white frosting. Tail carried curled up against back when squirrel is at rest; often held high, with the base vertical and tip curled backward when squirrel is moving. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent (and often barely visible). Nipples: 0 + 0 + 2 + 2 = 4. Geographic Variation Four subspecies were recognized by Amtmann (1966): F. a. anerythrus: W Uganda and DR Congo south to Mt Kabobo. Dorsal pelage reddish-brown, ventral pelage ochraceous. F. a. bandarum: Central African Republic. Dorsal pelage buff-tinged, ventral pelage pale grey to orange. F. a. mystax: S Cameroon to Gabon. Dorsal pelage warm brown, ventral pelage orange. F. a. raptorum: SW Nigeria. Dorsal pelage grey-tinged, ventral pelage whitish. Amtmann (1966) placed far western forms (e.g. mandingo; Sierra Leone), previously attributed to this species, in F. pyrropus. Similar Species Funisciurus pyrropus. Larger (HB: 190–230 mm); longer nose; limbs bright to dull reddish; ventral pelage white. Distribution Endemic to Africa. Rainforest BZ (West Central, East Central and South Central Regions) and surrounding forest– savanna mosaics. Recorded from Benin, Nigeria, Cameroon,

Funisciurus anerythrus

Equatorial Guinea, S Central African Republic, Gabon, DR Congo and Congo. Habitat Permanently or seasonally flooded waterside habitats of swamps, gallery forests, seasonally flooded riversides, and islands in rivers. Especially abundant in Raphia palm swamps. In Gabon, it is not found in terra firma forest even where this abuts directly on riversides (Emmons 1975, 1980). In Nigeria, prefers lower strata of secondary forest and tangled vegetation where there is a dense growth of shrubs and creepers; may also be seen climbing in medium-sized trees and oil palms (Happold 1977). Abundance Common in suitable habitats, where it can reach higher densities than does any other species of African tree squirrel. Adaptations Diurnal and scansorial. Forages on the ground and at all levels within the low-canopy of seasonally flooded forests (mean height: 3.8 m [1–13 m], n = 20). Builds round nests (20–24 cm diameter) of leaves lined with fibres. In habitats dominated by Raphia palms, the outer shell of the nest is often built with palm leaflets. The nests are exposed, but partially hidden in dense clumps of vegetation in the branches of trees or on the rachis of palm leaves. Most nests are built in vegetation that overhangs water (Emmons 1975). Foraging and Food Omnivorous. Feeds on fruits and seeds (77% dry mass of stomach contents in Gabon, n = 15), arthropods (20%) and minor amounts of green plant tissues and mushrooms. In Raphia palm swamps, these squirrels feed intensively on the orange pericarp 49

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of Raphia fruits, and they can be seen feeding and climbing around in the crowns of palms. They temporarily cache these palm nuts by wedging them into above-ground crevices. The arthropod portion of the diet is dominated by ants (92% occurrence) and termites (58% occurrence), but many other taxa are also eaten (Emmons 1980). Social and Reproductive Behaviour These squirrels often live in dense populations, and in Gabon were seen with other conspecifics 50% of the time (n = 431 observations). Individuals travelling in pairs (28% of observations) followed each other closely, groomed each other, and rested in contact. Up to six individuals were seen feeding at the same fruit tree. A captive heterosexual pair shared a nest box and showed contact behaviour, but competed aggressively for food (Emmons 1980).The social organization is unknown, but the above observations are consistent with monogamous pair formation. Vocalizations are common (see Emmons 1978 for details). The low intensity alarm call consists of frequency-modulated chucks emitted in groups of 1–6, but usually singly (45% of 166 call bouts). The high intensity call is an astonishing sound unlike that of any other species: a long series of stereotyped groups of 2–4 rhythmic pulses followed by 1–2 long, low frequency whistles (dada dada dadada . . . dadaweeeeeeou). Only the final whistles can be heard at a distance. During a low intensity alarm call, the tail is jerked sharply downward from its resting position against the back, then slowly returns up to its normal posture. As the tail is flicked down, the hindfeet are simultaneously stamped on the substrate, followed by stamping of the forefeet as the tail is raised.

(1–2, n = 6; Dubost 1968, Emmons 1979a) in Gabon. In E DR Congo, pregnancies occur in all months of the year, with the highest pregnancy rate at the end of the dry season and beginning of the wet season (Aug–Oct) (Rahm 1970). Predators, Parasites and Diseases These squirrels have been implicated as hosts for the monkeypox virus (Jezek & Fenner 1988). Conservation IUCN Category: Least Concern. Its widespread distribution and commonness in many habitats would suggest that the appropriate category is Least Concern. Measurements Funisciurus anerythrus HB: 182 (167–198) mm, n = 19 T: 159 (150–170) mm, n = 16 HF: 43.8 (42–46) mm, n = 11 E: 16 (15–16) mm n = ?* WT: 219 (190–240) g, n = 14 GLS: 47.6 (46.1–49.0) mm, n = 3 GWS: 26.2 (25.7–27.1) mm, n = 3 P3–M3: 8.7 (8.5–8.9) mm, n = 4 Gabon (Emmons 1975, L. Emmons unpubl.) *Rosevear 1969 Key References

Emmons 1978, 1979a, 1980; Rahm 1970. Louise H. Emmons

Reproduction and Population Structure Embryo number: 1.2 (1–2, n = 77 litters; Rahm 1970) in E DR Congo, and 1.5

Funisciurus bayonii LUNDA ROPE SQUIRREL Fr. Funisciure de Bocage; Ger. Bocages Baumhörnchen Funisciurus bayonii (Bocage, 1890). Jornal de Sciencias Mathematicas Physicas e Naturaes da Academia de Lisboa, ser. 2, 2: 3. ‘... du Duque de Bragança’. N Angola (exact locality not known).

Taxonomy Originally described in the genus Sciurus, but reallocated on the basis of skull characters to Funisciurus by Hayman (1951). Synonyms: none. Chromosome number: not known. Description Small to medium-sized dark squirrel with indistinct white side-stripe. Dorsal pelage uniformly dull olive, speckled with black. Indistinct side-stripe from shoulder to rump, buff to whitishbuff.Ventral pelage grey tinged with buff. Upper surface of head, foreand hindlimbs similar to dorsal pelage. Pale eye-ring. Tail long (ca. 100% of HB), grizzled, black and ochre without obvious rings, darker than body. Skull: short muzzled, resembling F. congicus in this respect; postorbital processes well developed; nasals broad; cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: not known. Geographic Variation None recorded. Similar Species F. congicus. Paler, also with side-stripe; probably not sympatric. 50

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Funisciurus carruthersi

Distribution Endemic to Africa. Zambezian Woodland BZ. Recorded from NE Angola and SW DR Congo south of Kasai and Congo rivers.

Abundance No information. Known only by a few specimens.

Measurements Funisciurus bayonii HB: 167, 190 mm, n = 2 T: 178, 170 mm, n = 2 HF: 42, 41 mm, n = 2 E: n. d. WT: n. d. GLS: 42 (41–44) mm, n = 5 GWS: 24.7 (24–26) mm, n = 5 P3–M3: 7.7 (7.5–7.8) mm, n = 5 Angola (Hayman 1951)

Remarks Apparently no other information.

Key References

Habitat Uncertain. According to Amtmann (1966), who based his conclusions on a vegetation map of the species range, the habitat is probably moist forest at low to medium elevations; and forest– savanna mosaic, woodland and savannas in south-western parts of the range (principally on Kalahari Sand).

Conservation

IUCN Category: Data Deficient.

Amtmann 1975; Hayman 1951.

Richard W. Thorington, Jr, Lindsay A. Pappas & Chad E. Schennum

Funisciurus carruthersi CARRUTHERS’S ROPE SQUIRREL (CARRUTHERS’S MOUNTAIN SQUIRREL) Fr. Funisciure de montagne; Ger. Carruthers Baumhörnchen (Bergstreifenhörnchen) Funisciurus carruthersi Thomas, 1906. Ann. Mag. Nat. Hist., ser. 7, 18: 140. ‘Rwenzori East, 6500 ft’ (= 1900 m), Uganda.

Taxonomy Funisciurus carruthersi appears to straddle a taxonomic boundary in sharing characteristics of both the Rope Squirrels Funisciurus and Bush Squirrels of the Paraxerus lucifer/byatti complex. In common with Funisciurus spp., adults have flat-crowned upper and lower cheekteeth. By contrast, the colouring, body size and ecological habits of F. carruthersi appear to ally it more closely with Paraxerus. Hopefully, molecular studies will help shed some light on an ambiguous taxonomic situation. Synonyms: birungensis, chrysippus, tanganyikae. Subspecies: three. Chromosome number: not known. Description Medium to large soft-furred olive-green squirrel with grey ventral pelage. Dorsal pelage grizzled olive-green and black, darker on mid-dorsal line; hairs long and soft, dark grey or black at base, yellowish-buff or olive-green subterminal band, and black tip. Long black guard hairs, especially on mid-dorsal line. Flanks paler, with fewer black-tipped hairs, and without side-stripes.Ventral pelage from throat to tail grey or greyish-white; hairs medium-grey at base, white or greyish-white at tip. Scrotum olive-green, as flanks. Head and cheeks similar to dorsal pelage. Conspicuous creamyyellow eye-ring. Ears small, pigmented, with sparse short hairs, and with sharply angled leading edge (so overall shape intermediate between the rather rectangular ear of Funisciurus and more rounded ears of Paraxerus). Forelimbs with four digits; hindlimbs with five digits (Digit 1 reduced); all digits have a strong sharp claw. Colour of pelage of limbs fades to brown before the moult. Tail long (ca. 100% of HB), ochre and black, with inconspicuous bands along length; hairs long, ochre at base, with black subterminal band and ochre at tip. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 1 + 2 = 6. Geographic Variation subspecies:

Amtmann (1966) lists the following

Funisciurus carruthersi

F. c. carruthersi: Rwenzori Mts. F. c. birungensis (including chrysippus): Mountains of SE Uganda, E DR Congo, Rwanda and NW Burundi. F. c. tanganyikae: Mountains around north end of L. Tanganyika. Similar Species Heliosciurus ruwenzori. Similar size; dorsal pelage grey, ventral pelage white; tail with grey and white bands. Heliosciurus rufobrachium. Similar size; pelage of limbs reddish or rufous; tail with conspicuous black and white bands. 51

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Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Recorded from the montane forests of the Rwenzori Mts and Albertine Rift Valley mountains in E DR Congo, Uganda, Rwanda and Burundi (Thorington & Hoffman 2005). Altitude range: 1500– 2800 m. Habitat Montane forests, especially in stands of Prunus (Pygeum) africanum, a common tree species in the region and, at the upper reaches of its altitudinal range, in forests of Hagenia abyssinica. Prefers areas of forest with a broken canopy where trees are smothered in climbers, and undergrowth is densely tangled. Does not adapt well to cultivated areas. In E DR Congo most observations were close to rivers (Rahm & Christiaensen 1963). Abundance Uncertain. Apparently common in all Prunus africanum stands throughout their range. Supposedly less common in higher reaches of montane forest. Abundance is difficult to assess because these squirrels are very shy and inhabit dense vegetation. Adaptations Diurnal and arboreal. Carruthers’s Mountain Squirrels are unusual among squirrels at higher altitudes in becoming active shortly after dawn as well as in the evening. They are less conspicuous than Sun Squirrels Heliosciurus living in the same habitat. The drab-coloured tail, in particular, which is commonly carried over the back, is less conspicuous, even when waved about. This suggests that the principal function of tail flicking is dispersion of scent from the anal glands. There may therefore be a bias in this species towards communication by olfactory rather than by visual signals. Their olive-green colouring provides well-matched camouflage on leafy branches and on the mossy, lichened trunks and branches of trees. Their colouring and noteworthy shyness may reflect greater vulnerability than Sun Squirrels to the abundant buzzards, hawks and hawk-eagles found in montane habitats. Foraging and Food Primarily frugivorous. Foraging occurs at all levels, even, rarely, on the ground. Recorded as feeding on the fruits of Bridelia spp., Alchornea spp., Carapa grandifolia and Strombosia scheffleri and the fruits of an unnamed gourd (Rahm & Christiaensen 1963). Traces of insects have been found in stomachs but very fine

mastication has prevented accurate identification of plant foods (Kingdon 1974). Social and Reproductive Behaviour Usually seen singly, occasionally in pairs.They have been heard to make a rasping, quacking cry, possibly an advertising or contact call, but its significance has not yet been determined. Reported to make large nests of plant material in liana tangles; these are lined with finely shredded bark torn from a local shrub (Kingdon 1974). Reproduction and Population Structure Very little information. There is a single record of a lactating " from SW Uganda in May (Kingdon 1974). An absence of records of juveniles suggests young remain well concealed. Predators, Parasites and Diseases Likely predators are hawkeagles Hieraaetus africanus and H. dubius, eagles, buzzards and, more rarely perhaps, genets and Golden Cats Profelis aurata (Kingdon 1974). Conservation IUCN Category: Least Concern. Previously considered as Vulnerable. Measurements Funisciurus carruthersi HB: 203 (187–236) mm, n = 7 T: 201 (180–231)mm, n = 6 HF: 48 (46–50) mm, n = 7 E: 19 (17–20) mm, n = 7 WT: 222 (200–280) g, n = 4 GLS: 48.7 (46.4–50.2) mm, n = 7 GWS: 28.0 (25.5–29.0) mm, n = 7 P3–M3: 8.55 (8.2–9.0) mm, n = 7 Body measurements: Uganda, DR Congo (BMNH) Weight: Rahm & Christiaensen 1963 Key References

Kingdon 1974; Rahm & Christiaensen 1963. Jonathan Kingdon

Funisciurus congicus CONGO ROPE SQUIRREL (STRIPED TREE SQUIRREL) Fr. Funisciure du Congo (Funisciure de Kuhl); Ger. Kongo Baumhörnchen Funisciurus congicus (Kuhl, 1820). Beiträge zur Zoologie und vergleichenden Anatomie 2: 66. ‘Congo’; no specific locality given, probably Angola (Hill & Carter 1941).

Taxonomy Originally described in the genus Sciurus. Seven subspecies have been described, but Amtmann (1975) considered them invalid because geographic variation in size and colouration is highly correlated with temperature and rainfall (Amtmann 1966). Funisciurus poolii included in this species by Amtmann (1975). Synonyms: damarensis, flavinus, interior, oenone, olivellus, poolii, praetextus. Subspecies: none. Chromosome number: not known. Description Small brown squirrel with long wide side-stripes. Pelage short and soft. Dorsal pelage grizzled yellowish-brown; hairs

black at base, yellowish-brown terminally, some with black tip. Each flank with wide white or cream side-stripe from shoulder to rump, and narrow pronounced brownish-black stripe below the white stripe.Ventral pelage white. Crown of head brown, cheeks and throat white. White stripes above and below eye, forming incomplete eyering; stripes buff or pale ochre on specimens from northern Angola. Ears with white, buff, or pale ochre hairs on outer margins. Outer surface of limbs buff, inner surface white. Tail long (ca. 105% of HB), thin and bushy, black and buff above, pure white below; hairs black at base with alternating bands of buff and black, buff tip above,

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Funisciurus congicus

sunset. On cool mornings they are especially active, probably in order to generate body heat. During hot days they bask in the sun, and may also hold the white underside of the tail over the back to prevent over-heating. These squirrels are very agile and move easily and quickly, jumping from branch to branch. Observations (n = 210) indicate that they prefer habitats on or near the ground: 39% on ground, 42% in subcanopy (2.5 m above ground) (Viljoen 1978). If disturbed on the ground they immediately flee into the trees. Nests are made in the forks of branches and are constructed of twigs, leaves and grass; sometimes nests are in the holes of trees. When running, the tail is held over the back in the form of a question mark (Viljoen 1978). Foraging and Food Omnivorous. Foraging takes place on the ground (39% of the time) and in the subcanopy, mostly in trees up to 2.5 m high. The diet includes seeds, fruits, stems and shoots, especially of Colophospermum mopane,Velvet Commiphora Commiphora mollis, Grewia bicolor and other trees. One individual was observed feeding on Mopane Caterpillars Gonimbrasia belina. Hard foods may be buried for eating later (Viljoen 1978, Smithers 1983). Funisciurus congicus

white below. At rest, tail held along surface of back, with terminal quarter curled over posteriorly. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 0 + 0 + 1 + 1 = 4. Geographic Variation Pelage varies geographically. Individuals from drier regions have paler colouration; those from more humid areas (e.g. C DR Congo) tend to be dark brownish-black. Similar Species F. bayonii. Slightly larger; uniformly dull olive, speckled with black; side-stripe indistinct. Other Funisciurus spp. Darker, often with rufous and rusty colouration. Paraxerus cepapi. Slightly larger (HB!: 145–203 mm); pelage grey, yellowish-brown or brown; without side-stripe. Distribution Endemic to Africa. Rainforest BZ (South Central Region), western parts of Southern Rainforest–Savanna Mosaic, and Zambezian Woodland BZ. Widely but patchily distributed in DR Congo (south of Congo R.), W Angola and NW Namibia. Habitat Coastal palm groves, mopane woodland and tall forest trees, especially along watercourses and on granite outcrops. Requires denser vegetation than Paraxerus cepapi, with which it is sympatric in parts of the geographic range. Abundance Common. In optimal habitat in Namibia, densities may reach 12–18 animals/ha (Viljoen 1978). Adaptations Diurnal, arboreal and terrestrial. In Namibia, these squirrels emerge from their nests soon after dawn (from 07:05h to 08:15h, depending on whether the day is clear or overcast) and return to their nests in early evening (18:00–18:50h), well before

Social and Reproductive Behaviour Congo Rope Squirrels are social animals, and live in small groups of up to four individuals. Individuals in the group keep in contact by a series of bird-like highpitched chirps, and by sight. Individual recognition is by the use of scent, and bonding within the group is maintained by scent and by mutual grooming. The alarm call, in response to a raptor, is a series of high-pitched whistles, which causes other individuals to ‘freeze’ instantly.When a ground predator is spotted, the squirrels emit birdlike chirps accompanied by flicking of the tail; a group may mob a predator by peering down at it while tail-flicking and chattering. Daily home-range of a group living in narrow riverine vegetation (20 m wide on either side of the river bank) is about 0.5 ha; one group moved 200 m on either side of the river during a day, a homerange of ca. 0.4 ha (Viljoen 1978, 1997). Reproduction and Population Structure Juvenile specimens in museum collections are most common in Apr–Jun and Oct–Jan suggesting two birth seasons/year. Litter-size: 2 (n = 8; Viljoen 1978). Predators, Parasites and Diseases Ectoparasites include two species of ticks (Haemaphysalis sp. and Rhipicephalus sp.) and one species of flea (Libyastus vates). Predators probably include snakes, mammalian carnivores and hawks. Conservation

IUCN Category: Least Concern.

Measurements Funisciurus congicus HB: 155.9 (142–168) mm, n = 50 T: 169.9 (153–190) mm, n = 50 HF: 40.2 (37–42) mm, n = 50 E: 16.7 (16–18) mm, n = 50 WT: 111 (108–113) g, n = 63 GLS: 38.8 (35.6–40.5) mm, n = 14 GWS: 22.0 (19.3–23.8) mm, n = 14 53

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P3–M3: 6.7 (6.1–7.0) mm, n = 13 Body measurements: Namibia (Shortridge 1934) Weight: throughout geographic range (Smithers 1983) Skull measurements: Angola (BMNH)

Funisciurus duchaillui

Key References Amtmann 1975; Smithers 1983; Viljoen 1978, 1997. Richard W. Thorington, Jr, Lindsay A. Pappas & Chad E. Schennum DU

CHAILLU’S ROPE SQUIRREL

Fr. Funisciure de Duchaillu; Ger. Duchaillu Baumhörnchen Funisciurus duchaillui Sanborn, 1953. Mammalia 17: 167. Poingi (01° 39´ S, 11° 46´ E), Gabon.

Taxonomy Described as a valid species from a single specimen; later placed as a synonym of F. isabella (Amtmann 1975, Hoffman et al. 1993, Thorington & Hoffman 2005). Reinstated as a full species, separable from F. isabella, on the basis of size and ecology by Brugière et al. (2005). Synonyms: none. Chromosome number: not known. Description Medium-sized arboreal squirrel with four stripes on back. Dorsal pelage olive-brown.Two side-stripes on either side of middorsal line; black from head to base of tail. Flanks grey to greyish-white; hairs with white tip. Ventral pelage grey; hairs with white tip. Head olive-brown. Limbs, fore- and hindfeet grey.Tail long (ca. 100% of HB), slender, with long hairs.When at rest, the tail is curled over the back and is frequently flicked. Skull: cheekteeth 5/4. Nipples: not known. Geographic Variation None recorded. Similar Species F. lemniscatus. Smaller (HB mean: 164 mm; GLS mean: 43.6 mm); ventral pelage white or buffy; two side-stripes on each flank from base of neck to rump; more widespread distribution. F. isabella. Smaller (HB mean: 158 mm, GLS mean: 40.9 mm); ventral pelage grey, white at tip; two side-stripes on each flank from ears/ head to rump/base of tail; more widespread distribution. Distribution Endemic to Africa. Rainforest BZ (West Central region [Gabon subregion]). Recorded from C Gabon, south of the Ogooué R. and south to the Massif du Chaillu. Appears to be allopatric to F. isabella. Habitat Rainforest, especially where trees of the families Cesalpiniaceae and Burseraceae are common. Abundance Moderately common within its restricted range; comprised 29% of sightings (n = 95; Brugière et al. 2005), and was the commonest of eight species observed (the second most numerous species was F. lemniscatus [22% of observations]). Remarks Usually seen singly or less frequently in pairs. Mostly observed in the lower storeys of the forest at 4–6 m above ground, but has been recorded in all storeys from ground level to >25 m (mean height: 10.2 m, median height: 6.0 m). When alarmed, escaped along branches or into higher levels of the forest, never along the ground (cf. F. lemniscatus) (Brugière et al. 2005). Observed eating fruits of Xylopia aethiopica and Dialium sp.

Funisciurus duchaillui

Conservation IUCN Category: Not Evaluated. Probably should be assessed as Data Deficient or Least Concern. Conserved within Lopé National Park, Gabon. Measurements Funisciurus duchaillui HB: 197 ± 13 (185–212) mm, n = 4 T: 210 ± 20 (190–230) mm, n = 3 HF: 45 ± 5.7 (42–50) mm, n = 4 E: n. d. WT: 200 ± 8.3 (180–220) g, n = 4 GLS: 47.2 ± 0.4 (46.9–48.3) mm, n = 3 GWS: 27.1 ± 1.0 (26.2–28.2) mm, n = 3 P3–M3: 7.8 ± 0.3 (7.5–8.2) mm, n = 3 Mean values ± 1 S.D. Gabon (Brugière et al. 2005, D. Brugière unpubl.) Key References

Brugière et al. 2005; Sanborn 1953. David Brugière

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Funisciurus isabella

Funisciurus isabella LADY BURTON’S ROPE SQUIRREL Fr. Funisciure d’Isabella (Funisciure de Gray); Ger. Lady Burtons Baumhörnchen Funisciurus isabella (Gray, 1862). Proc. Zool. Soc. Lond. 1862: 180, pl. 24. Cameroon Mountain, Cameroon. 7000 ft (2130 m).

Taxonomy Originally described in the genus Sciurus. Named after the wife of Sir Richard Burton, the British Consul in Fernando Poo at the time (Rosevear 1969). Taxonomy is reviewed in Amtmann (1966) and Rosevear (1969).The form duchaillui, placed as a synonym of F. isabella by Thorington & Hoffman (2005), is considered as a full species here (see above). Synonyms: dubosti. Subspecies: none. Chromosome number: not known. Description Small brown squirrel with four black side-stripes. Dorsal pelage brown, slightly grizzled with buff; hairs black with buff tips. Two black side-stripes on each flank: inner side-stripe from between ears to base of tail; outer side-stripe from base of neck/ shoulder to rump; buffy-brown pelage between all black dorsal side-stripes, but paler than on shoulders. Ventral pelage grey; hairs grey at base, white at tip. Tail long (ca. 100% of HB), slender, with long hairs, buff at base, black distally, with frosted buff tip. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Males tend to be slightly larger than "". Nipples: 0 + 0 + 1 + 1 = 4. Geographic Variation None recorded. Funisciurus isabella

Similar Species F. lemniscatus. Inner black side-stripes do not extend onto neck; pelage between inner side-stripes is darker (dull brown) than between outer and inner side-stripes (pale yellow). Distribution Endemic to Africa. Rainforest BZ (West Central Region). Recorded from S Cameroon, Equatorial Guinea (Rio Muni), Gabon, SW Central African Republic and NW Congo. Also Brazzaville, S Congo. Habitat Lady Burton’s Squirrels are found in dense, brushy or viny thickets within the Rainforest BZ. Preferred habitats include dense secondary growth alongside roads, abandoned gardens and plantations, and other habitats with closed vegetation below 5 m (Emmons 1975, 1980). They do not live in tall mature rainforest. In some parts of their range, they share this habitat with F. lemniscatus. Abundance Common where habitat is optimal, but distribution is patchy. Adaptations Diurnal and scansorial. Forage above the ground (90% of sightings), but generally stay below 10 m in dense thickets and vine tangles where often the most numerous species of squirrel (Emmons 1980). Nests found where there were many F. isabella, but no F. lemniscatus, closely resembled nests of F. lemniscatus in structure and placement (see F. lemniscatus). Bates (1905, cited in Rosevear 1969) stated that nests consist of a ball of dry leaves and fibres.

Foraging and Food Omnivorous. In Gabon, diet is fruits and seeds (81% of dry matter of stomach contents, n = 14), green plant tissue (9.2%) and arthropods (6%), with minor amounts of fungi. Arthropods consumed are chiefly ants (79% occurrence), termites (59% occurrence) and lepidopteran larvae (29%) (Emmons 1980). Social and Reproductive Behaviour Usually seen alone or in pairs, occasionally in threes (69% alone, 21% pairs, 10% threes; n = 29 sightings). Two captive "" shared a nest box and groomed each other (Emmons 1980). There is no detailed information on social organization. Lady Burton’s Squirrels call readily and often (see Emmons 1978 for details). The low intensity alarm call is a series of chucks emitted singly (45% of 71 groups) or in groups of two (41%) to four. The unique high intensity alarm call is a series of linked, frequencymodulated pulses that form a long warbling sound. The most frequent number of warbles in a call is 7 (60% of 95 calls), followed by 5 (30%), but calls may consist of 2–10 warbles. Warbles alternate between shorter (about 160 ms) and longer (about 310 ms) pulses. Each series of warbles may be preceded by 1–14 stereotyped short warbles. Because of the density of their habitat, Lady Burton’s Squirrels are more often heard than seen, and their presence is most easily determined by identifying their calls. Reproduction and Population Structure Litter-size seems restricted to one (n = 11; Dubost 1968, Emmons 1979a).

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Predators, Parasites and Diseases No information. Conservation IUCN Category: Least Concern. Previously considered as Near Threatened. Measurements Funisciurus isabella HB (!!): 158 (150–170) mm, n = 7 HB (""): 152 (143–165) mm, n = 16 T (!!): 159 (150–180) mm, n = 6 T (""): 163 (135–185) mm, n = 14 HF (!!): 35 (34–38) mm, n = 7 HF (""): 36 (34–38) mm, n = 16

E: 14.5 (12–16) mm, n =?* WT (!!): 116 (97–140) g, n = 8 WT (""): 104 (90–116) g, n = 10 GLS: 40.9 (39.6–41.9) mm, n = 3 GWS: 22.4, 22.4 mm, n = 2 P3–M3: 7.1 (6.7–7.7) mm, n = 5 Gabon (Emmons 1975, L. Emmons unpubl.) *Rosevear 1969 Key References

Emmons 1978, 1980. Louise H. Emmons

Funisciurus lemniscatus RIBBONED ROPE SQUIRREL Fr. Funisciure rayé; Ger. Streifiges Baumhörnchen Funisciurus lemniscatus (Le Conte, 1857). Proc. Nat. Acad. Sci. Philadelphia 9:11. Equatorial Guinea (Rio Muni).

Taxonomy Originally described in the genus Sciurus. Taxonomy reviewed in Amtmann (1966) and Rosevear (1969). Synonyms: mayumbicus, sharpei. Subspecies: two. Chromosome number: not known. Description Small brown squirrel, similar to F. isabella, with four black side-stripes on each flank. Dorsal pelage brown, hairs finely banded black and buffy. Two black side-stripes on each flank from base of neck to rump; inner side-stripes separated by dull brown (on mid-dorsal line); outer side-stripe separated from inner side-stripe by pale yellow pelage. Ventral pelage white or buffy. Tail long (ca. 80% of HB), bushy, with banded hairs, grizzled buff and black, with yellowish-buff below. Tail curled up over the back when the squirrel is at rest. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 0 + 0 + 1 + 1 = 4. Geographic Variation Two subspecies were recognized by Amtmann (1966) and Thorington & Hoffmann (2005): F. l. lemniscatus: north of Ogoué R., Gabon. Ventral pelage white. F. l. mayumbicus: south of the Ogoué R., Gabon. Ventral pelage buffy. Similar Species F. isabella. Slightly smaller (HB !!: 158 [150–170] mm); two inner black side-stripes extend to between the ears; buffy-brown pelage between all black side-stripes. The alarm calls of this species and F. lemniscatus are distinctive. Paraxerus alexandri and P. boehmi. Considerably smaller (mean HB 120 mm for P. alexandri, 102 mm for P. boehmi); tawny-orange or brown mid-dorsal stripe from shoulders to rump, bordered on each side by a black side-stripe and a yellow/cream side-stripe. Distribution Endemic to Africa. Rainforest BZ (West Central Region [Gabon sub-region]). Recorded from Cameroon (S of Sanaga R.), Equatorial Guinea (Rio Muni), Gabon, Congo, SW Central African Republic, extreme SW DR Congo, and Angola (Cabinda).

Funisciurus lemniscatus

Habitat Lowland evergreen humid rainforests; rare in secondary or disturbed vegetation (where F. isabella is the common species). Abundance Common in favourable habitats. Adaptations Diurnal and scansorial.The feet are long and narrow, well suited for terrestrial travel. Ribboned Rope Squirrels forage chiefly on the ground (47% of 167 sightings) and in low vegetation below 5 m; they are rarely seen above 5 m (4% of sightings; Emmons 1975, 1980).They build round nests (mean diameter 21 cm) of single, large, dead leaves taken from the ground, and lined with a ball of fine plant fibres (about 11 cm diameter). Nests have 2–3 entrances. Five individuals followed by radio-tracking used 17 nests: 11 exposed in small treelets (mean height 3.8 m, range 2–10 m), three in hollows

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of standing trees or lianas (mean height 1.9 m, range 1.5–2.2 m) and three in hollow logs that lay on the ground. Exposed leaf nests are placed in the top of saplings where branches divide, in the top of stumps, in free-hanging tangles of lianas, or against tree trunks where an epiphyte, liana or branch provides a support. If disturbed at night by noise or vibration while in an exposed nest, will jump out quickly and run to refuge in another nest nearby (Emmons 1975). Tree-cavity dens had small entrances just large enough to admit the squirrel, and were too small to hold normal-sized nests. Radio-collared squirrels left their nests at dawn and returned in the late afternoon or at nightfall (mean return time 18:22h; range 16:16–18:49h, n = 14) and thus had a long mean daily activity period of 11.22 h (Emmons 1975, 1980).

Ribboned Rope Squirrels call readily and often (details in Emmons 1978). The low intensity alarm call is a series of chucks given in groups of 1–8, but most often in triplets (36% of 61 groups). The high intensity call is a similar series of more prolonged and frequencymodulated chucks or chips, which are linked together in groups of 1–7, but usually in pairs (43% of 83 calls) or triplets (47%). The linked calls drop in frequency between successive calls, and are often preceded by single chucks. Low intensity alarm calling is associated with a downward tail-flick display like that described for F. anerythrus. Reproduction and Population Structure Embryo number: 1.8 (1–3, n = 4; Emmons 1979a); 1.7 (1–2, n = 10; Dubost 1968). Predators, Parasites and Diseases No information.

Foraging and Food Omnivorous. Insects can be captured by extending the highly extensible tongue into confined spaces (as into a deep vial when in captivity) (L. Emmons unpubl.). In Gabon, feeds on fruits and seeds (59% of dry mass of stomach contents, n = 15) and arthropods (36%). Arthropods consumed are mainly termites (100% occurrence) and ants (60%). Sometimes large numbers of termites are consumed, contributing up to 100% of a stomach content, indicating opportunistic feeding behaviour (Emmons 1980). Social and Reproductive Behaviour Ribboned Rope Squirrels were usually seen alone (59% of 222 sightings), but sometimes in pairs (21%), threes (11%) and fours (7%). They are commonly observed in pairs or small groups, which join in alarm calling or foraging together, with individuals spaced at least 1 m apart, but more often 5–20 m apart. A captive heterosexual pair did not share nest boxes, the dominant often attacking the subordinate and defending food and space. A ! and ", with overlapping homeranges, that were followed by radio-tracking for eight days (Emmons 1975, 1980) stayed within the same 0.5 ha for two days, but spaced 15–40 m apart. Home-ranges (assessed by radio-tracking) were 0.94 ha (subadult "), 1.0 ha and 1.24 ha (two adult !!) and 1.6 ha (one adult "). During activity, !! moved at a mean rate of 51 m/h (n = 9 days of activity), and "" at 43 m/h (n = 20 days of activity).

Conservation

IUCN Category: Least Concern.

Measurements Funisciurus lemniscatus HB (!!): 164 (150–177) mm, n = 13 HB (""): 160 (153–173) mm, n = 10 T (!!): 132 (115–145) mm, n = 9 T (""): 135 (122–145) mm, n = 9 HF (!!): 38 (35–40) mm, n = 13 HF (""): 38 (33–41) mm, n = 11 E: n. d. WT (!!): 140 (123–158) g, n = 12 WT (""): 141 (132–155) g, n = 7 GLS: 43.6 (42.1–43.9) mm, n = 3 GWS: 23.8 (22.9–24.6) mm, n = 3 P3–M3: 7.3 (6.9–7.7) mm, n = 6 Gabon (Emmons 1975, L. Emmons unpubl.) Key References

Emmons 1975, 1978, 1980. Louise H. Emmons

Funisciurus leucogenys RED-CHEEKED ROPE SQUIRREL Fr. Funisciure à oreilles noires; Ger. Orangenköpfiges Baumhörnchen Funisciurus leucogenys (Waterhouse, 1842). Ann. Mag. Nat. Hist., ser. 1, 10: 202. Bioko I., Equatorial Guinea.

Taxonomy Originally described in the genus Sciurus. The species name, which means ‘white-cheeked’, is a misnomer; Waterhouse apparently intended the more appropriate name erythrogenys (‘redcheeked’) (Rosevear 1969). Synonyms: auriculatus, beatus, boydi, erythrogenys, oliviae. Subspecies: none. Chromosome number: not known. Description Medium-sized squirrel with grey-brown pelage, red cheeks and one side-stripe on each flank broken into spots. Pelage colour varies geographically (see below). Dorsal pelage softtextured, dark grey-brown to brownish-black; yellow on shoulders. Single side-stripe on each flank, whitish-yellow, broken into spots. Ventral pelage white or pale orange-red. Head and cheeks bright

orange-red; black speckling on crown of head. Ears rounded, black on outer surface; dark postauricular patch often present. Thighs and forearms greyish. Hindfeet more strongly built than forefeet. Tail long (ca. 79% of HB), bushy, hairs long; black or brownish-black with white tips above, red below. Tail constantly erect and frequently carried curled over back. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 2 + 2 = 8. Geographic Variation Rosevear (1969) described four variations as subspecies; here considered as colour variants related to altitude and climate. (1) Ventral pelage white and side-stripe indistinctly broken into stripes (Bioko I.; leucogenys). (2) Ventral 57

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Habitat Lower strata in relatively undisturbed rainforest habitats and forest relics in rainforest–savanna mosaic (Sanderson 1940, Happold 1987). Of 19 individuals live-trapped in Dazing-Sangria, Central African Republic, eight were in unlogged mixed-species forest, eight in monodominant Gilbertiodendron dewevrei forest, two on skidder trails (i.e. trail used by a bulldozer to drag a log to the nearest road) and one on a secondary logging road (Ray 1996; J. C. Ray & J. R. Malcolm unpubl.). Abundance

No information.

Remarks Diurnal and arboreal (J. C. Ray & J. R. Malcolm unpubl.). During trapping at three heights in Dzanga-Sangha, Central African Republic, captured on ground and understorey (ca. 2 m), but not at 15 m (n = 9; Malcolm & Ray 2000; J. R. Malcolm unpubl.). Nests in holes entered either from tree hollow or from beneath vegetation or other suitable cover (Sanderson 1940). Vocalizations described as ‘excited chattering with a staccato quality’ (Rosevear 1969). Young born in simple nests lined with grass and leaves in holes among roots of trees. Pregnancies recorded in dry and early wet seasons (Dec, Feb and May) in Dzanga-Sangha. Embryo number: 1 (n = 3; J. C. Ray & J. R. Malcolm unpubl.).

Funisciurus leucogenys

pelage tinged with orange, side-stripe broken into spots, crown of head red, with obvious black patches behind ears, underside of tail brilliant red (Ghana, Togo, Nigeria and W Cameroon; olivae). (3) Ventral pelage tinged with orange, crown blackish-red, black patches behind ears; greyish-white ‘frosted’ mantle on shoulders (Upper Cameroon, Mt Cameroon; auriculatus). (4) Similar to auriculatus, but darker with buffy-grey mantle on shoulders (higher altitudes on Mt Cameroon and highlands; boydi). Similar Species F. anerythrus. Pelage drab; pale side-stripe not broken into spots; ventral pelage greyish; no dark postauricular patch; more common in secondary habitats. F. pyrropus. Pale side-stripe not broken into spots, thighs and forearms bright red; no dark postauricular patch. Distribution Endemic to Africa. Rainforest BZ (West Central Region, and extreme east of Western Region). Recorded from Ghana (east of Volta R.),Togo, Benin, Nigeria, Cameroon, Equatorial Guinea (Rio Muni) and Central African Republic (upper Sanga R.), Bioko I.

Conservation IUCN Category: Data Deficient. Deforestation and degradation of forests by logging and clearing constitute potential threats. Measurements Funisciurus leucogenys HB: 209.3 (181–225) mm, n = 14 T: 165.7 (151–178) mm, n = 12 HF: 51.6 (49–54) mm, n = 14 E: 20.1 (18–21) mm, n = 14 WT: 237.3 (171–298) g, n = 12 GLS: 51.2 (48.0–52.9) mm, n = 11 GWS: 27.8 (25.7–29.0) mm, n = 11 P3–M3: 23.6 (21.7–24.7) mm , n = 11 Dzanga-Sangha, Central African Republic (J. C. Ray & J. R. Malcolm unpubl., USNM) Key Reference Rosevear 1969. Justina C. Ray

Funisciurus pyrropus FIRE-FOOTED ROPE SQUIRREL (RED-LEGGED ROPE SQUIRREL) Fr. Funisciure à pattes rousses; Ger. Rotfüssiges Baumhörnchen Funisciurus pyrropus (F. Cuvier, 1842). In: E. Geoffroy Saint-Hilaire and F. Cuvier, Histoire Naturelle des Mammifères, vii, No. 66. Tab. 4: 240. Gabon.

Taxonomy Originally described in the genus Sciurus. The type locality was described as ‘et elle venoit de l’ile Fernandopô, dans le gulfe de Guinée’. However, the species is not known from Fernando Poo (now Bioko I.) and since the animal was a pet, it probably came from the mainland (see Hoffman et al. 1993). Type locality is likely to be Gabon or Equatorial Africa (Rio Muni). Rosevear (1969)

refers to the species as Funisciurus pyrrhopus (with an ‘h’), which is an unjustified emendation (Hoffmann et al. 1993). Taxonomy is reviewed in Amtmann (1966) and Rosevear (1969). Rosevear lists five subspecies, but does not include any forms east of the Sanaga R. in Cameroon, and Amtmann (1966) recognizes the nine subspecies given below. Some of the named forms (leonis, mandingo) were placed

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in F. anerythrus by Rosevear (1969). The allocation and validity of named forms of Funisciurus squirrels is in need of revision. Synonyms: akka, emini, erythrops, leonis, leucostigma, mandingo, nigrensis, niveatus, pembertoni, rubripes, talboti, victoriae, wintoni. Subspecies: nine. Chromosome number: not known. Description Beautiful, medium-sized, long-nosed squirrel with a dark back, pale side-stripe on each flank, and bright reddish limbs. Dorsal pelage grizzled greyish or blackish; hairs banded black with buff tip. Head below crown, muzzle, forelimbs below shoulders, and hindlimbs below hips brilliant rufous to dull rusty-red; hairs pure rufous or rusty-red. White or pale grey side-stripes. Ventral pelage pure white or off-white. Eyes ringed with buff. Ears pale behind. Tail long (ca. 80% of HB) and bushy; hairs banded, black at base, rufous distally, with white-frosted tip.When squirrel is at rest, tail is curled up over back; when moving, tail is carried with base vertical and tip curled backwards or horizontally straight behind body. Skull: cheekteeth 5 /4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 0 + 0 + 1 + 1 = 4. Geographic Variation Highly variable, as attested by the large number of named forms (see also above). Amtmann (1966) recognizes nine subspecies:

Funisciurus pyrropus

Abundance F. p. akka: E Congo, Uganda. Sides and crown without red; limbs and muzzle dull red; underparts washed orange. F. p. leucostigma: S. Ghana. Red parts of pelage duller and brownishred; sides below side-stripes red; crown not red. F. p. leonis: Liberia and Sierra Leone. Red parts of pelage deep rufous; sides red. F. p. mandingo: Gambia. Dorsal pelage pale, straw-coloured sprinkled with black; limbs and ears orange. F. p. nigrensis: Nigeria from Cross R. to Niger R. Head brownish. F. p. niveatus: Côte d’Ivoire. Red parts orange-red rather than rusty. F. p. pembertoni: N Angola. Greyish, with limbs red. F. p. pyrropus: S Cameroon to Mayumbe forest, Congo. Brightly coloured. F. p. talboti: Mt Cameroon and SE Nigeria. Red on flanks mixed with greenish-brown. Similar Species F. anerythrus. Smaller (HB: 167–198 mm); pelage brown without red or rufous on limbs; whooping birdlike alarm call. F. leucogenys. Limbs greyish similar in colour to back; black postauricular patch. Distribution Endemic to Africa. Widespread but disjunct in Rainforest BZ (Western,West Central and East Central Regions), and Rainforest–Savanna Mosaics. Recorded from NW Angola, Burundi, Cameroon, Congo, Côte d’Ivoire, Equatorial Guinea (Rio Muni), Gabon, Gambia, SW Ghana, W Guinea, Guinea-Bissau, Liberia, SE Nigeria, Rwanda, S Senegal, Sierra Leone, Uganda and DR Congo. Although sympatric with F. anerythrus in the east of its range (Nigeria eastwards), it is allopatric in the west of its range (from Ghana westwards (Grubb et al. 1998). Habitat

Tall evergreen forest and older secondary forests.

Common in suitable habitat.

Adaptations Diurnal and terrestrial. Fire-footed Rope Squirrels have long, narrow feet, suitable for terrestrial living. They forage on the ground and on fallen logs and brush below 1.5 m. Unlike other squirrels, they build nests on the ground and in burrows. Of 17 nest sites used by four individuals (found by radio-tracking), 14 were in underground burrows and three in hollow logs on the ground. Six burrows were of characteristic structure and were probably dug by the Fire-footed Rope Squirrels themselves. These were simple tubes, often constructed within a termite nest, with an entrance at each end and a central nest chamber (illustrated in Emmons 1975). The other burrows were probably excavated by other mammals, such as Cricetomys emini (1 den) and Atherurus africanus (1 den), and were likely to be used opportunistically by these squirrels. Radio-tracked individuals left their nest soon after daylight and returned to it at 15:18–18:52h (n = 17 times of return; Emmons 1980), and hence had a mean daily activity period of 9.86 h. Foraging and Food Omnivorous. The long muzzle is probably associated with hunting insects in small crevices. In Gabon, feeds on fruit and seeds (83% of dry matter of stomach contents, n = 12) and insects (12%) (Emmons 1980).The insect portion of the diet consists chiefly of ants (100% occurrence) and termites (92% occurrence). These are consumed in large numbers – soldiers, workers, eggs and larvae together – suggesting that the squirrels find and raid ant and termite nests. Social and Reproductive Behaviour Fire-footed Rope Squirrels appear to be mainly solitary: 80% of individuals (n = 56 sightings) were of single individuals, and 14% were in groups of two that were widely spaced and engaged in mobbing. In captivity, a heterosexual pair slept in different nest boxes and did not groom each other until they had been together many months. A ‘mating chase’, 59

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when several !! pursued a ", has been observed (Emmons 1980). Home-ranges (assessed by radio-tracking) were 5.2 ha (one adult !), 1.0 ha (a lactating ") and 2.3 ha (a subadult "). Movements were slow, at a mean rate of 35 m/h for two "" and 61 m/h for two !! (Emmons 1975). Vocalizations are frequent (Emmons 1978). The low intensity alarm is a loud single chuck, or more rarely, a double chuck.The high intensity alarm is distinctive and consists of a staccato, machine gunlike series of chucks, lasting about 20–40 seconds (tatatatatatatatatatata . . . ).This call superficially resembles that of Epixerus ebii, and is unlike the calls of any sympatric Funisciurus species. The tail-flick display during low intensity alarm is similar to that described for F. anerythrus. Reproduction and Population Structure Litter-size usually seems to be one (range 1–2), but there are few records (Emmons 1979a).

Conservation

IUCN Category: Least Concern.

Measurements Funisciurus pyrropus HB: 211 (190–230) mm, n = 11 T: 167 (153–180) mm, = 9 HF: 46 (44–49) mm, n = 11 E: 18 (17–18) mm, n = ?* WT: 283 (260–334) g, n = 9 GLS: 52.7 (50.4–56.7) mm, n = 5 GWS: 28.0 (26.5–29.8) mm, n = 5 P3–M3: 8.8 (8.1–9.2), n = 6 Gabon (Emmons 1975, L. Emmons unpubl.) *Rosevear 1969 Key References

Emmons 1975, 1978, 1980. Louise H. Emmons

Predators, Parasites and Diseases No information.

Funisciurus substriatus KINTAMPO ROPE SQUIRREL Fr. Funisciure de Kintampo; Ger. Kintampo-Baumhörnchen (Togo Streifenhörnchen) Funisciurus substriatus de Winton, 1899. Ann. Mag. Nat. Hist., ser. 7, 4: 357. ‘near Kintampo, Gold Coast hinterland, 800 feet (240 m)’ (= Kintampo, Ghana).

Taxonomy This form is usually listed as a valid species (Amtmann 1975; Hoffmann et al. 1993), but there is underlying doubt about species limits in Funisciurus. This species closely resembles Funisciurus anerythrus, a species originally described from W Uganda. It is not clear whether Funisciurus substriatus, distributed from E Ghana to Benin, is reproductively isolated from populations further east and south and which are here described as Funisciurus anerythrus. Synonyms: none. Chromosome number: not known. Description Medium-sized plain-coloured squirrel with single faint pale stripe on each flank, very similar to F. anerythrus. Dorsal pelage greenish-yellow or ochre, heavily speckled with black; especially pale in Burkina. Single side-stripe on each flank, faint, whitish, bordered ventrally by darker stripe. Ventral pelage white, occasionally suffused with pale ochre; hairs dark grey at base. Foreand hindlimbs similar to dorsal pelage. Tail long (ca.100% of HB), similar but darker than back, with noticeable rings of black and buff; dorsal hairs black with short pale tips; lateral hairs with long pale buff tips, ventral hairs ochre. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 0 + 0 + 1 + 1 = 4.

Funisciurus substriatus

Geographic Variation None recorded. Similar Species F. anerythrus. Very similar in form and colour; distinguished mainly by distribution; ventral pelage does not show the ochre to orange colouration of populations of Funisciurus anerythrus to the east and south of Cameroon.

Distribution Endemic to Africa. Guinea Savanna BZ and Northern Rainforest-Savanna Mosaic and some relict forests in Sudan Savanna BZ. Recorded from Burkina, E Ghana, Togo and Benin. Does not appear to extend eastwards into Nigeria (where replaced by F. anerythrus), or westwards to Côte d’Ivoire. Habitat Woodland savanna and remnant forests; rocky habitats and along rivers in Burkina (specimen labels).

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Abundance No information. Remarks Apparently no other information available. Conservation

IUCN Category: Data Deficient.

Measurements Funisciurus substriatus TL: 326.3 (300–348) mm, n = 28 T: 154.7 (130–178) mm, n = 28 HF: 43.1 (40–45) mm, n = 33

E: 17.5 (13–19) mm, n = 33 WT: 142.5 (115–187) g, n = 28 GLS: 44.2 (42.5–45.3) mm, n = 10 GWS: 24.2 (23.4–24.6) mm, n = 8 P3–M3: 7.2 (6.9–7.5) mm, n = 10 Burkina, E Ghana, Togo, Benin (USNM) Key References

Amtmann 1975; Hoffman et al. 1993.

Richard W. Thorington, Jr & Chad E. Schennum

GENUS Heliosciurus Sun Squirrels Heliosciurus Trouessart, 1880. Le Naturaliste, 2nd year, 1: 292. Type species: Sciurus gambianus Ogilby, 1835.

Heliosciurus rufobrachium.

Heliosciurus comprises six species of small to large tree squirrels widely distributed in sub-Saharan Africa except for arid or treeless regions. There are representatives in rainforest and montane forest (three spp.) and in savanna habitats (three spp.). Two species are widespread while four have rather small ranges. The vernacular name ‘Sun Squirrel’ is a literal translation of the generic name, and refers to the habit of these squirrels of living at higher levels in the forest where they are more likely to encounter sunny habitats. The genus is characterized by moderate size (HB: 170–270 mm, larger than Myosciurus and Funisciurus, but smaller than Protoxerus and Epixerus), slender build, and long tail, usually longer in length than HB. The pelage of the back and flanks is speckled and there is no lateral side-stripe (cf. Funisciurus). The head is relatively large, with small ears held close to the skull. The limbs are relatively long. The colouration of Heliosciurus squirrels is typically greyish or brownish, but one species has rufous or reddish limbs and shoulders (H. rufobrachium) and another has reddish-brown or rufous-brown body pelage (H. mutabilis). The tail is covered with long hairs, each alternately banded with dark and pale bands; the tail is extended

Figure 12. Skull and mandible of Heliosciurus gambianus (BMNH 62.346).

backwards in line with the body when the squirrel is running along branches, and hangs downwards when the squirrel is resting (cf. Funisciurus). Females have three pairs of nipples (not four as in Allosciurus, Protoxerus and Epixerus). Baculum absent. The length of the skull is larger than that of Myosciurus, overlaps extensively with that of Allosciurus, Paraxerus and Funisciurus, but is smaller than in Protoxerus and Epixerus. The skull is characterized by orange ungrooved incisors (except H. rufobrachium – slightly grooved), fossa for the origin of anterior deep masseter muscle extending onto the rostrum, supraorbital notch which is closed on margin of orbit forming a foramen piercing the frontal bone, large sphenopalatine foramina, anterodorsal process of premaxilla, which 61

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Table 12. Species in the genus Heliosciurus. Arranged according to amount of red colouration on the pelage. Species

Red colouration on pelage

Colour of ventral pelage

HB mean (range) (mm)

HF mean (range) (mm)

GLS mean (range) (mm)

Number of cheekteeth (upper/ lower)

H. gambianus

None

White, cream or buff; sparse

196 (180–230)

44 (40–50)

37 (45–51)

4

H. undulatus

None

Whitish-grey to ochre

233

56

32

4

H. punctatus

None

Grey

ca. 390 (355–524)

46 (37–50)

47 (37–50)

4

H. ruwenzorii

None

Creamy-buff, with white mid-ventral stripe

240–260

54

51–52

5

H. mutabilis

Rufous cinnamon (new pelage)

Creamy-buff; sparse

236 (217–269)

53 (46–60)

54 (52–56)

4

H. rufobrachium

Shoulders and limbs rufous to reddish (very varied)

Brown, reddish, rufous, grizzled as dorsal pelage

ca. 230 (205–249)

47–49 (40–56)

53 (50–54)

4

rises to about level with anterolateral angle of nasal, prominent masseteric tubercle (unlike all other African tree squirrels), and the posterior end of bony palate is approximately in line with the posterior end of M3 (Figure 12). There are four cheekteeth in each ramus (4/4), although Heliosciurus ruwenzorii (5/4) has a very small additional tooth (P3). Dental formula: I 1/1, C 0/0, P 1/1, M 3/3 = 20 (except H. ruwenzorii – see above; cf. Paraxerus). Two species are widespread (H. gambianus, H. rufobrachium) and four species have rather constricted ranges. All are arboreal. Heliosciurus ruwenzorii lives in montane forest, H. rufobrachium and H. punctatus in lowland rainforest, H. undulatus and H. mutabilis in forests and woodlands of East Africa, and H. gambianus lives primarily in savannas of West Africa. The species are mostly allopatric, but H. punctatus is entirely sympatric with H. rufobrachium, and the range of H. gambianus is parapatric with that of H. rufobrachium at least in West Africa, though the two species are ecologically segregated. These squirrels are mostly seen singly or in pairs. Sun Squirrels produce a large variety of vocal noises, which are mostly species-specific.

/4

/4

/4 /4

/4

/4

Notes

Widespread savannas N and S of Rainforest BZ Lowland and montane forests SE Kenya, NE Tanzania, Mafia and Zanzibar Is. Rainforest and forest relicts, Liberia to Ghana Montane forests E DR Congo, Rwanda, Uganda, NW Burundi Savannas of eastern Africa; eastern part of Zambesian Woodland BZ Widespread throughout Rainforest BZ and forest relicts

Heliosciurus forms a monophyletic clade with Allosciurus, Protoxerus and Epixerus (Moore 1959). There has been some disagreement concerning the allocation of subspecies to species. The taxa brauni, coenosus, emissus and lualabae, including the populations in the rainforest south of the Congo R., were allocated to H. gambianus by Amtmann (1975) but to H. rufobrachium by Thorington & Hoffmann (2005). Heliosciurus ruwenzorii was formerly placed in subgenus Aethosciurus (Ellerman 1940), whose type species (poensis) is now included in Paraxerus.The genus is placed in the subfamily Xerinae and tribe Protoxerini (which also includes all the other African sciurids except for Atlantoxerus and Xerus – the Ground Squirrels – which are placed in the tribe Xerini) (Thorington & Hoffmann 2005). Species in the genus are distinguished by presence/absence of reddish colouration of the pelage, colour of ventral pelage, bushiness of the tail, body and skull size, and habitat (Table 12). Peter Grubb

Heliosciurus gambianus GAMBIAN SUN SQUIRREL Fr: Heliosciure de Gambie; Ger. Gambisches Sonnenhörnchen Heliosciurus gambianus (Ogilby, 1835). Proc. Zool. Soc. Lond. 1835: 103. ‘brought from Gambia’, possibly near Fort St Mary, Gambia.

Taxonomy Originally described in the genus Sciurus. Geographic variation in pelage colour over an extensive range has led to the description of 20 forms of H. gambianus (Hoffman et al. 1993), many of which are synonyms. The rainforest representative, H. punctatus, originally regarded as a subspecies (e.g. Rosevear 1969), is now considered to be a separate species (Roth & Thorington 1982). Synonyms: abassensis, albina, annularis, annulatus, bongensis, canaster, dysoni, elegans, hoogstraali, kaffensis, lateralis, limbatus, loandicus, madogae, multicolor, omensis, rhodesiae, senescens, simplex. Subspecies: none. Chromosome number: not known.

Description Medium-sized arboreal greyish-brown squirrel with long banded tail, and mostly without any obvious bright colours or markings; considerable geographic variation in colour (see below). Pelage short, slightly coarse. [Description of form gambianus] Dorsal pelage and flanks grizzled pale brown or buff, flecked with black; dorsal hairs with alternating black and buff bands, usually with black tip. Ventral pelage sparse (so skin visible in parts), hairs longer than on dorsal pelage, pure white, cream or buff. Forehead and muzzle similar to dorsal pelage. Chin, throat, chest, as ventral pelage. Eyes large, dark. Ears small and rounded, covered with short hairs,

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Heliosciurus gambianus

Heliosciuirus gambianus.

situated close to head. Fore- and hindlimbs short, grizzled buff without any rufous colour (cf. H. rufobrachium). Forefeet with four digits (D1 vestigial), each with long claw; hindfeet with five digits, each with long claw. Tail long (ca. 110% of HB), buff with up to 15 black circular bands, some clearly defined, others obscure; hairs buff with 3–4 alternating black bands. Skull: cheekteeth 4/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle prominent. Nipples: 0 + 1 + 1 + 1 = 6. Geographic Variation Considerable geographic variation in pelage colour, although these differences in colour are not recognized as subspecific. For example: H. g. gambianus: West Africa. See Description above. H. g. dysoni: Ethiopia, S Sudan. Dark brown with bright rufous around genitalia, around base of tail and on undersurface of tail. H.g. elegans: Uganda. Medium brown; hairs with wide whitish-cream bands provide a flecked pattern on pelage (rather than a grizzled pattern). H. g. rhodesiae: Zambia. Greyish-cream; hairs longer and softer than in other forms. A larger and heavier form than those in East and West Africa. Similar Species H. rufobrachium. Larger (HB: 212–249 mm); limbs red; rainforest habitats; mostly allopatric. H. mutabilis. Larger (HB: 217–269 mm); limbs brownish; savanna habitats; allopatric. H. punctatus. Similar in size (HB: ca. 180–190 mm); pelage dark; rainforest and mosaic habitats; restricted to Rainforest BZ (Western Region). Distribution Endemic to Africa. Guinea and Sahel Savanna BZs, Northern Rainforest–Savanna Mosaic, Southern Rainforest–Savanna Mosaic, and Zambezian Woodland BZ. Recorded from Senegal and Gambia eastwards across West and central Africa to S Sudan, Eritrea and Ethiopia; and from Angola, S DR Congo, N Zambia (west of the Muchinga escarpment) and extreme SW Tanzania (Ufipa Mts). Outlier populations (which appear to be isolated from the main geographic range) on Jebel Marra (W–C Sudan) and C Tanzania (Tabora).

Heliosciurus gambianus

Habitat Wooded savanna, especially where savanna trees are denser and taller than average because of better soil and access to water (Rosevear 1969). Preferred habitats include Guinea savanna of West Africa (Rosevear 1969), savanna woodlands and secondary forest in Uganda (Delany 1975), and Brachystegia woodland in Zambia (Ansell 1960). Also recorded from rather dry locations in E Uganda (e.g. Karamoja) and NW Kenya (e.g. Lodwar, west of L. Turkana). Along the margins of the Rainforest BZ, Gambian Sun Squirrels have moved into areas where forests have been replaced by palm plantations and other savanna-like habitats (Happold 1987). Here they may occur sympatrically with H. rufobrachium and Paraxerus spp. (Rosevear 1969). Abundance Widely distributed and not uncommon in suitable habitats. Probably the most frequently seen of the treeliving savanna squirrels. In many localities, it is the only species of arboreal squirrel. Quantitative information on abundance not available. Adaptations Arboreal and diurnal. Gambian Sun Squirrels are very agile, running along branches, and jumping from one tree to another where the canopies are close together. In more open woodlands, they descend to the ground to forage and to travel (usually with a short bounding gait) to adjacent trees. They are active only during the day, especially the early morning and late afternoon, and they rest at night in nests, lined with leaves, in holes of trees. Gambian Sun Squirrels are remarkably tolerant of a wide range of environmental conditions, from well-wooded moist savannas to dry semi-arid savannas. There is no detailed ecological and behavioural information on this widespread and common species. Foraging and Food Mainly vegetarian, but also omnivorous.The diet includes fruits, nuts and insects, and occasionally eggs, geckos, lizards and nestlings are consumed (Ansell 1960, Delany 1975). 63

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Social and Reproductive Behaviour Usually observed as single individuals, suggesting they are solitary. Vocalizations are important for communication and include high-pitched squeaks, various chattering and trilling notes, and a long ‘ker ... ker ... ker’ emitted when alarmed (Delany 1975). Reproduction and Population Structure Embryo number: 5 (n = 1), 1 (n = 1) in Zambia (Ansell 1960). Young (about onethird grown) observed in Zambia in Sep and Oct. Young individuals in Karamoja (Uganda) in Feb (Kingdon 1974). Young born in nests in holes in trees; at birth young are almost naked and the eyes are closed. Predators, Parasites and Diseases No detailed information, but likely to be preyed upon by diurnal birds of prey.

Measurements Heliosciurus gambianus gambianus HB: 196.6 (180–230) mm, n = 10 T: 214.4 (189–240) mm, n = 10 HF: 44.5 (40–50) mm, n = 9 E: 15.3 (14–16) mm, n = 9 WT: 220 g, n = 1 GLS: 47.3 (44.9–51.0) mm, n = 10 GWS: 26.2 (24.4–27.6) mm, n = 10 P4–M3: 8.4 (8.0–8.9) mm, n = 10 West Africa (BMNH) The southern African form, rhodesiae, is slightly larger, e.g. HB: 211.1 (197–230) mm, T: 244.7 (222–268) mm. Key References

Happold 1978; Rosevear 1969. D. C. D. Happold

Conservation IUCN Category: Least Concern. Unlikely to be threatened: a relatively common species with a very large geographic range.

Heliosciurus mutabilis MUTABLE SUN SQUIRREL Fr. Heliosciure variable; Ger. Mutables Sonnenhörnchen Heliosciurus mutabilis (Peters, 1852). Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin 17: 273. ‘Africa orientalis, Boror, 17° Lat. Austr.’ (= Boror, 19 km NW of Quelimane, Mozambique).

Taxonomy Originally described in the genus Sciurus. The form mutabilis, originally described as a valid species, is considered either as a subspecies of H. gambianus (e.g. by Ellerman [1940] and Swynnerton & Hayman [1950]) or of H. rufobrachium (e.g. by Rosevear [1963], Smithers & Lobão Tello [1976], Ansell [1978], Smithers & Wilson [1979], and De Graaff [1981]). Grubb (1982) showed that mutabilis is a valid species clearly separable from H. gambianus and H. rufobrachium, a systematic opinion followed by Ansell & Dowsett (1988), Hoffman et al. (1993) and in this volume. Grubb (1982) lists five subspecies; Hoffman et al. (1993) and Thorington & Hoffman (2005) list these forms as ‘synonyms’ without distinction between synonyms and subspecies (see below). Synonyms: beirae, chirindensis, shirensis, smithersi, vumbae. Subspecies: five (but see below). Chromosome number: not known. Description Large brown or rufous-brown squirrel, often with patches of contrasting colour on different parts of the body, which result in a scruffy unkempt appearance. Pelage long (15–20 mm on mid-back), slightly coarse. Dorsal pelage and flanks brown or rufous-brown, abundantly flecked with cream. Dorsal hairs with five bands: brown at base, two paler bands and a broad buff band in centre, and cream at tip. Ventral pelage sparse; hairs long, cream or buff. No side-stripe. Head, cheeks and limbs similar to dorsal pelage. Fore- and hindlimbs (outer and inner surfaces) grey or brown (not russet or red as in H. rufobrachium). Tail long (ca. 112% of HB), dark blackish-brown with up to ten obscure cream or buff bands. Tip of tail may be rufous or rich cinnamon (especially when pelage is old). Striking colour changes occur when moulting: newly moulted pelage is cinnamon, rufous or chestnut, and brightly coloured; as the pelage ages, it becomes brown or blackish-brown, and dull. In many

individuals moulting occurs in patches, so the dorsal pelage appears as a patchwork of brown, black, cinnamon and rufous, partly bright, partly dull – a pattern quite unlike that seen in any other species of squirrel. Hence pelage shows seasonal changes in colour and pattern, as well as geographical variation. Skull: cheekteeth 4/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle prominent. Nipples: 0 + 1 + 1 + 1 = 6. Geographic Variation Mutable Sun Squirrels show considerable variation in pelage colouration within populations (mainly due to moulting patterns – see above) and between populations. In Malawi, for example, the following variation is visible in a series of specimens: (a) crown of head and upper back black, mid-back and flanks bright cinnamon, tail rufous with paler bands; (b) dorsal pelage cinnamon, flecked with buff; (c) crown of head, dorsal pelage and tail dark rufous or chestnut brown, flanks cinnamon flecked with pale buff, tail without any discernible bands. In other parts of the range (e.g. E Zimbabwe) some individuals are almost black. The pelage of populations living in montane habitats is particularly long and intensely pigmented, and without cream flecking (P. Grubb unpubl.). Grubb (1982) recognized five subspecies but suggested that much of the colour variation is clinal. Because variation within populations may obscure geographic variation, only the distribution of each of these taxa is provided: H. m. shirensis: S and E Zambia, N Malawi, SW Tanzania. The most widespread form. H. m. beirae: S Tanzania and coastal Mozambique. H. m. chirindensis: highland regions of E Zimbabwe and adjacent Mozambique.

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Heliosciurus punctatus

Habitat Very varied: montane forests, Brachystegia woodland savanna, especially where trees are dense, thickets, riparian forests and coastal forests. Sometimes occurs in mopane woodlands (Dowsett 1969). Abundance

Uncertain; may be quite common in suitable habitats.

Adaptations Arboreal and diurnal. Runs quickly and nimbly among denser branches and twigs, and in the canopy of trees. Nests in holes of trees. Foraging and Food Mostly vegetarian. Feeds on wild fruits, berries, nuts, fresh green shoots and flowers; occasionally eats insects and eggs (Smithers 1966, Smithers & Wilson 1979). Social and Reproductive Behaviour Normally observed as solitary individuals or in pairs. When alarmed, makes loud ‘clucking’ sounds and flicks tail, and sometimes lies flat along the length of a branch or hides in a tree hole (Smithers 1966, 1986a).

Heliosciurus mutabilis

H. m. mutabilis: S Malawi and adjacent Mozambique. H. m. vumbae: SE Zimbabwe and adjacent Mozambique.

Reproduction and Population Structure Young born in ‘summer months’ in Zimbabwe. Litter-size: up to four (Smithers 1986a). Predators, Parasites and Diseases No information. Conservation

Similar Species Paraxerus cepapi. Smaller (HB: 145–203 mm); dorsal pelage dull grey or yellowish-brown; no side-stripe; cheekteeth 4/4. Paraxerus palliates. Similar size (HB: 187–221 mm); brightly coloured with rufous or red on ventral ventrum, cheeks and/or tail; no side-stripe; cheekteeth 4/4. Paraxerus flavovittis. Smaller (HB: 165–176 mm); dorsal pelage rustybrown or red-brown; white side-stripe bordered by lower black stripe; cheekteeth 4/4. Distribution Endemic to Africa. Eastern parts of Zambezian Woodland BZ and southern parts of Coastal Forest Mosaic BZ, including some highland regions. Recorded from S Tanzania, E Zambia, Malawi, E Zimbabwe, and S and C Mozambique. May occur in N Mozambique, but no definite records. Recorded from sea level to ca. 2100 m. In Tanzania, replaced by H. undulatus north of Rufigi R.

IUCN Category: Least Concern.

Measurements Heliosciurus mutabilis HB: 236.3 (217–269) mm, n = 10 T: 265.0 (241–302) mm, n = 10 HF: 53.5 (46–60) mm, n = 10 E: 16.7 (13–19) mm, n = 10 WT: 380 g, n = 1 GLS: 54.2 (52.2–56.2) mm, n = 10 GWS: 32.0 (30.4–34.5) mm, n = 10 P4–M3: 10.3 (9.7–11.1) mm, n = 10 Malawi, Zambia, Tanzania (BMNH) Key References

Grubb 1982; De Graaff 1981. D. C. D. Happold

Heliosciurus punctatus PUNCTATE SUN SQUIRREL (SMALL SUN SQUIRREL) Fr. Heliosciure de forêt; Ger. Geflecktes Sonnenhörnchen (Kleine Sonnenhörnchen) Heliosciurus punctatus (Temminck, 1853). Esquisses Zoologiques sur la Côte de Guiné, p. 138. ‘dans toutes les forêts de la Guiné’. Guinea coast, no exact locality given. Ingoldby (1927) suggested type locality as ‘Secondi and Bibiani, Gold Coast’, Ghana.

Taxonomy Originally described in the genus Sciurus. Considered as a subspecies of H. gambianus by Ingoldby (1927), Ellerman (1940), Rosevear (1969) and Amtmann (1975), but as a valid species by Allen (1939), Roth & Thorington (1982), Hoffman et al. (1993) and Grubb et al. (1998). Rosevear (1969) commented that punctatus was ‘obviously merely a form of gambianus darkened through residence

in the moister climate of the forest’. Included in H. gambianus by Kingdon (1997). Synonyms: savannius. Subspecies: two. Chromosome number: not known. Description Medium-sized rather dark squirrel, with long ringed tail. Pelage long and soft. Dorsal pelage and flanks dark brown 65

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grizzled with buff; hairs sepia at base, bands of dark and buff distally, with black tip. Ventral pelage greyish. Insides of thighs dark grey. Tail long (ca. 120% of HB), slender, banded with dark and pale rings; hairs dark with white tip; underside paler than the upperside. Skull: cheekteeth 4/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle prominent; comparatively larger braincase than in H. gambianus. Nipples: 0 + 1 + 1 + 1 = 6. Geographic Variation H. p. punctatus: forest regions from Liberia to Volta R., Ghana. Darker coastal form. H. p. savannius: savanna regions of Côte d’Ivoire. Paler inland form. Similar Species H. gambianus. Similar but with paler ventral pelage; smaller braincase; different distribution. H. rufobrachium. Larger; reddish colouration on limbs. Distribution Endemic to Africa. Rainforest BZ (Western Region) and Northern Rainforest–Savanna Mosaic. Recorded from Sierra Leone, E Liberia, S Côte d’Ivoire and S Ghana (west of the Volta R.). Habitat Primary and secondary rainforest (including open areas), relict forests in Rainforest–Savanna Mosaic, and southern parts of the Guinea Savanna BZ. Abundance No information. Remarks

Apparently no other information available.

Conservation

IUCN Category: Data Deficient.

Measurements Heliosciurus punctatus TL (!!): 383.3 (355–412) mm, n = 32 TL (""): 396.6 (365–524) mm, n = 28 T (!!): 201.8 (170–220) mm, n = 32

Heliosciurus punctatus

T (""): 207.3 (112–230) mm, n = 28 HF: 45.9 (37–50) mm, n = 60 E: 15.9 (14–19) mm, n = 59 WT (!!): 166.2 (109–222) g, n = 30 WT (""): 169.1 (114–256) g, n = 27 GLS: 47.2 (43.5–49.7) mm, n = 14 GWS: 25.2 (22.9–26.6) mm, n = 12 P4–M3: 8.4 (8.0–8.8) mm, n = 13 Ghana, Côte d’Ivoire, Liberia (USNM) Key References

Rosevear 1969; Roth & Thorington 1982.

Richard W. Thorington, Jr & Chad E. Schennum

Heliosciurus rufobrachium RED-LEGGED SUN SQUIRREL Fr. Héliosciure à pieds roux; Ger. Rotbeiniges Sonnenhörnchen Heliosciurus rufobrachium (Waterhouse, 1842). Ann. Mag. Nat. Hist., ser. 1, 10: 202. Fernando Poo (= Bioko I., Equatorial Guinea).

Taxonomy Originally described in the genus Sciurus. Rosevear (1969) and Amtmann (1966) recognized two species from the gambianus-like group of Sun Squirrels: H. gambianus for small, palecoloured squirrels from dry habitats, and H. rufobrachium for large, darker squirrels from wetter habitats. Subsequently Grubb (1982) allocated specimens of this species from S Tanzania, Malawi, Mozambique, Zambia and Zimbabwe to H. mutabilis (see species profile). Hoffman et al. (1993) list 26 synonyms, attesting to the large amount of variation within the species (see below, and Thorington & Hoffman 2005). Synonyms: acticola, arrhenii, aschantiensis, aubryi, benga, brauni, caurinus, coenosus, emissus, hardyi, isabellinus, keniae, leakyi, leonensis, libericus, lualabae, maculatus, medjianus, nyansae, obfuscatus, occidentalis, pasha, rubricatus, rufo-brachiatus, semlikii, waterhousii.

Subspecies: possibly 16. Chromosome number: not known. Description Large dark brown to grey squirrel with red-tinged limbs and a slender faintly banded tail with pale yellow and blackish bands. Dorsal pelage dark brown, grey or rusty-red, grizzled with buff; hairs have 3–5 bands of dark brown and buff. Variation in the colour of bands produces the many local variations in overall pelage colour.Ventral pelage pale brown, whitish-brown, reddish or orange, grizzled as in dorsal pelage. Shoulders, limbs, inner surface of hindlimbs vary from bright rusty-red to grizzled brown or grey. Head smallish, slightly flattened. Ears short. Eyes large, often bordered by a pale eye-ring. Tail long (ca. 195% of HB), slender, faintly banded with paler bands; carried straight out behind or drooping downward

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Heliosciurus rufobrachium

over a branch, not curled up against back. Skull: cheekteeth 4/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle prominent. Nipples: 1 + 0 + 1+ 1 = 6. Geographic Variation There is much geographical variation in colour, especially in the overall colour of the pelage, and in the colour and intensity of colour of the limbs. Of the 32 named forms, Amtmann (1966) synonymized several forms and listed 16 subspecies (four of them now included in H. mutabilis) without detail. H. r. arrhenii: N Kivu, DR Congo. H. r. aubryi: SW Central African Republic, Gabon, DR Congo between Tshuapa and Kasai rivers. H. r. caurinus: Guinea-Bissau. H. r. hardyi: Côte d’Ivoire. H. r. isabellinus: E Nigeria to Togo. H. r. keniae: western slope of Mt Kenya. H. r. leonensis: Sierra Leone. H. r. maculatus: Liberia. H. r. medjianus: DR Congo. H. r. nyansae: Nyando R. valley, Kenya, Tanzania west of L. Victoria. H. r. obfuscatus: SE Nigeria, Mt Cameroon. H. r. pasha: DR Congo. H. r. rufobrachium: Bioko I. H. r. semlikii: Semliki R., DR Congo. Similar Species H. mutabilis. Paler, less rufous on limbs; multi-coloured pelage during moult; allopatric to H. rufobrachium. H. gambianus. Paler without any reddish colouration; mainly savanna habitats. H. ruwenzorii. Broad, mid-ventral white stripe; restricted distribution only near Albertine Rift Valley. Protoxerus stangeri. Much larger, with grizzled grey head, conspicuously bushy tail and sharply delineated almost naked yellow belly. Distribution Endemic to Africa. Widespread in Rainforest BZ (Western, West Central and East Central Regions), Northern and Eastern Rainforest–Savanna Mosaics, and parts of Guinea Savanna BZ. Recorded from Benin, Burundi, Cameroon, Central African Republic, Congo, Côte d’Ivoire, Equatorial Guinea, Gambia, Ghana, Guinea, Guinea-Bissau,W Kenya, Liberia, Nigeria, Rwanda, Senegal, Sierra Leone, N Tanzania, Togo, Uganda, DR Congo, Bioko I. Habitat Habitats with large trees in lowland evergreen moist rainforests, secondary forests, plantations and gardens in Rainforest BZ; forest outliers and some relict forests in savanna. Abundance Common. Adaptations Diurnal and arboreal. Red-legged Sun Squirrels live in the canopy and middle levels of the forest.They are morphologically adapted for arboreal life, with a long back, short limbs and short broad feet. Red-legged Sun Squirrels nest within tree hollows and prefer those with small entrances. Fifteen nest holes in Gabon were at a mean height of 8.8 m (range 1–20 m). Nests within the tree hollows are constructed of sprays of green leaves attached to their

Heliosciurus rufobrachium

twigs (Emmons 1975). In Gabon, Red-legged Sun Squirrels left their nests at dawn, but often returned well before dark (mean hours of activity 9.46 h); mean time of nest entry 15:55h; range 12:21– 18:31h; n = 13). Foraging and Food Omnivorous. In Gabon, feed on fruits and seeds (89% by dry mass of stomach contents), green vegetative parts of plants (6%) and arthropods (5%) (n = 15 stomachs). Although the nutritional return is evidently low, Red-legged Sun Squirrels spend much of their time hunting for arthropods (76% of 38 observations, Emmons 1980) by searching intently along branches and lianas, poking their nose into crevices and cavities, and rummaging around in epiphytes and suspended debris. Red-legged Sun Squirrels have a predatory, mongoose-like appearance, and they sometimes move with a slow, sneaking, weasel-like gait. In captivity they quickly captured and ate birds flying within their cages, killing them with bites to the head; and they readily ate bird’s eggs as well as arthropods (Emmons 1975). Arthropods identified in the diet included ants, lepidopteran larvae, Coleoptera and others. Social and Reproductive Behaviour Red-legged Sun Squirrels in the wild have been observed singly (60% of observations), in pairs (32%) and in threes (5%) (n = 128 sightings). In captivity they were gregarious and contact-loving: two adult !! and a " always crammed themselves tightly together to sleep in the same nest box, and they frequently groomed each other and rested in physical contact, often draped over one another. They had a strict dominance hierarchy with respect to food.Wild pairs of undetermined sex were seen to forage, groom each other, play and rest together. Two radio-collared !! and two "" nested and travelled alone (Emmons 1975, 1980). The social organization is thus unclear, but bonded pairs seem likely. Squirrels of this species are not highly vocal and are heard calling more rarely than are other species of squirrel (Emmons 1978). The 67

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low intensity alarm call is a single- or double-pulsed chuck or bark, given in groups of 1–3, most often singly. The squirrel may call repeatedly for a few minutes. The distinctive high intensity alarm call sounds somewhat like the cooing of a dove. It has two parts: a low amplitude, descending frequency whine immediately followed by a short, rapid trill of low frequency pulses. During a calling bout, usually only one, or at most a few, calls are emitted at long intervals. Alarm chucks are emitted with a visual display similar to that of Paraxerus poensis. As the call is emitted, the tail is jerked upward in a stiff C shape, or with a somewhat circular sweep, and the feet are stamped on the substrate. If the squirrel is sitting with the tail hanging below, the tail may be jerked stiffly in random directions. Reproduction and Population Structure Litter-size: 1–2, with more litters of one than of two (Rahm 1970). Pregnancy rate seems low; none of 11 adult "" collected in all seasons in Gabon was pregnant (L. Emmons & G. Dubost unpubl.).

Measurements Heliosciurus rufobrachium HB (!!): 237.3 (212–249) mm, n = 12 HB (""): 230.5 (205–241) mm, n = 8 T (!!): 248 (210–265) mm, n = 12 T (""): 248 (230–260) mm, n = 8 HF (!!): 47 (45–56) mm, n = 12 HF (""): 49 (40–55) mm, n = 8 E: n. d. WT (!!): 356 (300–420) g, n = 12 WT (""): 351 (290–387) g, n = 7 GLS: 52.8 (50.6–54.5) mm, n = 5 GWS: 30.8 (29.1–31.8) mm, n = 5 P4–M3: 9.9 (9.4–10.4) mm, n = 5 Gabon (Emmons 1975, L. Emmons unpubl.) Key References

Emmons 1978, 1980; Rosevear 1969. Louise H. Emmons

Predators, Parasites and Diseases No information. Conservation

IUCN Category: Least Concern.

Heliosciurus ruwenzorii RWENZORI SUN SQUIRREL Fr. Héliosciure de Rwenzori; Ger. Ruwenzori Sonnenhörnchen Heliosciurus ruwenzorii (Schwann, 1904). Ann. Mag. Nat. Hist., ser. 7, 13: 71. Wimi Valley, Rwenzori, E DR Congo.

Taxonomy Originally described as Sciurus rufobrachiatus ruwenzorii. Formerly included in the genus Aethosciurus, which is here included in Paraxerus following Moore (1959). Synonyms: ituriensis, schoutedeni, vulcanius. Subspecies: four. Chromosome number: not known. Description Medium to large grey squirrel with broad white stripe running down the underside from the throat to the genitals; geographical variation in pelage colour (see below). Pelage thick and dense. Dorsal pelage and flanks medium grey, slightly grizzled.Ventral pelage (on either side of white stripe) creamy-buff to olivaceous. Hairs pure white in ventral stripe. Head and outer surface of limbs grey. Chin, throat and chest white. Tail long (ca. 110% of HB) and slender, markedly banded with alternating grey and white bands. Skull: cheekteeth 5/4 (anterior premolar P3 small); posterior end of bony palate in line with posterior end of M3; masseteric tubercle prominent. Nipples: 0 + 1 + 1 +1 = 6. Geographic Variation H. r. ituriensis: E DR Congo (mountains west of L. Albert near Djalasinda and Djugu). Darker ventrally, showing less contrast with the dorsal pelage than in H. r. ruwenzorii; tail blacker; dorsal surfaces of feet less brown. H. r. ruwenzorii: W Uganda and E DR Congo (Rwenzori Mts, 1980– 2590 m). See Description above. H. r. schoutedeni: E DR Congo (mountains from west of L. Edward to west of L. Kivu, area around Kahuzi-Biega N. P.), NW Rwanda (Parc National des Volcans, as vulcanius), SW Uganda. Brown on feet and muzzle; ventral pelage beige on either side of the white stripe.

Heliosciurus ruwenzorii

H. r. vulcanius: NW of L. Tanganyika (Mt Kandashomwa, 2330 m, Itombwe area), NW Burundi (Kibira N. P.), SW Rwanda (Nyungwe N. P., usually above 2000 m). Dorsal pelage sootybrown, finely speckled with pale buffy; ventral pelage washed with ochraceous on either side of white stripe; rufous on feet.

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Heliosciurus undulatus

Similar Species Heliosciurus spp. Third upper premolar absent; without white ventral stripe. Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Montane forests of E DR Congo, W Rwanda, W and SW Uganda and NW Burundi.

Social and Reproductive Behaviour Usually seen alone or in pairs (Rahm & Christiaensen 1963). Reproduction and Population Structure Little information. One pregnant " with three large young found in Mar (Rahm & Christiaensen 1963).

Predators, Parasites and Diseases No information. Habitat Montane and bamboo forests (1600–2700 m) (H. r. ruwenzorii); montane forests, gallery forests and forest edges and Conservation IUCN Category: Least Concern. lightly wooded areas (H. r. schoutedeni, H. r. vulcanius), transitional and disturbed montane forests (H. r. ituriensis), cultivations (H. r. Measurements vulcanius). Heliosciurus ruwenzorii HB: 217 (192–242) mm, n = 19 Abundance Uncertain. Formerly common in the forests of T: 240 (220–267) mm, n = 19 Rwenzori Mts. HF: 54.9 (51–58.5) mm, n = 18 E: 17.7 (15–20) mm, n = 15 Adaptations Diurnal and arboreal. Lives mainly in lower WT: 286 (249–318) g, n = 8 vegetation rather than in the canopy. One nest was made of grass and GLS: 51.3 (48.8–54.4) mm, n = 24 leaves (Rahm & Christiaensen 1963). When travelling, the tail is held GWS: 29.2 (27.5–30.4) mm, n = 23 horizontally, in line with the body (Kingdon 1974). Vocalizations P3–M3: 9.1 (8.3–10.1) mm, n = 25* Throughout geographic range (AMNH, FMNH, LACM, MCZ, include a loud chattering call (Thomas & Wroughton 1910). USNM) Foraging and Food Vegetarian and occasionally insectivorous. *RMCA Near L. Kivu, at the edges of the forest, the diet includes the fruits of several species of trees – Parinari holstii, Syzygium cordatum, Key References Kingdon 1974; Prigogine 1954; Rahm & Conopharyngia holsteii, Carapa sp. and Urera hypselodendron – as well Christiaensen 1963. as the lichen Usnea (Rahm & Christiaensen 1963). In farmlands and J. Kerbis Peterhans & Richard W. Thorington, Jr plantations, feed on guavas, papaya, bananas and palm nuts. Stomachs contained fragments of leaves, stems and insects. Local people say that these squirrels store food (Rahm & Christiaensen 1963).

Heliosciurus undulatus ZANJ SUN SQUIRREL Fr. Héliosciure de Zanj; Ger. Zanj Sonnenhörnchen Heliosciurus undulatus (True, 1892). Proc. U. S. Nat. Mus. 15: 465. ‘Male. Mount Kilima-Njaro, 6000 feet (1800 m). Female. Kahé, south of Mount Kilima-Njaro’. Tanzania.

Taxonomy Originally described in the genus Sciurus. Considered a subspecies of Heliosciurus rufobrachium (e.g. Kingdon 1974) until separated from this species by Grubb (1982). Does not intergrade with the populations of H. rufobrachium to the south or west.The form keniae, included in this species by Kingdon (1974), is now retained in H. rufobrachium (Thorington & Hoffman 2005). Synonyms: daucinus, dolosus, marwitzi, shindi. Subspecies: none (Grubb 1982). Chromosome number: not known. Description Large tawny-grey squirrel with long ringed tail. Dorsal pelage grizzled tawny-grey; hairs banded black and orange, with white subterminal band. Ventral pelage whitish-grey to ochre. Face, nose and feet similar to dorsal pelage, but suffused with pale grey-ochre to orange-ochre.Tail long (ca. 120% of HB), slender, with 10–14 black bands alternating with pale bands, tail hairs ca. 40 mm. Pelage colour varies geographically (see below). Skull: cheekteeth 4 /4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle prominent. Nipples: not known.

Geographic Variation Individuals from higher altitudes are darker and richer in colour; those from the north of the range are paler, and those from the south are duller and greyer (Grubb 1982). Similar Species H. ruwenzorii. White ventral stripe; Rwenzori and Albertine Rift Valley only. H. rufobrachium. Reddish colouration of ventral pelage, and on limbs; different distribution. H. mutabilis. Darker colour, with seasonal changes in pelage colour; different distribution. Distribution Endemic to Africa. Somalia–Masai Bushland BZ (perhaps also northern parts of Coastal Forest Mosaic BZ). Recorded from SE Kenya and NE Tanzania, including Mafia and Zanzibar Is. Does not overlap with H. rufobrachium or with H. mutabilis. Occurs up to 1800 m on Mt Kilimanjaro. 69

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Habitat No information. Remarks Omnivorous. Fruits, seeds, palm dates, leaves, buds, with insects important seasonally (Kingdon 1997). Nests in hollow trees or branches. Conservation

IUCN Category: Data Deficient.

Measurements Heliosciurus undulatus HB: 233 ± 20.7 mm, n = 18 T: 281 ± 28.1 mm, n = 18 HF: 56.2 ± 3.2 mm, n = 17 E: 17.1 ± 2.1 mm, n = 16 WT: n. d. GLS: 53.9 ± 1.4 mm, n = 16 GWS: 31.6 ± 0.8 mm, n = 18 P4–M3: n. d. Throughout geographic range; mean ± 1 S.D. (Grubb 1982) Key References

Grubb 1982; Kingdon 1974, 1997.

Heliosciurus undulatus

Chad E. Schennum & Richard W. Thorington, Jr

GENUS Myosciurus African Pygmy Squirrel Myosciurus Thomas, 1909. Ann. Mag. Nat. Hist., ser. 8, 3: 474. Type species: Sciurus minutus Du Chaillu, 1860 (= Sciurus pumilio Le Conte, 1857).

Myosciurus is a monotypic genus, with restricted distribution in rainforests of Cameroon, Equatorial Guinea, Gabon and NW Congo. The most obvious character is the very small size (‘not much bigger than a man’s thumb’ – Rosevear 1969) and the absence of any externally visible pollex (Digit 1 of forefoot).The skull is small, broad and rounded, with rounded orbits, semicircular zygomatic arch, large auditory bullae, and narrow rostrum (Figure 13). Masseteric tubercle absent. Dental formula is I 1/1, C 0/0, P 1/1, M 3/3 = 20.

Myosciurus pumilio.

Figure 13 Skull and mandible of Myosciurus pumilio (BMNH 5.5.23.25).

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Myosciurus pumilio

Pygmy squirrels similar in size to the African Myosciurus are found in South America and the East Indies. The remarkable similarity between these forms is thought to be the result of convergent evolution, and does not to imply any close phylogenetic relationship

between them (Rosevear 1969). There is a single species, Myosciurus pumilio. Louise H. Emmons

Myosciurus pumilio AFRICAN PYGMY SQUIRREL Fr. Écureuil pygmée; Ger. Afrikanischer Zwerghörnchen Myosciurus pumilio (Le Conte, 1857). Proc. Acad. Nat. Sci. Philadelphia 9: 11. ‘headwaters of the Ovenga River’, Gabon.

Taxonomy Originally described in the genus Sciurus (Rosevear 1969). Synonyms: minutulus, minutus. Subspecies: none. Chromosome number: not known. See also Gharaibeh & Jones (1996). Description Very small rufous-brown squirrel, about the size of a mouse. Pelage soft, of moderate length. Dorsal pelage rufousbrown, grizzled with buff; hairs dark grey or black at base, rufousred or yellowish-red terminally. Ventral pelage pale brown. Head similar to dorsal pelage, with prominent pale buff eye-ring. Ears brown, with inner and outer margins bright pale buff. Fore- and hindfeet elongated and narrow; forefeet with no external evidence of a Digit 1, although the vestigial, much reduced bones persist within the wrist (Emmons 1979b). Tail moderate (ca. 86% of HB), slender, hairs rufous at base tipped with black above. Tail is held straight out behind the body, never over the back. Skull: incisors slightly proodont; cheekteeth 4/4; posterior end of bony palate slightly posterior to M3; masseteric tubercle absent. Nipples: 0 + 0 + 1 + 1 = 4. Geographic Variation None recorded. Similar Species No other squirrel in Africa is as small as this species. The next largest squirrel is Paraxerus alexandri (HB: 102.5 [91–114] mm, T: 110.3 [93–126] mm).

Myosciurus pumilio

Distribution Endemic to Africa. Rainforest BZ (West Central Region, parts of Eastern Nigerian and Gabon sub-regions). Recorded from S Cameroon, NW Congo, Equatorial Guinea (Rio Muni and Bioko I.) and Gabon. There is no evidence that the species occurs in eastern Nigeria (see Happold 1987). Habitat Restricted to evergreen moist rainforests and secondary forests. Uses all levels of the vegetation from near the ground to the canopy, but mostly uses 0–5 m (Emmons 1980). Abundance Mostly rare, with a patchy distribution and small geographic range. May be fairly common in some localities. Adaptations Diurnal and arboreal. The tiny body size is probably an adaptation to a specialized life-style of feeding on both the top and bottom surfaces of large tree trunks and branches. The elongated toes and the loss of Digit 1 on both fore- and hindfeet appear to be adaptations associated with a lizard-like locomotion, with the long limbs splayed sideways and the body flattened against the substrate, held by the hooked claws (Emmons 1979b). The nest is undescribed, but individuals have been seen entering tree holes. Foraging and Food Omnivorous.African Pygmy Squirrels have a highly specialized foraging behaviour. They forage almost incessantly, moving rapidly over the surfaces of tree trunks and branches, pulling off small chips of bark, and holding them in the forepaws while something is scraped with the teeth from the bark surface before the chip is dropped. Three stomach contents included bark scrapings (30%), fruit (33%) and ants and termites (37%). The nutritive material sought from bark is unidentified, and it may be a bacterial or fungal film (Emmons 1979b, 1980). The Neotropical Pygmy Squirrel Sciurillus pusillus and Bornean Pygmy Squirrels Exilisciurus exilis and E. whiteheadi have virtually identical behaviour; they feed on a yellow material, probably fungus or bacteria growing on exudate, under the bark of particular species of living trees, especially certain legumes (Emmons & Feer 1997, L. Emmons unpubl.). In contrast, African Pygmy Squirrels forage on the surface of bark pulled from both dead and living trees. These squirrels, unlike many other squirrels, are not known to cache food. Social and Reproductive Behaviour Pygmy Squirrels are apparently solitary: they are usually sighted alone (87%; n = 45 observations) and only rarely are two active on the same tree (13%). The simple alarm call is a low amplitude pipping sound consisting of widely spaced short (18 ms, n = 14) pulses, which are emitted repeatedly (mean interval between pulses 279 ms, n =12) (Emmons 71

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1978). The squirrel usually does not stop foraging when it calls. The tail is carried stiffly straight behind the body when the squirrel is calling and its base is twitched from side to side, while the tip stays more or less centred. Because these squirrels spend most of their time splayed against trunks (and only the dorsal surface is normally seen), the side-to-side tail movement is an effective display. Reproduction and Population Structure 2 (n = 1; Emmons 1979a).

Embryo number:

Predators, Parasites and Diseases No information. Conservation IUCN Category: Least Concern. Previously considered as ‘Vulnerable’.

Measurements Myosciurus pumilio HB: 66 (61–74) mm, n = 6 T: 56 (45–60) mm, n = 6 HF: 18.7 (18–20) mm, n = 6 E: 8 (7–8) mm, n = ?* WT: 16.5 (12.4–20) g, n = 6 GLS: 21.3 (20.3–22.1) mm, n = 6 GWS: 13.5 (12.9–14.1) mm, n = 6 P4–M3: 2.8 (2.7–2.9) mm, n = 4 Gabon Body measurements and weight: Emmons 1975, L. Emmons unpubl. Skull measurements: BMNH *Rosevear 1969 Key References

Emmons 1979b, 1980. Louise H. Emmons

GENUS Paraxerus Bush Squirrels Paraxerus Forsyth Major, 1893. Proc. Zool. Soc. Lond. 1893: 189. Type species: Sciurus cepapi A. Smith, 1836.

Paraxerus alexandri.

A genus of very small to medium-sized tree squirrels with 11 species widely distributed over much of Africa south of the Sahara, except for arid areas and savanna habitats north of the Rainforest BZ. All species are arboreal; most species live in rainforest or montane forests, but two live in wooded savanna habitats. The genus is best represented in eastern Africa, where nine species are recorded. Most species are geographically and/or ecologically separated. The genus is characterized by very small to medium size (HB: 91– 114 mm for the smallest to 145–203 mm for the largest species), and a long well-haired tail mostly longer than the head and body. Pelage markings and colouration show considerable variation. The muzzle is slightly elongated, and the ears are mostly relatively longer than in Funisciurus and Heliosciurus (as expressed by percentage of GLS). Depending on the species, there is no side-stripe, one sidestripe or two side-stripes, and the side-stripes are pale or dark. Females have three pairs of nipples (for those species in which the number has been recorded). The skull is similar to that of Funisciurus

Figure 14. Skull and mandible of Paraxerus cepapi (HC 2172).

but without the specialized condition of the cheekteeth. There are five subhypsodont upper cheekteeth and four lower cheekteeth, all with prominent cusps; the posterior end of the bony palate is in line with the posterior end of M3; supraorbital foramen absent, and the masseteric tubercle is small and insignificant (Figure 14). Dental formula: I 1/1, C 0/0, P 2/1, M 3/3 = 22 (cf. Heliosciurus).

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Ecological characteristics of species in the genus also show considerable variation. Three species (P. alexandri, P. boehmi, P. peoensis) live in rainforest, four species (P. cooperi, P. lucifer, P. vexillarius, P. vincenti) live in montane forests, two species (P. flavovittis, P. palliatus) live in the coastal forests and neighbouring savannas of eastern Africa, and two species (P. cepapi, P. ochraceus) live in savanna woodlands. Likewise, diet and reproductive characteristics also

exhibit considerable variation. The ecology of most of these species is poorly known. Paraxerus, Funisciurus and Myosciurus form a monophyletic clade. Paraxerus is related to Funisciurus, and the species of Paraxerus are more variable in ecology, size and in the number of transbular septae than Funisciurus species. From external appearance and habitat, Kingdon (1974) thought F. carruthersi was more like species of Paraxerus (P. 2.

1.

3.

4.

5. 7. Courtship behaviour of Paraxerus palliatus. 1. one tail flagging approaches other 2. then grooms 3. alert 4. rests 5. resume grooming hindquarters and genitalia 6. leave off and then self groom side by side 7. resume grooming 8. sudden excitement; " curls tail 9. ! mounts

6. 9.

8.

Table 13. Species in the genus Paraxerus. Arranged in order of increasing number of side-stripes on each flank. (n. d. = no data). Species

Side-stripes on each flank

Red colouration on pelage

HB mean (mm)

HF mean (mm)

GLS mean (mm)

P. poensis

None

None

153

33

38

P. cepapi

None

None

175

44

42

P. cooperi

None

203

42

46

P. lucifer

None

222

52

55

P. palliatus

None

187.7a

43.9a

45.3a

P. vexillarius

None

ca. 230

ca. 50

53

P. vincenti

None

212

46

50

Namuli Mt, Mozambique only

P. ochraceus

None (or one, pale) One, black, bordered ventrally by creamyyellow One, white, bordered ventrally by black

Golden-rufous on thighs Head and dorsal pelage bright rufous Cheeks, limbs and ventral pelage bright rufous Limbs, feet and tip of tail rufous-orange Rich rufous on limbs, feet, ventral pelage and around eye None

155

40

41

None

102

26

31

None

172

39

n. d.

None

120

30

35

Savannas; SE Ethiopia to Tanzania Rainforest of NE DR Congo and W Uganda; edges of ears white; mid-dorsal colour tawny-brown SE Kenya to N Mozambique; savannas and forests Rainforest BZ (East Central Region) and relicts to N Zambia; edges of ears not white; mid-dorsal colour tawny-brown

P. alexandri P. flavovittis

P. boehmi

a

Two, black, separated by yellow

Notes Rainforests; Sierra Leone to Zaire; mostly greenish-olive Widespread Zambezian Woodland BZ; savanna and thickets Montane forest in Cameroon only Montane forests in Malawi and Tanzania; black patch on dorsum Coastal regions S Somalia to South Africa Montane habitats NE and C Tanzania

P. palliatus bridgemani; P. p. ornatus is ca. 20% larger in all measurements.

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cooperi, P. vexillarius) than other species of Funisciurus, and considered Paraxerus to be a subgenus of Funisciurus. Paraxerus has in the past been confused with Heliosciurus. Ellerman (1940), for example, included Heliosciurus ruwenzorii and species of Paraxerus (P. poensis, P. lucifer, P. vexillarius) in a subgenus (Aethosciurus) of Heliosciurus. The skulls of the two genera (Paraxerus and Heliosciurus) are superficially similar but are distinguished by absence of the supraorbital foramen and small insignificant masseteric knob in Paraxerus (cf. supraorbital foramen present and large masseteric knob in Heliosciurus) and the form of the naso-premaxillary suture. Paraxerus can also be differentiated from Heliosciurus (Ellerman 1940) by the form of the zygomatic plate, which is shorter with the ridge stopping abruptly over the infraorbital foramen, and not approaching the superior border of the rostrum in Paraxerus (cf. zygomatic plate more strongly ridged, ridge

extending forward in Heliosciurus); and by the cheekteeth, which are usually more cuspidate in Paraxerus, especially in the lower tooth row, though P. poensis is conservative and resembles Heliosciurus. Relationships of the genus have recently been re-assessed. Moore (1959) placed the genus in the tribe Funambulini (with non-African genera), whereas Thorington & Hoffmann (2005), on the basis of molecular studies by Mercer & Roth (2003) and Steppan et al. (2004), place it in the tribe Protoxerini with all the other African squirrels (except Xerus and Atlantoxerus). Species in the genus are distinguished by size, presence/absence of side-stripes, presence/absence of reddish colouration of pelage, and habitat (Table 13). Peter Grubb

Paraxerus alexandri ALEXANDER’S BUSH SQUIRREL Fr. Écureuil de brousse d’Alexander; Ger. Alexander Buschhörnchen Paraxerus alexandri (Thomas and Wroughton, 1907). Ann. Mag. Nat. Hist., ser. 7, 19: 376. Upper Welle, River Iri, Gudima, DR Congo.

Taxonomy Originally described in the genus Funisciurus. Referred to as Tamiscus alexandri by Rahm (1966, 1970). Synonyms: none. Chromosome number: not known. Description Very small, snub-nosed, greenish-brown squirrel with white edges to ears and five long stripes on back and flanks. Dorsal pelage grizzled yellow, black and grey, giving appearance of greenish-brown at a distance; hairs dark grey at base, subterminal band yellow, usually with black tip. Some long pure black guard hairs. Wide tawny-orange mid-dorsal stripe from shoulders to mid-back bordered on each side by a thin black stripe and a thin creamy-yellow stripe. Ventral pelage similar but paler than dorsal pelage, often with irregular patches and streaks of yellow; hairs dark grey at base, yellow or buff at tip. Head similar in colour to dorsal pelage. Vibrissae long. White eye-ring around eye; not obvious in some individuals. Edges of ears finely covered with short white hairs. Limb extremities proportionally large compared with other small squirrels. Forelimbs greenish-brown; four slim long digits each with thin long sharp claw. Hindlimbs greenish-brown, with five digits, Digit 1 reduced; digits and claws similar to forelimbs. Tail long (ca. 100% of HB), well covered with short hairs, indistinctly marked with irregular brown and ochre bars, tapered towards tip. Seasonal changes in intensity of colouration (due to seasonal moulting or to fading?) have been noted: specimens collected between Nov and Feb were brighter than those in Apr, May and Sep. Skull: cheekteeth 5 /4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 1 + 2 = 6. Geographic Variation None recorded. Similar Species P. boehmi. Slightly larger; tawny-orange mid-dorsal stripe from shoulders to rump, bordered on each side by two black stripes separated by a yellow or cream stripe; ears pigmented without white hairs; comparatively longer nose; relatively smaller foreand hindfeet.

Paraxerus alexandri

Funisciurus lemniscatus. Slightly larger, longer-nosed squirrel of darker overall colour. Dull brown mid-dorsal stripe bordered on each side by two black stripes from base of neck to rump, the outer stripe separated from the inner stripe by a pale yellow stripe; longer, more strongly patterned tail. Distribution Endemic to Africa. Rainforest BZ (East Central Region) and parts of Eastern Rainforest–Savanna Mosaic. Recorded from NE DR Congo and Uganda. Distribution extends (east to west) from the Victoria Nile to the Lualaba R., and (north to south) from the Mbomou R. to the Lukuga R., at altitudes of 500–1500 m.

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Paraxerus boehmi

Habitat Lowland rainforest with a preference for tall relatively mature forest. Apparently common in Ironwood Cynometra alexandri (Caesalpiniaceae), a climax species that forms extensive, nearly monospecific stands in many parts of its geographic range. Sometimes occurs in abandoned plantations within the rainforest in E DR Congo (Rahm 1970).

Social and Reproductive Behaviour Commonly seen alone but not infrequently in pairs. Larger numbers have not been noted. Typically silent, but may be the originator of a sharp, rather birdlike twitter sometimes heard when Alexander’s Bush Squirrels are nearby. The large, mobile and white-coloured ears suggest that ear movements might be significant during social interactions.

Abundance Commonly seen in mature forests. Locally, populations may be dense. Rare or absent in young forests regenerating after felling.

Reproduction and Population Structure In E DR Congo pregnant "" have been recorded in Mar, Apr, Jul and Sep (Rahm 1970). In Uganda, pregnant "" were taken in Sep and Nov and juveniles in Apr, Oct and Nov (Kingdon 1974). Embryo number: usually one, sometimes two (Rahm 1970).Testes of specimens collected in Uganda vary in size; whether fluctuating testis size follows a seasonal trend has yet to be determined.

Adaptations Arboreal and diurnal. Alexander’s Bush Squirrels have a compact body, with relatively large hands and feet, sharp claws and thin relatively long digits enabling them to scuttle, spreadeagled, over the very extensive surfaces of large forest trees. There is some indication that they prefer smoother types of bark. They are commonly seen on the bare surfaces of boles and large branches of very large forest trees such as Ironwood, Mahogany, Khaya and Mututu Klainedoxa, and may be seen emerging from hollow branches. It is uncertain whether they build nests or live in holes in trees. They live at many levels of the forest, but more often on larger branches than other squirrels. The reduced nasal region suggests that they rely less upon scent than most other species of squirrels, and they appear to be able to find scent clues only at very close quarters. The incisors are relatively small, suggesting a poor ability to gnaw; this is consistent with a diet of micro-fauna and flora (see below). Foraging and Food Omnivorous. Commonly seen moving in fits and starts over the surfaces of branches. Food appears to be found at many levels of the forest but apparently not normally in the leafy canopy. Movements are fast, and a foraging squirrel explores continuously, stopping only briefly to consume the food items it encounters. Alexander’s Bush Squirrels mostly forage alone, perhaps because of the dispersed nature of their food supply.The diet includes small ants and other insects (about 50% in a small sample). The exact composition of plant matter (such as that retrieved from stomach contents) has yet to be determined. It is possible that lichens may be a significant part of their diet. Traces of tree resins have been recorded (Kingdon 1974; J. Kingdon unpubl.). The diet is more insectivorous than that of sympatric Funisciurus pyrropus (Rahm 1970).

Predators, Parasites and Diseases The most likely predators are accipiterine hawks but hornbills perhaps represent a hazard, especially for animals sheltering in crevices or nests. Snakes may also be predators occasionally. Ectoparasites have not been noted by collectors. Conservation IUCN Category: Least Concern. Previously considered as Near Threatened. The habitat is declining in area due to logging of forests. Measurements Paraxerus alexandri HB: 102.5 (91–114) mm, n = 10 T: 110.3 (93–126) m, n = 10 HF: 26.2 (23–28) mm, n = 10 E: 13.2 (12–14) mm, n = 9 WT: 46 (40–72) g, n = 9* GLS: 30.6 (29.2–33.1) mm, n = 8 GWS: 17.9 (17.4–18.6) mm, n = 8 P3–M3: 5.2 (4.8–6.1) mm, n = 8 Measurements: Uganda and DR Congo (BMNH) Weight: Rahm 1966 Key References

Kingdon 1974; Rahm 1966, 1970. Jonathan Kingdon

Paraxerus boehmi BOEHM’S BUSH SQUIRREL Fr. Écureuil de brousse de Boehm; Ger. Boehms Buschhörnchen Paraxerus boehmi (Reichenow, 1886). Zool. Anz. 9: 315. ‘Marungu (Inner-Afrika)’ (= Marungu, SE DR Congo).

Paraxerus boehmi.

Taxonomy Originally described in genus Sciurus. One form, provisionally treated here as a subspecies (P. b. vulcanorum, see below) has, with some justification, been regarded as a separate species by some authorities (e.g. Schouteden 1946, as Tamiscus vulcanorum). This interesting situation deserves further study in the field, supplemented by molecular work to determine whether more than one species is involved. Referred to as Tamiscus emini by Rahm (1966) and Rahm & Christiaensen (1963). Synonyms: antoniae, emini, gazella, lunaris, tanganyikae, ugandae, vulcanorum. Subspecies: four. Chromosome number: not known. 75

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Description Small olive-coloured squirrel with five long stripes. Pelage varies geographically (see below). Dorsal pelage grizzled yellow, black and grey giving appearance of olive or greenish-brown at a distance. Wide mid-dorsal stripe, tawny-orange, from shoulders to rump, bordered on each flank by two black side-stripes separated by white or cream (giving the impression of an additional stripe). Ventral pelage paler than dorsal pelage. Head similar in colour to dorsal pelage with three pale longitudinal stripes (sometimes indistinct): one above and below each eye, and one across cheek. Ears small, rounded, deeply pigmented, and without white hairs. Fore- and hindlimbs greenish-brown; forelimb with four digits, each with thick claw; hindlimb with five digits each with sharp thick claw. Tail long (ca. 125% of HB), well covered with short hairs; mottled black and ochre and indistinctly barred; tapers towards tip. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: not known. Geographic Variation Pelage colour and pattern of stripes vary geographically. Lowland and upland forms meet and reportedly overlap in range at about 2000 m. Four subspecies are recognized: P. b. boehmi: lowland forest west of Albertine Rift Valley. Pelage darker and sparse; white stripe conspicuous (see Description above). P. b. emini: lowland forest east of Albertine Rift Valley. Pelage paler and sparse; white stripe conspicuous. P. b. gazellae: lowland forest margins in S Sudan. Pelage paler, somewhat ‘bleached’, but otherwise resembles P. b. boehmi. P. b. vulcanorum: montane forests of the Albertine Rift (Rwenzori Mts and the mountains between L. Edward and L. Tanganyika). Pelage darker olive/russet, dense and long; upper black side-stripe wide; lower black side-stripe narrow, white stripe (in between the black side-stripes) narrow and not conspicuous. This subspecies may be a distinct montane species (J. Kingdon unpubl.). Similar Species P. alexandri. Very small (smaller than P. boehmi); wide tawny-orange mid-dorsal stripe from shoulders to mid-back bordered on each side by thin black stripe and thin creamy-yellow stripe; vivid white ears; relatively larger fore- and hindfeet. Funisciurus lemniscatus. Slightly larger, mainly terrestrial squirrel of darker overall colour. Dull brown mid-dorsal stripe (but redder than in P. boehmi) bordered on either flank by two black stripes from base of neck to rump, the outer stripes separated from inner stripes by pale yellow band; longer, more strongly patterned tail. Distribution Endemic to Africa. Rainforest BZ (East Central Region). Recorded from E DR Congo, Uganda, S Sudan, NW Tanzania and N Zambia. The range of subspecies P. b. vulcanorum includes the montane forests of the Albertine Rift Valley. Records from W Kenya need confirmation. An outlier population is present in Bahr el Ghazal (at two localities west of Malek) in C Sudan (F. Dieterlen unpubl.). Habitat Typically found in the undergrowth and lower storeys of rainforest, notably in thick tangles of lianas and sometimes on the ground. The montane form prefers disturbed areas with dense undergrowth. Although mainly a true rainforest species it has

Paraxerus boehmi

been recorded from wooded savanna in E DR Congo (Rahm & Christiaensen 1963), as well as on the edges of plantations and in tangles along roadsides (Rahm 1966). Abundance Common in suitable habitats. The subspecies P. b. vulcanorum is especially common in montane areas. Adaptations Arboreal and diurnal. Boehm’s Bush Squirrels are well adapted to moving fast through dense tangles of vegetation by virtue of their tapered limbs and small hands and feet.They can travel in any direction over trunks and branches, even hanging upside-down to examine the underside of branches. The inner toe of the hindfoot is partially opposable, increasing the squirrel’s ability to grip plant stems and twigs. The robust muzzle and incisor teeth are well suited to tearing bark, moss and lichen and, it seems, to catching, gnawing and carrying food items. Boehm’s Bush Squirrels make unusually large nests (which resemble the nests of birds) that are dense agglomerations of fine twigs and grasses with a chamber in the middle lined with finer materials, including strands of soft shredded bark. They are built in thick tangles 2–8 m from the ground (Rahm & Christiaensen 1963). As with some other squirrels, the tail is likely to serve as a scent disperser for pheromones emanating from the anal glands. Sustained flicking of the tail is often accompanied by a bird-like chitter. This has been interpreted as an alarm but is just as likely to serve as a combined olfactory/auditory/visual set of signals directed at conspecifics. Foraging and Food Frugivore and insectivore. Animals forage by travelling along branches, tearing moss, lichen and bark with their incisor teeth. However, unlike other species of squirrels, the forefeet seem to be less commonly used to hold food, which is usually seized by the mouth. Of 30 stomachs investigated in Uganda, seven contained insects alone (mainly ants but also caterpillars and beetles), ten contained a large proportion of insects, while eight

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Paraxerus cepapi

had traces of insects; 12 contained vegetable matter (including the fruits of Rubus and epiphytic mushrooms) and ten contained large quantities of tree resin (Kingdon 1974). In E DR Congo, stomach contents contained resins from Albizzia trees, and the remains of fruits, caterpillars, beetles and other insects (Rahm & Christiaensen 1963). Feeding on insects is unusual for squirrels.

Predators, Parasites and Diseases The most likely predators are accipiterine hawks (especially the Long-tailed Hawk), hornbills and snakes. They might also be vulnerable to nocturnal arboreal carnivores such as Genetta spp. and Palm Civet Nandinia binotata, especially while in their nests. They are host to a particular coccidial parasite, Wenyonella parva (Van den Berghe 1938).

Social and Reproductive Behaviour Normally solitary, but also observed in pairs and trios. Boehm’s Bush Squirrels sometimes forage close to another individual for some time. They never aggregate, but the regularity of sightings suggests that they remain fairly evenly dispersed, and perhaps are more densely distributed in suitable habitats. Reproductive !! pursue "" relentlessly, and copulation has been seen even while hanging on a vertical trunk.

Conservation

Reproduction and Population Structure In E DR Congo, where rainfall occurs in all months, Rahm (1970) recorded reproductively active and pregnant "" in all months of the year. The percentage of pregnant "" varied monthly from 10 to 35% (total n = 222 "" examined, range 8–41 ""/month) with the highest percentages occurring towards the end of the dry season and the beginning of the wet season. Rahm (1970) commented that rainfall did not seem to exert a strong influence on the timing of reproduction. In Uganda, Kingdon (1974) provided confirmatory evidence, with records of pregnant "" in Jan, Jun, Nov and Dec (total n = 4), and lactating "" in Jan, May, Jun and Jul (total n = 5). Embryo number: 1 (90%) or 2 (10%) (n = 61 pregnant ""; Rahm 1970). This pattern of reproduction indicates that the population contains young, subadult and adult individuals, and reproductively active and inactive individuals, in all months of the year.

IUCN Category: Least Concern.

Measurements Paraxerus boehmi emini HB: 120 (111–126) mm, n = 10 T: 152 (140–160) mm, n = 9 HF: 30.7 (28–34) mm, n = 9 E: 13 (11–14) mm, n = 10 WT (!!): 69 (48–80) g, n = 9 WT (non-pregnant ""): 79 (72–83) g, n = 6 GLS: 34.9 (33.9–36.6) mm, n = 10 GWS: 19.5 (18.1–21.7) mm, n = 10 P3–M3: 5.8 (5.3–6.4) mm, n = 10 Measurements: Ituri Forest, DR Congo (BMNH) Weight: near L. Kivu, E DR Congo (Rahm & Christiaensen 1963) Mean HB measurements from E DR Congo (HB !!: 135 [110– 145] mm; "" 137 [129–145] mm; Rahm & Christiaensen 1963) are greater than those from Ituri Key References Kingdon 1974; 1997, Rahm 1970; Rahm & Christiaensen 1963. Jonathan Kingdon

Paraxerus cepapi SMITH’S BUSH SQUIRREL Fr. Écureuil de brousse de Smith; Ger. Smiths Buschhörnchen Paraxerus cepapi (A. Smith, 1836). Report on the Expedition and Exploration of Central Africa, p. 43. Marico River, Rustenberg District, W Transvaal, South Africa.

Taxonomy Originally described in the genus Sciurus. Kingdon (1974) proposed that Paraxerus cepapi and Paraxerus palliatus hybridize in the wild and form hybrid populations;Viljoen (1989) disagreed, pointing out distinctive differences in behaviour and habitat use between the two species. Many subspecies have been described: Amtmann (1975) lists ten subspecies although recognizing that not all of them are valid, and Ansell & Dowsett (1988) subsequently synonymized soccatus with yulei. Synonyms: cepate (lapsus for cepapi), bororensis, carpi, cepapoides, chobiensis, kalaharicus, maunensis, phalaena, quotus, sindi, soccatus, yulei. Subspecies: nine. Chromosome number: not known. Description Medium-sized yellowish-brown grizzled squirrel without any bright colours or markings. Dorsal pelage and flanks grey, yellowish-brown or brown; hairs annulated with alternating bands of yellow and black, usually with black tip. Ventral pelage dull white, tending to yellow or buff on the chest. No side-stripe. Head with indistinct upper and lower white eye-stripes; cheeks pale yellowishbrown. Limbs short, similar in colour to flanks; well-developed digits with sharp claws. Tail long (ca. 95% of HB), bushy, with long

hairs annulated with alternating bands of black and yellowish-brown. Pelage colour varies geographically. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 0 + 1 + 1 + 1 = 6. Geographic Variation Amtmann (1975) lists the following subspecies (see also above): P. c. bororensis: Namabieda, Boror, north of the Zambezi R., Mozambique. Darker and more chestnut than P. c. cepapoides. Sides of body and lower part of hind legs greyer. P. c. carpi: Junction of Messenguez and Zambezi rivers, Mozambique. Small size (HB: 158 mm; T: 150 mm). Paler than P. c. cepapi, with thighs and mid-line of underside of tail orange-yellow, feet whitish or whitish-yellow. P. c. cepapi: Transvaal, S Botswana and S Zimbabwe. See Description. P. c. cepapoides: Zimbiti, Beira, Mozambique. More rusty coloured than P. c. cepapi, with the upper parts of body and thighs having a more tawny hue. 77

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P. c. chobiensis: N Botswana and N Namibia. Ventral pelage and toes whiter than in P. c. cepapi. P. c. phalaena: Ovamboland, Namibia, and SW Angola. Dorsal pelage pale grey, with crown, shoulders, hips and legs also grey. Foreand hindfeet pale buffy-white, paler than in P. c. cepapi. P. c. quotus: SE Katanga and N Zambia. Darker colouring with no suffusion of colour on flanks. P. c. sindi: Tete district, Zambezi R., Mozambique and S Malawi. Thighs and underside of mid-line of tail ochre. Ventral pelage white. P. c. yulei: NE Zambia, N Malawi and W Tanzania. A large subspecies (type specimen HB: 205 mm). Dorsal colour pale coarsely grizzled tawny, greyer over the shoulders, sides paler greyishtawny; ventral colour white, more greyish on the belly; fore- and hindfeet greyish-white to whitish-yellow. Similar Species P. ochraceus. Darker, with many white-tipped hairs in dorsal pelage; mostly in E Tanzania and Kenya (further north than P. cepapi). Distribution Endemic to Africa. Zambezian Woodland BZ. Recorded from S Angola, Zambia, SE DR Congo, Malawi, SW Tanzania, S Mozambique, N Namibia, N Botswana, Zimbabwe and South Africa (former Transvaal). Habitat Mixed woodlands and thickets, especially in stony, hilly country, especially in Colophospermum (mopane), Brachystegia (miombo) and Acacia woodlands, and in riverine forests. These squirrels prefer areas where there are trees with suitable holes for nesting. Abundance Widespread and fairly common. In sandveld habitat in South Africa, density was 2.08–2.58 individuals/ha (biomass of 449–557 g/ha) (Viljoen 1986). Adaptations Diurnal and arboreal. Smith’s Bush Squirrels avoid activity during the hottest times of the day. Winter mornings usually begin with basking and grooming in the sun. Both auto- and allogrooming are significant social behaviours (see below). They are very alert and can be quick on the ground and swift in the trees when danger is near. Although mainly arboreal, individuals will descend to the ground to forage for fallen fruits. They are very active, running quickly along branches and jumping from tree to tree. At night, squirrels retire to their holes, where they nest in territorial groups.While holes in trees are favourite nest sites, they will also nest in holes in the ground, in rocky crevices and in roofs of houses. Nests are lined with grass and leaves, and the squirrels frequently clean them out, possibly to reduce the number of parasites in the nest (Smithers 1983). Foraging and Food Predominantly vegetarian; sometimes omnivorous. Bush Squirrels forage in the trees and on the ground, and suitable foods are held in the forefeet while eating. In South Africa (Viljoen 1977b) they feed opportunistically on a wide variety of seeds, berries, flowers, stems, leaves and gum, changing the diet according to the season. Seeds and gums of Acacia, and seeds and flowers of Aloes are favourite foods. Analysis of stomach contents showed that insects (mostly termites) formed about 30% of the diet on an annual basis (range 0–85%), and 79% of stomachs (n = 49) contained some insect material. Over a period of several

Paraxerus cepapi

months, more than 30 species of plants were utilized in the diet (Viljoen 1975). In East Africa, they are reported to feed on fruits of Sclerocarya, Pterocarpus and Kigelia, Aloe and Euphorbia leaves, as well as unspecified bulbs, nuts, seeds, insects and bird’s eggs (Kingdon 1974). In gardens and plantations, they feed on mangoes and many other cultivated fruits (De Graaff 1981). Seeds are cached close to grass tufts and tree trunks, thus facilitating dispersal and germination of savanna trees (Viljoen 1997). Smith’s Bush Squirrels drink water, both in the wild and in captivity, usually from holes in trees (Viljoen 1975). Smith’s Bush Squirrels are a ‘colonizing species’ (Viljoen 1983a), living in vegetation that grows quickly and produces many fruits and seeds (in contrast to tall mature forest); hence a squirrel can find sufficient daily food in a small area (see above). Social and Reproductive Behaviour Social and territorial. Average group size is five animals (rather large for squirrels), usually including one or two adults and several juveniles. Group size in South Africa varied seasonally, from an average of 2/group in Sep to 12/group in Nov when adults were accompanied by young, and in different years (Viljoen 1977a). Pairs and solitary animals are commonly seen.They are territorial, except during the mating season when strange squirrels are tolerated. Territory size in termitaria thickets is 0.3–1.26/ha, although within this area an individual or group may feed within only 150 m2 (Viljoen 1986, 1997). Territory size varies with habitat, being smallest in termitaria thickets and larger in woodlands. Territorial behaviour includes vocalizations, chasing of strangers and scent marking. Scent marking is very common and can include mouth-wiping, urination and anal-dragging (Viljoen 1983b). A dominance hierarchy was observed by Viljoen (1977a), especially with regards to feeding and access to food. In captivity, individuals are very aggressive towards each other; fighting results in many torn ears and sometimes an individual is killed. Dominance– subordinance relationships are important in the social organization of these squirrels, and hence behaviour such as chasing, fighting, mutual

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Paraxerus cooperi

grooming and scent-marking play important roles in maintaining group bonds and territorial boundaries. Vocal communications are well developed (Viljoen 1983b). Most vocalizations begin with a loud call (usually a ‘click’, ‘rattle’ or ‘whistle’) and continue with decreasing intensity and descending pitch. Different types of calls may be joined in succession, depending on the circumstances. Four calls have been described: (a) Whistle call – a high frequency call, with 6–7 notes emitted at about 1 sec intervals; rather similar to a bird call, indicating extreme alarm. (b) Rattle and clicks, a series of different calls indicating alarm and alertness, and also used in territorial defence. These complicated calls results in ‘chir-chir-chir’ alarm rattles and clearer ‘click-clickclick’ calls, spread out over several seconds. (c) Murmurs – a lowpitched nasal sound emitted during courtship and mating by the male. Clicking noises are also emitted by both ! and " during courtship, and by " when communicating with young. (d) Grunting and growling – when disturbed at the nest. Mating behaviour occurs in the morning. Mating chases are initiated by enthusiastic calls of ", a sound similar to the alarm rattle; ! also has a distinct mating call, and clicking and tail flicking are common in both sexes. Male allogrooms " during mating, and they both autogroom afterwards (Viljoen 1977a). Reproduction and Population Structure In Botswana, pregnant "" recorded in every month except May and Sep (n = 256 ", Smithers 1971); the number of pregnant or lactating "" was significantly lower during the cold dry months (4.5%) than during the warm wet months (51%). In South Africa, reproduction is seasonal and most young are born in Oct–Jan (Viljoen 1997). While one litter/year is most common, inter-birth interval in captive animals is 60–63 days (n = 4; Viljoen, 1977a). Gestation: 56–58 days (longer than for most squirrels). Litter-size: 2 (1–3), sample size not recorded. At birth, young are comparatively precocious. Eyes open by Day 7–8, young begin to climb out of the nest by Day 19, take solid food by Day 21 and are fully weaned by Day 29–42.

Sexual maturity by 6–10 months. Both parents groom their young. Subadults are usually evicted from the group when sexually mature. Infanticide has been observed occasionally by !! in a group, after which !! may try to mate with "" (De Villiers 1986). Predators, Parasites and Diseases Predators probably include raptors, snakes and probably some carnivorous mammals. Ectoparasites include a species of chigger, two species of mites, seven species of ticks, four species of fleas and one species of sucking louse (details in De Graaff 1981). Blood parasites include several bacteria (transmitted by ticks), which are responsible for a variety of fevers. Viljoen (1977b) noted that all the squirrels she studied were heavily infected with the parasitic nematode Syphacia paraxeri. Conservation

IUCN Category: Least Concern.

Measurements Paraxerus cepapi HB: 175.5 (145–203) mm, n = 38 T: 169 (116–215) mm, n = 61* HF c.u.: 43.0 (26–49) mm, n = 61 E: 19.0 (16–21) mm, n = 128 WT: 192.3 (76–265) g, n = 52 GLS: 44 (43–45) mm, n = 8 GWS: 26 (25–26) mm, n = 8 P3–M3: 7.4 (6.8–7.9) mm, n = 15 T, HF and WT: Transvaal, South Africa (Rautenbach 1978) HB, E and P4–M3: throughout geographic range (USNM) GLS and GWS: Botswana (Smithers 1971) *"" only Key References De Graaff 1981; Smithers 1971; Viljoen 1975, 1977a, b, 1983a, b, 1997. Lindsay A. Pappas & Richard W. Thorington, Jr

Paraxerus cooperi COOPER’S BUSH SQUIRREL (COOPER’S MOUNTAIN SQUIRREL) Fr. Écureuil de brousse de Cooper; Ger. Cooper Buschhörnchen Paraxerus cooperi Hayman, 1950. Ann. Mag. Nat. Hist., ser. 12, 3: 262. Kumba Division, Rumpi Hills, Cameroon.

Taxonomy Although originally described in the genus Paraxerus, Rosevear (1969) placed this species (together with poensis) in the genus Aethosciurus even though he admitted that the status of the genus was questionable. Eisentraut (1976) placed cooperi in a separate genus, Montisciurus, because it has many palatal ridges between the molars. Synonyms: none. Chromosome number: not known.

and hindfeet rufous. Thighs deep rufous. Tail long (ca. 100% of HB), blackish golden-green above, without bands or rings, golden-yellow stripe (similar in colour to ventral pelage) below. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: not known. Geographic Variation

Description Medium-sized dark squirrel with pale goldenrufous thighs. Pelage very soft and long. Dorsal pelage blackishbrown speckled with creamy-buff, becoming more olive-green and golden on flanks; dorsal hairs dark grey or black at base, subterminal band bright buff or gold, with black tip. Ventral pelage medium grey at base, golden-yellow at tip. Head similar to dorsal pelage. Ears darkly pigmented, mostly naked, with yellow-tipped hairs on outer surface. Lips and cheeks golden. Forearms, upper surface of forefeet

None recorded.

Similar Species P. poensis. Smaller (HB: 148–161 mm), lacks rufous colour on limbs and feet. Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Recorded only from forested montane regions of S Cameroon (Kupe Mts, Oku Mts, Rumpi Hills; Eisentraut 1973). 79

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Abundance No information. In the late 1960s, the species may have been far more abundant than previously thought (M. Eisentraut in Rosevear 1969). Remarks One individual (the holotype) was feeding on the succulent flowers of the guttiferaceous tree Pentadesma butyracea and was noticeably fat. This record is the only direct evidence of squirrels feeding on flowers in W Africa (Rosevear 1969). Conservation IUCN Category: Data Deficient. Previously considered as Vulnerable. Measurements Paraxerus cooperi HB: 203 (192–212) mm, n = 12 T: 179 (161–200) mm, n = 12 HF: 42 (41–45) mm, n = 12 E: 16 (15–17) mm, n = 12 WT: n. d. GLS: 45.9 (44.4–46.6) mm, n = 12 GWS: 26.7 (26.2–27.3) mm, n = 12 P3–M3: 8.8 (8.3–9.0) mm, n = 12 Oku Mts, Cameroon; !! only (Eisentraut 1973)

Paraxerus cooperi

Habitat Lower storeys of forest remnants in the Bamenda highlands at altitudes above 1400 m (Rosevear 1969).

Key Reference Rosevear 1969. Richard W. Thorington, Jr & Chad E. Schennum

Paraxerus flavovittis STRIPED BUSH SQUIRREL Fr. Écureuil de brousse rayé; Ger. Gestreiftes Buschhörnchen Paraxerus flavovittis (Peters, 1852). Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin 17: 274. ‘Africa orientalis. Mossimboa, Quitangonha, a 11° ad 15° Lat. Aust’. Mocimboa, NE Mozambique.

Taxonomy Originally described in the genus Sciuirus. Commonly spelled flavivittis, but this was an unjustified emendation by Peters (1852). Four subspecies usually recognized, but individual and seasonal variations suggest that mossambicus is a synonym of flavovittis (see Hinton 1920). Synonyms: exgeanus, ibeanus, mossambicus. Subspecies: three. Chromosome number: not known. Description Medium-sized arboreal squirrel with a lateral whitish side-stripe. Pelage short and slightly coarse. Dorsal pelage rusty-brown or red-brown; hairs black with one or two wide rustybrown bands, usually black at tip. White to yellowish side-stripe, 9–10 mm wide, bordered below by a darker stripe. Flanks (below dark side-stripe) olive-brown. Ochraceous hairs tint the forelimbs, and ochraceous colouration may extend across the shoulders forming a mantle, and even onto the crown and into the lumbar region. Ventral pelage white or off-white. Head similar to dorsal pelage, though usually not ochraceous. Cheek with two indistinct white bands, one above the eye and one below, from nasal region to base of ear. Fore- and hindfeet whitish. Tail long (ca. 97% of HB), bushy towards tip, with black and white rings at distal end. Pelage colour

varies seasonally or individually, and also during the moult. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: not known. Geographic Variation P. f. exgeanus: SE Tanzania. Side-stripe narrower (5 mm wide) and not as long as in flavovittis. P. f. flavovittis: NE Mozambique (Mossimboa). See Description above. P. f. ibeanus: NE Tanzania and SE Kenya. Side-stripe slightly broader than in exgeanus, but shorter; facial stripes faint. Similar Species Paraxerus spp. (other striped species). Mostly smaller in body size. P. cepapi. No side-stripe. P. ochraceus. No side-stripe. Distribution Endemic to Africa. Coastal Forest Mosaic BZ and nearby regions of Zambezian Woodland BZ. Recorded from SE Kenya, E and SE Tanzania, SE Malawi and N Mozambique.

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Paraxerus lucifer

Foraging and Food Omnivorous. Forages on the ground and in the trees. Feeds on fruits, seeds, buds, leaves, roots and some animal matter. In farmland, eats millet and other grains (Kingdon 1974, 1997). Social and Reproductive Behaviour Little information. Usually seen in pairs or as " with her young (Kingdon 1974). Reproduction and Population Structure Young and juveniles have been recorded in Mar, Apr, Jun and Sep (Kingdon 1974).Young are born in nests inside hollow trees; one nest was made of coconut fibres and grass (A. Loveridge in Kingdon 1974). Both pale and dark side-stripes are present in young when HB 90–100 mm (labels, BMNH). Predators, Parasites and Diseases

No information.

Conservation IUCN Category: Least Concern. Previously considered as Least Concern.

Paraxerus flavovittis

Habitat Savanna and forest, especially where there are Uapaca trees, and hardwood trees with holes (suitable for nesting). May also occur in cultivations. Abundance Common and widespread between the Rufigi and Rovuma rivers in Tanzania. Particularly numerous where there are many old hardwood trees (Kingdon 1974). Adaptations Diurnal and arboreal. These squirrels nest in the holes and hollows of hardwood trees, and sometimes nest in the roofs of houses (Kingdon 1974).

Measurements Paraxerus flavovittis HB: 172.7 (165–176) mm, n = 6 T: 168.8 (160–175) mm, n = 4 HF: 39.2 (35–40) mm, n = 6 E: 16.6 (15–18) mm, n = 5 WT: n. d. GLS: 40.1 (38.6–42.2) mm, n = 8 GWS: 22.5 (21.1–23.7) mm, n = 8 P3–M3: 4.7 (6.7–7.9) mm, n = 8 Body measurements: Lumbo, Mozambique (Hinton 1920) Skull measurements: Tanzania (BMNH) Key References

Kingdon 1974, 1997.

Chad E. Schennum & Richard W. Thorington, Jr

Paraxerus lucifer BLACK-AND-RED BUSH SQUIRREL Fr. Écureuil de brousse rouge et noir; Ger. Tanganjika-Buschhörnchen Paraxerus lucifer (Thomas, 1897). Proc. Zool. Soc. Lond. 1897: 430. Kombe Forest, Misuku Mts, Malawi.

Taxonomy Originally described in the genus Xerus (Paraxerus). Paraxerus lucifer is closely related to three other species, P. vincenti, P. palliatus and P. vexillarius. Synonyms: none. Chromosome number: not known.

at base with subterminal black band and bright rufous tip; below hairs bright rufous without banding; tail bands very indistinct. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 0 + 1 + 1 + 1 = 6.

Description Large bright rufous squirrel, sometimes with blackish patch on back. Pelage long and dense. Dorsal pelage bright rufous or russet with large patch of rufous-black in centre of back; dorsal hairs dark grey on basal half, bright rufous on terminal half, with black tip; numerous long pure black guard hairs in centre of back. Flanks bright rufous; hairs pale grey on basal half, bright rufous on terminal half. Ventral pelage, throat and chest dove-grey; hairs dove-grey with silver or whitish tip. Crown of head, cheeks and chin similar to dorsal pelage. Forelimbs and hindlimbs very bright rufous. Tail moderately long (ca. 60% of HB), similar in colour to dorsal pelage; above hairs pale rufous

Geographic Variation

None recorded.

Similar Species P. palliatus. Ventral pelage rufous. P. vexillarius. Ventral pelage grey, limbs rufous, sometimes with orange tail tip; Kilimanjaro and Usambara Mts, Tanzania only. P. vincenti. Ventral pelage and limbs rufous; Namuli Mountain, Mozambique only. P. cepapi.Yellowish brown without any rufous colour; widespread. P. flavovittis. Lateral whitish side-stripe. 81

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Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Recorded from montane habitats in N Malawi (Misuku Hills, Nyika Plateau), SW Tanzania (Poroto Mts, Nkuka Forest) (Ansell 1978, Ansell & Dowsett 1988). Not yet found in the Mafinga and Makutu Mts of NE Zambia where it might be expected to occur (Ansell 1978). Habitat Restricted to montane forests in isolated montane regions that have a high annual rainfall (Kingdon 1974). Abundance Maybe numerous in some habitats (Kingdon 1974). Adaptations Diurnal and arboreal. Tend to be rather noisy, with a loud and distinctive call. Foraging and Food Omnivorous. May forage on the ground.The diet includes vegetable matter, fruit, nuts, termites and ants (Kingdon 1974). Social and Reproductive Behaviour

No information.

Reproduction and Population Structure A single " caught in Sep was both pregnant and lactating, indicating two litters in close succession. Of eight "" examined in Mar and Apr, none was reproductively active (Kingdon 1974). Predators, Parasites and Diseases Ectoparasites reported: fleas of the genus Libyastus (Ansell & Ansell 1973). Conservation IUCN Category: Data Deficient. Previously considered as Least Concern. Measurements Paraxerus lucifer HB: 222 (201–241) mm, n = 10 T: 202 (186–218) mm, n = 10

Paraxerus lucifer

HF: 52.2 (48–55) mm, n = 13 E: 19.7 (15–22) mm, n = 12 WT: 496 (300–675*) g, n = 7 GLS: 55.6 (53.8–56.4) mm, n = 12 GWS: 31.2 (30.0–31.7) mm, n = 10 P3–M3: 9.3 (8.8–9.6) mm, n = 15 Nyika Plateau, Zambia (Ansell & Ansell 1973) *Three individuals of 1.5 lb (= 675 g) Key References

Ansell 1978; Kingdon 1974, 1997.

Chad E. Schennum & Richard W. Thorington, Jr

Paraxerus ochraceus OCHRE BUSH SQUIRREL Fr. Écureuil de brousse ocre; Ger. Ockerfarbiges Buschhörnchen Paraxerus ochraceus (Huet, 1880). Nouvelles Archives, Museum d’Histoire Naturelle, Paris, ser. 2, 3: 54. ‘Cette petite espèce provient de Bagamoyo, station de mos missionnaires, sur côte de Zanguebar ...’ Bagamoyo (06° 25´ S, 38° 54´ E), Tanzania.

Taxonomy Placed in genus Funisciurus (subgenus Paraxerus) by Kingdon (1974). Synonyms: affinis, animosus, aruscensis, augustus, capitis, electus, ganana, jacksoni, kahari, pauli, percivali, salutans. Amtmann (1975) recognized eight subspecies, but two of them were considered provisional; Kingdon (1974) recognized the five subspecies listed below. Chromosome number: not known.

behind the body. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 0 + 1 + 1 + 1 = 6.

Description Small squirrel with dull yellowish-brown pelage and slightly ringed tail. Dorsal pelage pale yellow, ochre or dark olive, grizzled. Pelage colour varies geographically (see below). Pale sidestripe in some subspecies. Ventral pelage yellow to off-white, not grizzled. Fore- and hindfeet same as dorsal pelage. Dorsal surface of fore- and hindfeet ochraceous.Tail long (ca. 100% of HB), with black and pale irregular bars and patches; tail is mostly held horizontally

P. o. aruscensis: NE Tanzania and SE Kenya. Pelage colour richer than P. o. ochraceus, with a yellow ventral pelage; without a side-stripe. P. o. electus: W Kenya. Pale form with white ventral pelage, perhaps without a side-stripe. P. o. ganana: S Ethiopia, NE Kenya and Tana River area. Small, pale, sandy-yellow coloured race, without a side-stripe.

Geographic Variation (1974):

Five subspecies recognized by Kingdon

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Paraxerus ochraceus

Paraxerus ochraceus.

P. o. jacksoni: S Kenya and most mountain forests in Kenya. The largest subspecies, generally darker than others with a rather green colouring and sometimes a pale side-stripe near the shoulder. P. o. ochraceus: Central and E Tanzania. A medium-sized subspecies; mostly sandy grizzled ochre, ventral pelage off-white; distinct side-stripe. Similar Species P. cepapi. Ventral pelage paler. Occurs further south, and probably is not sympatric with P. ochraceus.

Paraxerus ochraceus

Social and Reproductive Behaviour Live in pairs or small groups. Courtship involves a lot of chasing, mutual grooming and arching of the tail over the body.

Distribution Endemic to Africa. Somalia–Masai Bushland BZ. Recorded from Kenya and N Tanzania. Outliers in Somalia (near to Kenya/Ethiopian border) and in S Sudan.

Reproduction and Population Structure Little information. Observations in Kenya suggest that reproduction occurs in most months of the year. Litter-size: 2–3. Two "" with young may nest together.Young emerge from nest when 3–4 weeks of age. A mother may carry young in her mouth at times (A. Root in Kingdon 1974).

Habitat Tolerant of many habitats; recorded from diverse wooded savannas, riverine forests in semi-arid regions, and thickets from near sea level to at least 2000 m. Also recorded from plantations (coffee, Grevillea, Eucaplytus) and in suburban gardens (e.g. in Nairobi).

Predators, Parasites and Diseases Buzzards, snakes and genets are likely predators. Conservation

IUCN Category: Least Concern.

Abundance Very common in some localities. Adaptations Diurnal and arboreal. Most activity is in the early morning and late afternoon, with a rest period at the hottest time of the day. Ochre Bush Squirrels are very active, running quickly along branches and between the ground and the tops of trees. Calls include a high-pitched metallic ‘Burr’, which is emitted when threatened, accompanied by flicking of the tail. Foraging and Food Mostly vegetarian. Forages in the trees and on the ground. The diet includes fruits, seeds, buds, flowers, roots, bulbs, Acacia gum and occasionally animal matter (Kingdon 1997).

Measurements Paraxerus ochraceus HB: (!!): 155.1 (123–190) mm, n = 30 HB: (""): 163.5 (143–175) mm, n = 20 T: 162.5 (113–183) mm, n = 48 HF: 39.7 (34–45) mm, n = 52 E: 15.2 (10–27) mm, n = 45 WT: n. d. GLS: 41.3 (40.0–42.1) mm, n = 9 GWS: 23.5 (22.0–24.5) mm, n = 10 P3–M3: 6.8 (6.5–7.2) mm, n = 11 Kenya (USNM) Key References

Kingdon 1974, 1997.

Richard W. Thorington, Jr & Chad E. Schennum

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Family SCIURIDAE

Paraxerus palliatus RED BUSH SQUIRREL Fr. Écureuil de brousse à ventre roux; Ger. Rotbaüchiges Buschhörnchen Paraxerus palliatus (Peters, 1852). Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin 17: 273. Mainland near Mocambique Island ‘Africa orientalis, Quintangonha, 15° Lat. Aust.’. Mozambique, near Mozambique I.

Taxonomy Originally described in the genus Sciurus. This species consists of a series of populations living in isolated forests. Kingdon (1974) suggested that some of the populations are hybrids between Paraxerus cepapi and P. palliatus, but Viljoen (1989) pointed out that this was unlikely because there are important differences between the species in behaviour and habitat. Paraxerus palliatus is closely related to three other species, P. lucifer, P. vexillarius and P. vincenti. The species exhibits considerable variation in pelage colour and pattern, such that Amtmann (1975) listed 11 subspecies but noted that this was probably an excessive number. Kingdon (1974) recognized three subspecies in the northern part of the range and Viljoen (1989) recognized four subspecies in southern Africa; all seven subspecies are recorded here. Body size and colouration seem to be influenced greatly by habitat, with larger darker squirrels in moist forest and smaller paler squirrels in dry forest (Viljoen 1989).The form vincenti, recognized as a valid species here, was classified as subspecies (P. p. vincenti) by Kingdon (1997). Synonyms: auriventris, barawensis, bridgemani, frerei, lastii, ornatus, sponsus, suahelicus, swynnertoni, tanae, tongensis. Subspecies: seven. Chromosome number: not known. Description Medium to large arboreal squirrel with rufous or yellowish ventral pelage, variable geographically (see below, and Measurements). (Description for P. p. palliatus.) Dorsal pelage brownish, grizzled with buff. No side-stripes. Ventral pelage bright rufous. Crown of head grizzled brown, cheeks rufous. Fore- and hindlimbs, fore- and hindfeet rufous. Tail long (ca. 90% of HB), bushy, grizzled brown at base, bright rufous on terminal two-thirds. Individuals in drier climates are smaller (mean 210 g) than those in humid climates (mean 380 g). Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 0 + 1 + 1 + 1 = 6. Geographic Variation P. p. tanae (including barawensis): S Somalia, E Kenya and N Tanzania (south to Pangani R.). Tail completely rufous-orange. P. p. frerei (including lastii): Mafia and Zanzibar Is. Similar to coastal populations of P. p. palliatus, but with black feet. P. p. palliatus (including suahelicus): E coastal Tanzania and N Mozambique. See Description above. P. p. swynnertoni: Chirinda Forest, E Zimbabwe. Dorsal pelage grizzled black and buff; ventral pelage cinnamon-rufous. Face coloured like dorsal pelage, cheeks like ventral pelage. Colouration resembles P. palliatus and P. p. ornatus, but the different colouration of the face, fore- and hindfeet, and its smaller size, distinguishes it from these two. P. p. bridgemani (including auriventris and tongensis): E coastal Mozambique (south of the Save R.). Dorsal pelage dark brown, grizzled; ventral pelage orange; generally paler and more ‘yellowish’ than other subspecies. (This may be a separate species – Amtmann 1975.)

Paraxerus palliatus

P. p. sponsus. Perhaps indistinguishable from P. p. palliatus with its brown dorsal pelage and bright rufous ventral pelage (see above), but geographically very close to P. p. bridgemani. P. p. ornatus: South Africa (Ngoye Forest, Eshowe District, Zululand). Large; dorsal pelage dark brownish-black grizzled with buff; ventral pelage orange-rufous; tail dark brownish-black tinged with rufous. Similar Species Paraxerus spp. Without brightly coloured ventral pelage (i.e. not rufous, orange-rufous, yellowish). P. vincenti. Similar in most respects, but restricted to Namuli Mountain, Mozambique. Distribution Endemic to Africa. Coastal Forest Mosaic BZ of eastern Africa and some adjacent parts of the Zambezian Woodland BZ, especially extending inland along riverine forests and on some inland montane forests. Recorded from S Somalia, E Kenya, E Tanzania, Malawi, E Mozambique, E Zimbabwe and South Africa (KwaZulu–Natal), Mafia and Zanzibar Is. Some populations occur far inland along riverine forests (e.g. along the Tana and Ruaha Rivers; Kingdon 1974). Habitat Dry to wet forests, preferring woodlands with shady thickets; dune forests and evergreen moist forests in Mozambique and South Africa. Montane populations up to about 2000 m (e.g. Mt Mlanje, Malawi).

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Paraxerus poensis

Abundance Varies according to locality. In favoured habitats in South Africa, mean density was 4.32 squirrels/ha (P. p. ornatus) and 2.15 squirrels/ha (P. p. bridgemani) (Viljoen 1986). Biomass varies with locality: in Ngoye Forest (South Africa) was 1659 g/ha (maximum) and 595 g/ha in Mkwakwa Forest (Viljoen 1986). Adaptations Diurnal and arboreal. Also spends a considerable amount of time on the ground, but the proportion of time spent foraging on the ground is uncertain. Nests in holes in baobabs and Kigelia trees. Foraging and Food Omnivorous, feeding on seeds, fruits, nuts and invertebrates. Seems to prefer the seeds to the fleshy parts of the fruit. Viljoen (1983a) reported that captive squirrels ate raw liver and biltong. Drinks water when available, but does not appear to be dependent on it. Red Bush Squirrels are scatter hoarders of larger seeds, although they do not hoard large quantities of food as do squirrels of more temperate climates. They scratch under bark to get at insects, and Viljoen (1983a) reported that in captivity, they displayed hunting behaviour when stalking invertebrates.

Reproduction and Population Structure In the wild "" have 1–2 young/litter, and probably one litter/year. In captivity, multiple litters per year are possible. One lactating " recorded in March in S Kenya (Kingdon 1974). In southern Africa, young born during the warm wet season (Aug–Mar) (Smithers 1983). Gestation: 60–65 days. At birth, young weigh 13–14 g. Eyes open Day 7–10. Leave nest about Day 18. Weaned by Day 40 (Viljoen 1980). Predators, Parasites and Diseases Uncertain, but it would be expected that snakes and arboreal mongooses would prey on young, and that hawks and raptors would be predators of adults. Conservation IUCN Category: Least Concern. Previously considered as Vulnerable.

Measurements Paraxerus palliatus HB (P. p. ornatus): 221.5 ± 9.2 mm, n = 30 HB (P. p. bridgemani): 187.7 ± 9.9 mm, n = 11 T (P. p. ornatus): 203.8 ± 8.6 mm, n = 86 T (P. p. bridgemani): 176.8 ± 9.1 mm, n = 28 Social and Reproductive Behaviour Although Red Bush HF (P. p. ornatus): 51.8 ± 1.7 mm, n = 103 Squirrels are quite common and diurnal, they are shy animals. HF (P. p. bridgemani): 43.9 ± 1.8 mm, n = 36 Normally observed as solitary individuals or in pairs (Ansell & Dowsett E (P. p. ornatus): 20.6 ± 1.4 mm, n = 75 1988), although several squirrels may nest together in larger groups E (P. p. bridgemani): 19.4 ± 1.5 mm, n = 32 (mean 3.1 ± 1.2; Viljoen 1986). They nest in tree holes. Home- WT (P. p. ornatus): 368.2 ± 22.4 g, n = 104 range varies according to habitat and sex: in evergreen moist forest, WT (P. p. bridgemani): 209.1 ± 19.2 g, n = 60 mean home-range (P. p. palliatus) was 3.18 ha (!!) and 2.19 ha GLS (P. p. ornatus): 50.9 ± 1.6 mm, n = 11 (""), and in coastal forest and thickets, mean home-range (P. p. GLS (P. p. bridgemani): 45.3 ± 0.8 mm, n = 13 bridgemani) was 4.17 ha (!!) and 0.73 ha (""). Communication GWS (P. p. ornatus): 29.4 ± 1.3 mm, n = 11 between animals is maintained by visual, auditory and olfactory GWS (P. p. bridgemani): 26.3 ± 0.6 mm, n = 13 signals. Viljoen (1983b) recorded lots of tail flicking and fluffing, P3–M3: 9.6 (9.3–9.9) mm, n = 9 especially in dense habitats.Vocalizations include an array of murmurs, P. p. ornatus: Ngoye forest, South Africa (Viljoen 1989) hisses, growls, clicks, twitters and barks. Urine dribbling and anal P. p. bridgemani: L. St Lucia, South Africa (Viljoen 1989) dragging are also common. Males make murmuring vocalizations P3–M3: Ngoye forest, South Africa (BMNH) when chasing "". This may serve as a trigger to stimulate oestrus. Measurements given as ± 1 S.D. Female builds nest in a tree hole lined with leaves. She keeps the nest very clean while young are being reared.When offspring are very young Key References Kingdon 1974; Smithers 1983; Viljoen 1983a, b, "" are extremely protective and respond aggressively toward 1986, 1989. intruders, including the male. Later, !, " and young form a family Richard W. Thorington, Jr, Lindsay A. Pappas & group. When young reach subadulthood (early winter in southern Chad E. Schennum Africa), parents drive them away from the nest (Viljoen 1980).

Paraxerus poensis GREEN BUSH SQUIRREL Fr. Petit Écureuil de brousse; Ger. Grünes Buschhörnchen Paraxerus poensis (A. Smith, 1834). South Afr. Quart. J., 2nd ser., 2: 128. Fernando Poo (= Bioko I., Equatorial Guinea).

Taxonomy Originally described in the genus Sciurus. The species has been placed, at different times, in the genera Aethosciurus, Heliosciurus, Funisciurus and Paraxerus. Thomas (1916a) placed poensis into a new genus, Aethosciurus, citing molar differences to separate it from Heliosciurus and Funisciurus. Hollister (1919) placed it in Heliosciurus because of dental similarity, as did Ellerman (1940), who noted the similarity to Paraxerus. Moore (1959) placed it in Funisciurus, noting some cranial similarities. Rosevear (1969)

restored it to Aethosciurus, stating that it possessed characters of both Heliosciurus and Funisciurus and could not be placed unambiguously in either. Amtmann (1966) allocated the species to Paraxerus, even though its teeth differ from those of some members of that genus. Here, following Hoffman et al. (1993) and Thorington & Hoffman (2005), it is placed in the genus Paraxerus (see also Rosevear 1969 for a review). Synonyms: affinis, musculinus, olivaceus, subviridescens. Subspecies: none. Chromosome number: not known. 85

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suggest they prefer vegetation above 5 m (Emmons 1980). Nests are undescribed; anecdotal reports suggest that Green Bush Squirrels build arboreal, exposed, leaf nests (Emmons 1979a) or nest in tree hollows (Rosevear 1969). In captivity, nest boxes were lined with finely teased fibres (Emmons 1975). Foraging and Food Omnivorous. Forages arboreally. Six of 12 observations of foraging behaviour in the wild were of insect hunting by intense searching of stems, bark and arboreal crannies. In Gabon, the diet is mainly fruit and seeds (88% dry mass of stomach contents, n = 8) and arthropods (11%). The arthropods eaten are a miscellaneous mixture as might be expected from random searching (Emmons 1975, 1980). In captivity, Green Bush Squirrels were adept at capturing flying insects, which they pursued eagerly, and they opened and ate the eggs of small birds (Emmons 1975).

Paraxerus poensis

Description Small, greenish-olive squirrel with a slender tail. Pelage soft, thick and dense. Dorsal pelage dark grizzled goldengreen; hairs black at base with greenish-yellow tips. No lateral side-stripe. Ventral pelage pale yellow, thickly furred; hairs grey at base with long yellow tip. Cheek with yellow above and below eye, bordering a darker stripe to base of ear. Fore- and hindlimbs dark golden-green. Tail long (ca. 105% of HB), slender, not bushy, slightly darker than dorsal pelage. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: 1 + 0 + 1 + 1 = 6. Geographic Variation None recorded. Similar Species P. cooperi. Slightly larger (HB: 192–212 mm); rufous colour on foreand hindlimbs; sympatric with P. poensis only in parts of Cameroon. Myosciurus pumilio. Much smaller, with inner and outer surface of ears white; sympatric with M. pumilio in west of range. Distribution Endemic to Africa. Rainforest BZ except for eastern part of the East Central Region. Distributed in three disjunct populations: (a) from Sierra Leone to Ghana, (b) Cameroon, Equatorial Guinea (Rio Muni and Bioko I.), Gabon and Congo and (c) N DR Congo. Paraxerus poensis is the only small, greenish arboreal squirrel in its geographic range. Habitat Lowland evergreen moist rainforests, secondary forest and tangles. Often seen in farmlands and plantations (Rosevear 1969). Abundance Frequently seen. Adaptations Diurnal and arboreal. Green Bush Squirrels are morphologically adapted for arboreal life, with short limbs and short, broad feet with strong, curved claws. Most observations

Social and Reproductive Behaviour Usually seen singly (68%, n = 44) or in pairs (18%) (Emmons 1980). The social organization is undescribed, but in captivity heterosexual pairs housed together showed extreme cohesion and many bonding behavioural characteristics (e.g. always sharing a nest box, grooming each other, resting in physical contact). After " gave birth, ! continued to share the nest box and showed strong parental behaviour towards young when they emerged. Female showed no aggression towards ! at parturition or any other time (L. Emmons, unpubl.). These behaviours suggest that these squirrels live in monogamous pairs. Vocalizations are described in detail in Emmons (1978). The single type of alarm call is a loud buzz composed of about 25 pulses emitted in a one-second burst; pulses are so close together that they are indistinguishable to the human ear. Calls are emitted singly, but can be repeated more than 100 times. The alarm call is always associated with a highly stereotyped visual display in which the squirrel sits, stands, or moves along a branch with the tail held out stiffly behind, its tip curved upward. While it emits a call, the squirrel freezes, then immediately jerks the tail sharply upward until its base is nearly vertical, maintaining a stiff C-shape. As the tail goes up, the hindfeet alone, or both fore- and hindfeet, are hopped or stamped, often moving the body slightly forward. The tail then relaxes to the horizontal and the display may be repeated (see also Heliosciurus rufobrachium). Reproduction and Population Structure Litter-size: 1–2, with more litters of one than of two (Emmons 1979a). Predators, Parasites and Diseases No information. Conservation

IUCN Category: Least Concern.

Measurements Paraxerus poensis HB: 153 (148–161) mm, n = 5 T: 160 (150–165) mm, n = 5 HF: 33 (30–35) mm, n = 5 E: 13 (11–15) mm, n = ?* WT: 104.5 (101–114) g, n = 4 GLS: 38.5 (37.7–39.1) mm, n = 3 GWS: 21.4, 21.7 mm, n = 2 P3–M3: 6.5 (6.3–6.9) mm, n = 4

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Paraxerus vexillarius

Gabon (Emmons 1975, L. Emmons unpubl.) *Rosevear 1969

Key References

Emmons 1978, 1980; Rosevear 1969. Louise H. Emmons

Paraxerus vexillarius SWYNNERTON’S BUSH SQUIRREL Fr. Écureuil de brousse de Kershaw; Ger. Lushoto-Buschhörnchen Paraxerus vexillarius (Kershaw, 1923). Ann. Mag. Nat. Hist., ser. 9, 11: 591. Lushoto, Wilhelmsthal, Usambara, Tanzania.

Taxonomy Originally described in the genus Funisciurus. It is possible that the two forms, vexillarius and byatti, given here as subspecies, are valid species (e.g. as in Allen 1939 and Ellerman 1940). Amtmann (1975) regarded them as forms of P. vexillarius and also commented that they may be distinct species. Inexplicably, Kingdon (1974) treated P. byatti as a subspecies of P. lucifer. See Allen and Loveridge (1933). Synonyms: byatti, laetus. Subspecies: two. Chromosome number: not known. Description Large squirrel with grizzled brownish dorsal pelage and sometimes with rufous-orange tip to tail. Dorsal pelage olivegreen to brown, grizzled. Ventral pelage dove-grey. Limbs and feet rufous-orange. Tail long (ca. 85% of HB), brown and whitish rings at base, bright orange at tip; hairs at base banded black and white. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: not known. Geographic Variation P. v. byatti: Mt Kilimanjaro, Tanzania. Very similar to P. v. vexillarius, but described as being more ochraceous-buff on the back and darker ventrally. This name is usually attributed to animals throughout the range of P. vexillarius lacking an orange tail tip, a condition found only among some individuals of the Usambara population, as suggested in the original description of P. vexillarius. Perhaps this subspecies should not be recognized. P. v. vexillarius: Usambara, Uluguru and Uzungwa Mts, Tanzania. Dorsal pelage grizzled brown, greyer on the sides, and dull rufous on the flanks and outside of thighs; arms and shoulders dark rufous, fore- and hindfeet tawny-ochraceous. Tail coloured like back for proximal one-fifth of its length; distally more buff; hairs sometimes tipped more white. Rufous on nose, around mouth, and a broad streak running through the eye to the ear. This name is

Paraxerus vexillarius

usually attributed only to animals with an orange tail tip, but this is probably a local variation within the Usambara population of P. vexillarius, as noted by Kershaw (1923). Similar Species P. palliatus. Rufous or orange ventral pelage; more widespread distribution. P. lucifer. Predominantly rufous with black dorsal patch; limited distribution in S Tanzania and N Malawi.

Paraxerus spp., pelage colours. Left to right: P. cooperi, P. lucifer, P. palliatus, P. poensis and P. vexillarius.

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Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Mountains of NE to S Tanzania (details as above). Habitat

Montane forest.

Abundance No information. Remarks Arboreal. Feeds on fruits and seeds. Conservation IUCN Category: Near Threatened. Survival of this species is threatened because the montane forests of the Usambara Mts are being fragmented and are decreasing in area as a result of expanding agriculture.

Measurements Paraxerus vexillarius HB: 230 (190–264) mm, n = 4 T: 195 (180–210) mm, n = 4 HF: 49.7 (45–52) mm, n = 4 E: 18, 15 mm, n = 2 WT: n. d. GLS: 54, 53 mm, n = 2 GWS: 33.6, 31 mm, n = 2 P3–M3: n. d. Tanzania (P. vexillarius type specimen and P. v. byatti type specimen [Kershaw 1923]; USNM [2]) Key References

Kingdon 1974, 1997.

Richard W. Thorington, Jr & Chad E. Schennum

Paraxerus vincenti VINCENT’S BUSH SQUIRREL (SELINDA MOUNTAIN SQUIRREL) Fr. Écureuil de brousse de Vincent; Ger. Vincents Buschhörnchen Paraxerus vincenti Hayman, 1950. Ann. Mag. Nat. Hist., ser. 12, 3: 263. Namuli Mountain, Mozambique. 5000 ft (= 1500 m).

Taxonomy Classified as a subspecies of Paraxerus palliatus by Kingdon (1994, 1997), but considered here to be a valid species because of its very limited distribution and its isolation from all subspecies of P. palliatus. Paraxerus vincenti is closely related to three other species, P. lucifer, P. palliatus and P. vexillarius. Synonyms: none. Chromosome number: not known. Description Medium-sized squirrel very similar to Paraxerus palliatus ornatus. Dorsal pelage blackish, grizzled. Ventral pelage rich rufous. Crown of head and cheeks darkish-brown, rufous around eye and nasal region. Fore- and hindlimbs and upper surfaces of feet similar to dorsal pelage. Tail long (ca. 100% of HB), blackishbrown tipped with rufous; hairs black at base, rufous at tip. Skull: cheekteeth 5/4; posterior end of bony palate in line with posterior end of M3; masseteric tubercle not prominent. Nipples: not known. Geographic Variation None recorded. Similar Species Paraxerus spp. Ventral pelage less intense and less bright. Hayman (1950) noted that P. palliatus has ‘brick-red’ ventral pelage whereas in P. vincenti is rich rufous. Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Recorded only from Namuli Mountain, N Mozambique (15°21´ S, 37°04´ E). Habitat

Moist evergreen forests (Viljoen 1989).

Remarks Apparently no other information available. Conservation

IUCN Category: Endangered.

Paraxerus vincenti

Measurements Paraxerus vincenti HB: 212 ± 5.9 mm, n = 5 T: 209 ± 8.9 mm, n = 5 HF: 46.6 ± 2.0 mm, n = 5 E: 21.1 ± 0.6 mm, n = 5 WT: n. d. GLS: 50.4 ± 0.4 mm, n = 5 GWS: 29.8 ± 0.3 mm, n = 5 P3–M3: n. d.

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Key References

Mozambique (Viljoen 1989) Measurements given as mean value ± 1 S.D.

Hayman 1950; Viljoen 1989.

Richard W. Thorington, Jr & Chad E. Schennum

GENUS Protoxerus Forest Giant Squirrel Protoxerus Forsyth Major, 1893. Proc. Zool. Soc. Lond. 1893: 189. Type species: Sciurus stangeri Waterhouse, 1842.

Monotypic genus. A very large tree squirrel with a wide distribution in the Rainforest BZ and some adjacent rainforest relicts in the Rainforest–Savanna Mosaics. The single species Protoxerus stangeri lives in the upper storeys of rainforest, feeds primarily on nuts and fruits, and tends to be solitary. The genus is characterized by very large size (HB 270–306 mm) and long tail, about the same length as the head and body. The pelage is longer and denser than in Heliosciurus spp., and the hairs have three or five bands, which result in a strongly speckled effect. Flanks without longitudinal stripes. The ventral surface is mostly hairless. The tail is long and bushy, and about as long as HB; each hair with 7–9 bands and white at the tip on the dorsal and lateral sides. Females have four pairs of nipples (as in Epixerus); !! have large baculum. The skull is characterized by large size (66–73 mm in length) and is larger than in any other African squirrel; four upper cheekteeth; supraorbital notch closed on margin of orbit forming a foramen piercing the frontal bone; posterior end of bony palate in line with posterior end of M3; anterodorsal process of premaxilla rises to abut evenly with anterolateral angle of nasal; pronounced masseteric ridge; and masseteric tubercle absent or very small (Figure 15).

Protoxerus forms a monophyletic clade with Heliosciurus, Allosciurus and Epixerus (Moore 1959). Protoxerus resembles the related genera Heliosciurus, Allosciurus and Epixerus in relative length of tail; differs from Heliosciurus but agrees with Epixerus in having a small masseteric tubercle (absent in some individuals) and four pairs of nipples. Protoxerus also resembles Epixerus in speckled pelage of upperparts, contrasting sparse pelage of underparts and banded pattern of tail. Some authors have allocated Allosciurus to Protoxerus as a synonym or subgenus (Moore 1959, Amtmann 1966, Thorington & Hoffman 2005), though in many features, Protoxerus stangeri is more similar to Epixerus than to Allosciurus. Protoxerus and Allosciurus have not been recorded to share derived character-states absent in other squirrels, so are not demonstrably sister-taxa. Epixerus ebii has sometimes been misidentified as Protoxerus stangeri but their skulls have been compared in considerable detail, and can be readily distinguished. For differences between Protoxerus and Allosciurus, see Allosciurus genus profile. Peter Grubb

Protoxerus stangeri.

Figure 15. Skull and mandible of Protoxerus stangeri (RMCA 16124).

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Protoxerus stangeri FOREST GIANT SQUIRREL (AFRICAN GIANT SQUIRREL) Fr. Écureuil géant de Stanger; Ger. Afrikanisches Riesenhörnchen Protoxerus stangeri (Waterhouse, 1842). Proc. Zool. Soc. Lond. 1842: 127 (publ. 1943). Fernando Poo (= Bioko I., Equatorial Guinea).

Taxonomy Originally described in the genus Sciurus. This species shows considerable geographic variation, which has resulted in 19 named forms; 11 of these may be considered as subspecies (Amtmann 1975, Thorington & Hoffman 2005), although the validity of these subspecies is uncertain. Synonyms: bea, calliurus, caniceps, centricola, cooperi, dissonus, eborivorus, kabobo, kwango, loandae, moerens, nigeriae, nordhoffi, notabilis, personatus, signatus, subalbidus, temmincki, torrentium. Subspecies: 11. Chromosome number: not known. Description Very large long-bodied squirrel with very long ringed tail; the largest African arboreal squirrel. Pelage short and stiff. Dorsal pelage medium brown, grizzled with yellow or buff; hairs banded, black at base, yellow or buff on terminal half, with black tip; some hairs have five alternating dark-pale bands. Ventral surface naked yellow skin; almost hairless. Head large and rounded with conspicuously large cheek muscles. Crown and nasal region of head similar to back except terminal half of hairs are white, giving a frosted appearance to the head. Cheeks thinly haired showing yellow skin below and behind eye; ears short, nearly naked, yellowish. Chest usually white. Limbs and feet short and robust. Tail long (ca. 100% of HB), extremely bushy, mostly black and white; hairs long with 7–9 bands of black and white, with white tips, which result in pale (but sometimes obscure) rings along the length of the tail; undersurface of tail varied, black and white, but not always forming rings. Tail carried straight out behind the body during locomotion, and hangs below body while squirrel is at rest; not normally curled against back. Skull and mandible large and heavily built; cheekteeth 4 /4; posterior end of bony palate in line with posterior end of M3; supraorbital ridge with small foramen; pronounced masseteric ridge; masseteric tubercle inconspicuous or absent . Females slightly larger, on average, than males. Nipples: 1 + 1 + 1 + 1 = 8. Geographic Variation Amtmann (1975) lists 11 subspecies, as follows, without comment: P. s. bea: Kakamega Forest, Kenya. P. s. centricola: between Congo and Oubangui rivers, DR Congo; forest relics in Uganda; Mt Kungwe, Tanzania; Equatoria District, S Sudan. P. s. eborivorus: E Nigeria south to Ogooué R., Gabon, and east to Oubangui R., DR Congo. P. s. kabobo: Montane forests of Mt Kabobo, E DR Congo. P. s. kwango: near Kasonga Lunda, Bandundu Province, SW DR Congo. P. s. loandae: N Angola. P. s. nigeriae: between Volta R., Ghana and Niger R., Nigeria. P. s. personatus: between Ogooué R., Gabon and Congo R., DR Congo. P. s. signatus: between Congo R. and Kasai R., DR Congo. P. s. stangeri: Bioko I. P. s. temmincki: Sierra Leone to Volta R., Ghana.

Protoxerus stangeri

Similar Species Epixerus ebii. Reddish head, slender body, long legs; ventral surface with yellow skin; mostly terrestrial. Heliosciurus spp. Smaller, crown the same colour as back, tail slender, ventral surface well haired, without visible yellow skin; arboreal. Distribution Endemic to Africa. Widely distributed in the Rainforest BZ and Rainforest–Savanna Mosaics from Sierra Leone to the Kavirondoro district of Kenya, and from N Angola to N Tanzania. Recorded from N Angola, Cameroon, Central African Republic, Congo, Côte d’Ivoire, Equatorial Guinea (Rio Muni and Bioko I.), Gabon, Ghana, W Kenya, Liberia, Nigeria, Sierra Leone, N Tanzania, Togo, Uganda and DR Congo. Habitat Lowland evergreen rainforest and rainforest outliers, mostly at lower altitudes. Within this area, occupies many forest types including tall mature forests, secondary forests, plantations and gardens with trees. Abundance Common. One of the most abundant and frequently seen species of rainforest squirrels. Adaptations Diurnal and arboreal, living mostly in the forest canopy and upper vegetation levels (Emmons 1980) but descending to the ground occasionally (J. C. Ray & J. R. Malcolm unpubl.). The short limbs with short, broad palmar and plantar surfaces are suitable for running along large branches and up and down tree trunks

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Protoxerus stangeri

(Emmons 1980). The incisor teeth are robust, and the masseter muscles are massive, as befits its diet of hard nuts (see below). A prominent glandular pocket in the inside corner of the lips may be used in scent marking. Forest Giant Squirrels nest in tree hollows and favour those with entrances just large enough for the squirrel to squeeze through (mean height 9.3 m; range: 4–17 m; n = 7). The nest within the hollow is constructed of twigs with sprays of attached green leaves (Emmons 1975). Animals leave their nests at dawn (range 06:00– 06:08h); in Gabon they have a relatively long activity period (mean 10.46 h/day) and return to their nests in the late afternoon or well before dusk (mean entry time 16:33h; range: 15:25–17:55h, n = 16; Emmons 1980).

of a mean of 12 modulated pulses that drop in frequency about 300 Hz from beginning to the end of a call, with a mean call duration of 1.3 s. Calls are repeated at intervals of 5–20 s (Emmons 1978). Alarm calls in this species are not accompanied by displays of the tail other than erection of the tail hairs; however, when approaching a strange object, Forest Giant Squirrels may fluff out the tail, raise the base vertically over the back with the tip curled posteriorly downwards, and slowly wave it from side to side (Emmons 1978).

Foraging and Food Primarily vegetarian. In Gabon, feeds mainly on seeds extracted from fruits of trees and lianas, notably from the thick-walled nuts of Panda oleosa, Coula edulis and species of Klainodoxa, Irvingia, Elaeis and many other species. The diet is supplemented by minor amounts of plant vegetative parts (9%) and insects (0.4%) (Emmons 1980). Many of the fruits whose seeds are eaten by Forest Giant Squirrels are primarily dispersed by elephants (Gautier-Hion et al. 1980). Ten of 60 observations of feeding in the wild were of insect-hunting behaviour (Emmons 1980), but wild-caught captive Forest Giant Squirrels had little predatory tendency and ignored both live and dead birds, and bird’s eggs (Emmons 1975).

Predators, Parasites and Diseases Preyed upon by eagles and other large raptors. Hunted for its meat where larger game is scarce.

Social and Reproductive Behaviour Mostly asocial. Forest Giant Squirrels forage, travel and nest alone; 80% of sightings were of solitary animals, the remainder were either aggressive encounters or mothers followed by young. Individuals appear to avoid each other and can chase conspecifics out of fruiting trees. In captivity, squirrels housed together avoid each other, and there is a complete absence of contact, mutual grooming, chasing, or sharing of nest boxes. Apart from these notes, which suggest a solitary life-style, the social organization is undescribed. An apparent ‘mating chase’ in which several calling !! pursued a presumably oestrous ", was seen once by Emmons (1980). Two subadult "" (followed by radio tracking) had home-ranges of 3.2 and 5.0 ha (Emmons 1975). Vocalizations most often heard in the wild are two types of alarm calls: the low intensity alarm consists of repeated soft to loud sniffs or sneezes, sometimes alternating with clicks of the incisors; and the unique, diagnostic, high intensity alarm call is a loud whinny composed

Reproduction and Population Structure There are few records; litter-size seems to be 1–2, with more litters of one than of two (Emmons 1979a). One juvenile (HB 172 mm) in Bwamba, Uganda, in Sep (label, BMNH).

Conservation IUCN Category: Least Concern. Could be threatened by destruction and fragmentation of rainforest, and by hunting where terrestrial game species are rare. Measurements Protoxerus stangeri HB (!!): 297 (270–306) mm, n = 13 HB (""): 300 (294–305) mm, n = 10 T (!!): 271 (250–335) mm, n = 11 T (""): 317 (300–340) mm, n = 10 HF (!!): 63.7 (61–67) mm, n = 10 HF (""): 64.0 (60–67) mm, n = 7 E: 22 (18–25) mm, n = ?* WT (!!): 685 (536–769) g, n = 13 WT (""): 701 (680–730) g, n = 8 GLS: 67.8 (65.9–70.0) mm, n = 7 GWS: 37.2 (34.5–38.7) mm, n = 7 P4–M3: 11.2 (10.5–11.6) mm, n = 5 Gabon (Emmons 1975, L. Emmons unpubl.) *Rosevear 1969 Key References

Emmons 1975, 1978, 1980. Louise H. Emmons

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GENUS Sciurus Squirrels Sciurus Linnaeus, 1758. Syst. Nat., 10th edn., 1: 63. Type species: Sciurus vulgaris Linnaeus, 1758.

The genus contains 28 species widely distributed in North America and South America with a few species in Europe, the Middle East, parts of Asia, and Japan. Since the genus is not indigenous to Africa, and is represented by only one introduced species in a limited area

of South Africa, details of genus are not given (see Thorington & Hoffman 2005 for further details). The single species in Africa is Sciurus carolinensis.

Sciurus carolinensis GREY SQUIRREL Fr. Écureuil Gris; Ger. Grauhörnchen Sciurus carolinensis Gmelin, 1788. In: Linnaeus, Syst. Nat., 13th edn, 1: 148. Carolina, USA.

Taxonomy This profile refers to the species in Africa only. For general information of the species extralimitally to Africa, see Koprowski (1994), Thorington & Hoffmann (2005) and Southern (1964). Chromosome number: 2n = 40, FN = 76. Description Very large grey squirrel with bushy tail. Hairs rather coarse. Dorsal pelage yellowish-brown or greyish-brown (summer) to silvery-grey (winter); hairs ca. 10 mm; scattered black guard hairs (ca. 15 mm). Often slightly darker (and/or browner) along middorsal region. Ventral pelage paler, usually grey to greyish-white; hairs less dense; ca. 8 mm. Flanks sometimes with rufous streak at junction of dorsal and ventral pelages. Head similar in colour to dorsal pelage. Ears small, rounded; small white or yellow tuft on each ear tip (winter only). Limbs short but moderately long for a

squirrel; similar in colour to dorsal pelage, sometimes with rufous on upper surfaces; digits with well developed claws. Tail long (ca. 75% of HB), densely covered with long hair (typically 30–40 mm, but up to 45 mm); very large and bushy; yellowish-brown, with subterminal black band and white tip (which gives a ‘frosted’ appearance); tail hairs can be flattened or erected depending on mood. When animal is at rest, tail is held horizontally over back, with the tip pointing vertically upwards. Upper toothrow with five cheekteeth (the anterior tooth being a very small peg-like P3, and sometimes absent) (Figure 16). Albino individuals may occur in some populations (Britain and South Africa). Nipples: 1 + 1 + 1 + 1 = 8. Geographic Variation

None recorded in Africa.

Similar Species Heliosciurus rufobrachium. Similar in size; pelage much redder; limbs rufous-red. Distribution Introduced from North America (via Britain) in ca. 1890–1900 by Cecil Rhodes (at Groote Schuur Estate in Cape Town). Confined to a small area of SW Cape Province, South Africa (in the regions of Cape Town, Stellenbosch, Paarl, Elgin, Swellandam and Ceres). Details of the expansion of Grey Squirrels within South Africa are documented by Davis (1950), Millar (1980) and Lever (1985). Natural expansion of this range is unlikely because of the surrounding unsuitable habitat (e.g. fynbos); currrently, the species occurs in suitable patches of habitat within an area of ca. 7000 km2, and is confined to urban, agricultural and afforested environments (Long 2003). Extralimitally widespread as an indigenous species in the USA and Canada; introduced into Britain, Ireland and to parts of N Italy; also to parts of Victoria State, Australia (1880–1973, but now extinct) (Lever 1985, Long 2003). Map not given. Habitat Woodlands with suitable food trees including oaks, selected species of pines, eucalypts, acacias etc. May also occur in fruit orchards if suitable woodlands are nearby. Does not occur in monocultures such as pine plantations, nor in indigenous forests. Figure 16. Skull and mandible of Sciurus carolinensis (BMNH 2004.96).

Abundance No information; tends to be common in suitable habitats in natural distribution and where introduced successfully.

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Adaptations Diurnal and arboreal, although may descend to the ground to feed and cross small open spaces. Builds spherical nests (dreys) of twigs, 30–60 cm diameter, amongst branches and twigs of trees; may sometimes live in natural cavities in trees. Activity outside nest varies seasonally and is influenced by temperature. In warmer months with longer days, activity is from after dawn until ca. 10:00h and after 16:00h until near dusk; in cooler months with shorter days, activity is mostly between 10:00h and 14:00h (Millar 1980 in Smithers 1983). In the northern hemisphere, Grey Squirrels enter torpor on cold days and do not leave the drey for several days at a time; whether torpor occurs in South African Grey Squirrels is not recorded. Foraging and Food Herbivorous, mostly frugivorous. Foods in South Africa include acorns, pine nuts and other deciduous fruits. Occasionally omnivorous; feeding, in addition, on bird’s eggs, fungi, pollen, insects, bark and leaves (Millar 1980 in Smithers 1983). The majority of the diet is provided by acorns and pine nuts, which together provide a balanced diet of carbohydrate, protein and fat (acorns: protein 4.9%, fat 5%, carbohydrate 84.5%; pine nuts: protein 31%, fat, 47.4%, carbohydrate 11%) (Millar 1980 in Smithers 1983). Social and Reproductive Behaviour Usually solitary; may occur in small groups (mother and young). May rest in dreys in small groups (especially in cooler weather).Young are born in the dreys. Reproduction and Population Structure In South Africa, mating occurs in Jul–Aug and in Nov–Dec; young are born in Aug–Nov (spring) and Dec–Feb (summer). Embryo number: 2.5 (1–4, mode 2 and 3). Litter-size tends to be smaller in spring (mean 2.2, n = 11) and

larger in summer (mean 2.6, n = 16) (Millar 1980 in Smithers 1983). General information from elsewhere: gestation 44 days; young weaned at ca. Week 7; breeding first occurs in second year of life when 11–16 months of age. Females sometimes have two litters per year. Predators, Parasites and Diseases Conservation hemisphere).

No information.

IUCN Category: Least Concern (in northern

Measurements Sciurus carolinensis TL: 498 (431–572) mm, n = 250 HB: (ca. 282 mm)* T: 216 (115–269) mm, n = 250 HF: 60 (51–67) mm, n = 179 E: 25–33 mm** WT: 579 (434–750) g, n = 256 GLS: 59.0 (54.6–62.9) mm, n = 13 GWS: 34.2 (32.8–36.1) mm, n = 12† P3–M3: 10.9 (10.1–11.4) mm, n = 10† South Africa (Millar 1980 in Smithers 1983) *Calculated mean (mean TL – mean T) **North America; mean and sample size not given (Koprowski 1994) †Southern England (BMNH); P4–M3 where P3 is absent Key References

Koprowski 1994; Millar 1980; Smithers 1983. D. C. D. Happold

GENUS Xerus Ground Squirrels Xerus Hemprich and Ehrenberg, 1833. Symb. Phys. Mamm., vol. 1, sig. Ee, pl. 9. Type species: Sciurus (Xerus) brachyotus Hemprich and Ehrenberg, 1832 (= Sciurus rutilus Cretzchmar, 1828).

The genus Xerus is endemic to Africa, and comprises four species distributed throughout the semi-arid regions of the continent. Placed in the Tribe Xerini (together with Atlantoxerus, the only other genus of ground squirrels). Typical habitats are semi-arid desert, grassland and lightly wooded savannas. Only one species, Xerus princeps, inhabits rocky, hilly ground. The genus is distinguished by its bristly fur and small ear pinnae. Skull characteristics include cheekteeth 5/4 although upper anterior premolar (P3) very small and often absent in adults; palate long (ca. 62% of occipital-nasal length) with the posterior end of bony palate well posterior to M3 (as in Atlantoxerus and unlike other sciurids); masseteric tubercle very prominent; lachrymal enlarged; jugal joining the lacrimal with a blunt truncation, supraorbital notch small (occasionally absent), posterior end of bony palate well posterior to M3, and masseteric tubercle prominent (Figure 17). Dental formula: I 1/1, C 0/0, P 2/1, M 3/3 = 22 (but see also Xerus erythropus). There are two pairs of nipples, although Xerus erythropus has three pairs (Moore 1961). Little is known about the biology and reproduction except for Xerus inauris. All four species are diurnal, terrestrial and semi-

Xerus erythropus.

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Figure 17. Skull and mandible of Xerus erythropus (BMNH 50.21). The first premolar is missing in this specimen.

fossorial. All, except for X. inauris, are asocial. Burrows are usually simple, with 2–6 openings. In Xerus inauris, groups of "" maintain complex burrows with multiple burrow openings (up to 60). All four species are mainly herbivorous, feeding on grasses, leaves, fruits and nuts. Xerus inauris and X. rutilus can breed throughout the year. Litter-size in the genus appears to range from 1 to 6 young/litter. The genus Xerus, together with Atlantoxerus and the non-African Spermophilopsis, was first placed in the tribe Xerini by Simpson (1945). Nadler & Hoffman (1974) supported the classification of Xerus rutilus as a member of the tribe Xerini rather than in a subfamily of its own and subsequently others have supported this classification. The currently recognized species have been described under numerous names, resulting in a long list of synonyms. The genus was split into two subgenera, Euxerus and Geosciurus, by Simpson (1945). Moore (1959), following Pocock (1923), supported Simpson’s classification; however, Moore suggests that Euxerus might deserve elevation to generic status. The genus is currently considered to contain three subgenera: Euxerus (X. erythropus), Geosciurus (X. inauris, X. princeps) and Xerus (X. rutilus) (Ellerman 1940, Moore 1959, Amtmann 1975, Thorington & Hoffman 2005). The species are distinguished on the presence or absence of a lateral side-stripe, the number of dark bands on tail hairs, presence or absence of an extra premolar (P3) in the skull, and the colour of the front incisors. Jane M. Waterman

Xerus erythropus STRIPED GROUND SQUIRREL (AFRICAN GROUND SQUIRREL, WEST AFRICAN GROUND SQUIRREL, GEOFFREY’S GROUND SQUIRREL) Fr. Écureuil fouisseur du Sahel; Ger. Gestreiftes Erdhörnchen Xerus erythropus (E. Geoffroy, 1803). In: Cat. Mamm. Mus. Hist. Nat., Paris, p. 178. Unknown, but neotype given as Senegal.

Taxonomy Originally described in the genus Sciurus and later placed in either the genus Euxerus or Xerus. Euxerus is now considered to be a subgenus by most authors. The species name was first cited as erythopus [sic] but the emended spelling is now accepted (see Grubb [2004] for a discussion of the specific name). For general account, see Herron &Waterman (2004). Synonyms: agadius, albovittatus, chadensis, fulvior, lacustris, lessoni, leucoumbrinus, limitaneus, maestus, marabutus, microdon, prestigiator. Subspecies: five. Chromosome number: 2n = 38 (Dobigny et al. 2002b). Description Large terrestrial squirrel with coarse pelage. Hairs sparse, especially on ventral surface; black skin underlying hairs easily visible. Dorsal pelage dark brown to pale cinnamon and sandy-yellow; hairs sandy coloured at base, sometimes with black or brown tip. Most geographical variation in colour is due to colour of hair tip (see below). Ventral pelage white or pale. Conspicuous white side-stripe on each flank. Head with narrow white stripes above and below the eyes. Ears small and rounded. Limbs and feet paler than dorsal pelage. Tail moderately long (ca. 80% of HB), dorsoventrally flattened, hairs long, with alternating black and white bands; hairs project vertically from tail to form large brush. Males slightly larger than females. Skull: cheekteeth 5/4 (anterior upper premolar (P3) may be shed in some

adults so cheekteeth 4/4); posterior end of bony palate considerably posterior to M3; masseteric tubercle prominent; outer surface of incisor teeth orange. Nipples: 0 + 0 + 1 + 2 = 6.

Xerus erythropus.

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Atlantoxerus getulus. Smaller (HB: 165–230 mm; GLS: 38.4–50.0 mm); pelage soft and less coarse; side-stripe on each flank and single mid-dorsal stripe; usually five upper cheekteeth; sympatric in W Morocco, but more widespread in N Africa, including montane habitats, than X. erythropus. Distribution Endemic to Africa. Widespread across sub-Saharan Africa in Sahel, Sudan and Guinea Savanna BZs, and Northern and Eastern Rainforest–Savanna Mosaics, from Senegal and Mauritania to E Sudan. Extends southwards into margins of Rainforest BZ where there is suitable habitat (see below). Small isolated population in W Morocco (near Agadir). Recorded from SW Morocco, S Mauritania, Senegal, Gambia, Guinea-Bissau, Guinea, Sierra Leone, Côte d’Ivoire, S Mali, Burkina, Ghana, Togo, Benin, SE Niger, Nigeria, N Cameroon, NE Congo, S Chad, Central African Republic, Sudan, NE DR Congo, Uganda, Rwanda, Eritrea,W Ethiopia (except montane regions) and W Kenya. Unconfirmed record in N Tanzania.

Xerus erythropus

Habitat Open habitats in semi-desert, grassland savannas, woodland savannas and coastal scrub, and also in grassy clearings in the Rainforest– Savanna mosaic and the northern edge of the Rainforest BZ. Lives in rocky areas, tree root crevices, termite mounds and self-dug burrows. Abundance

Geographic Variation Many forms of X. erythropus have been recognized, all based on differences in pelage colour. Amtmann (1975) and Thorington & Hoffman (2005) recognized six subspecies, and Rosevear (1969) recognized five subspecies. Populations in high to moderate rainfall areas are dark rufous-brown or dark chocolate-brown, those in arid regions are pale oatmeal or sandy; all shades of pelage colour between these extremes are recorded from areas of intermediate rainfall. Six subspecies, of uncertain validity, are listed here. X. e. chadensis: Lake Chad (N Nigeria, W Chad, N Cameroon) and W Sudan. Dorsal pelage buff to creamy-buff. Paler than other subspecies. X. e. erythropus: mainly Sahel Savanna BZ, from Senegal to S Mauritania to NE Nigeria (and perhaps further east). Also SW Morocco. Dorsal pelage pale reddish-brown to sandy-yellow; some hairs on back have black tips. X. e. lacustris: NE DR Congo and Uganda.The darkest brown subspecies. Rosevear (1969) suggests that this subspecies is the same as X. e. maestus (see Taxonomy). X. e. leucoumbrinus: Sudan Savanna BZ from Senegal to Ethiopia, including NW Kenya. Dorsal pelage dark red-brown. X. e. limitaneus:W Sudan. Dorsal pelage similar to leucoumbrinus, darker than chadensis. Said to be larger than leucoumbrinus and chadensis. X. e. microdon: Rosevear (1969) suggests this subspecies ranges from Senegal to Kenya but Amtmann (1975) suggests it only occurs in SW Kenya. Dorsal pelage dark brown. Rosevear (1969) questions the validity of subspecific status for this form, suggesting that it is the same as X. e. leucombrinus.

Common. Densities not known.

Adaptations Diurnal and terrestrial. Burrows consist of a central chamber and about three entrance tunnels, and are simpler than those of X. inauris. During the heat of the day, Striped Ground Squirrels move into areas of shade or shelter in their burrows. They also press the ventral surface of the body on shaded or cool sand as a means of losing body heat (Ewer 1966, Linn & Key 1996). Foraging and Food Herbivorous. Feeds mainly on leaves, flowers, roots, seeds, soft fruits, and nuts. The diet may also include insects (especially termites) and meat (Ewer 1966). Food in much of the habitat is patchy and unpredictable, and hence individuals tend to forage alone. Striped Ground Squirrels are scatter hoarders, burying food well away from the burrow entrances (Ewer 1966, 1968). In some regions they are regarded as a pest because of their consumption of maize (Key 1990).

Social and Reproductive Behaviour Asocial. Adults usually live singly or in small family groups. Interactions between individuals are usually brief – two individuals approach and touch nasal areas and then move on. Overt aggression is rare. Often during a meeting, one individual is submissive and the other dominant, but detailed information on these interactions is lacking (Linn & Key 1996). In an area of periodic drought, when resources were very patchy, home-range size of three "" averaged 12.4 ha. In an area of higher rainfall, home-range of a single " was 3.34 ha and for four !! was 9.43 ha. Individuals forage widely within the home-range and often do not return to the same sleeping burrow each night. Home-ranges overlap considerably between conspecifics and there is no evidence of territoriality. Very tolerant of conspecifics and even burrows are Similar Species not defended from visitations by a succession of other individuals X. rutilus. Smaller body size: side-stripe absent: lacks the extra premolar (Linn & Key 1996). characteristic of X. erythropus (premolar is often shed in adult X. Nothing is known about mating behaviour, although mating chases erythropus); marginally sympatric in parts of eastern Africa. have been noted (Linn & Key 1996). Young animals participate in 95

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social play but this decreases as they reach maturity. Vocalizations are rare. Juveniles emit distress calls and individuals will give alarm calls while running toward the burrow if they are threatened. Scent marking by rubbing cheek glands on objects has been described as more common in "" than !! (Ewer 1966, 1968). This species appears to be displacing X. rutilus in parts of Kenya, possibly because it is more aggressive and larger (Key 1990); however, O’Shea (1976) reported that the two species share burrows. Reproduction and Population Structure Almost nothing is known about reproduction and growth. Probably breeds prior to one year of age (Ewer 1966). Litter-size: usually 4 (2–5). Eyes open ca. Day 26. First moult ca. Day 55. Eat solid food ca. Week 5 (Ewer 1966). May live up to six years in captivity (Kingdon 1974). Predators, Parasites and Diseases Major predators include hawks, eagles, snakes (especially Puff Adder Bitis arietans), servals, wild cats and jackals (Kingdon 1974). Can harbour a number of diseases, including Streptobacillus moniliformis, which causes rat-bite fever (Linn & Key 1996), Trypanosoma (Herpetosoma) xeri (Marinkelle & Abdalla 1978) and many species of ticks.

Measurements Xerus erythropus HB (!!): 259 (160–390) mm, n = 45 HB (""): 250 (193–430) mm, n = 48 T (!!): 200 (124–269) mm, n = 47 T (""): 197 (138–226) mm, n = 48 HF (!!): 67 (51–75) mm, n = 48 HF (""): 66 (47–75) mm, n = 48 E (!!): 16 (7–24) mm, n = 46 E (""): 17 (7–19) mm, n = 45 WT (!!): 513 (346–750) g, n = 26 WT (""): 429 (226–565) g, n = 30 GLS: 60.8 (57.4–65.7) mm, n = 127 GWS: 32.4 (30.7–34.5) mm, n = 17 P3–M3: 12.1 (10.9–13.5) mm, n = 17 Body measurements and weight: throughout geographic range (USNM) Skull measurements: throughout geographic range (BMNH). In those specimens (7 of 17) that do not have the very small P3, the measurement is P4–M3 Key References Rosevear 1969.

Ewer 1966; Kingdon 1974; Linn & Key 1996;

Conservation IUCN Category: Least Concern. Unlikely to be threatened because of widespread distribution and commonness.

Jane M. Waterman

Xerus inauris CAPE GROUND SQUIRREL (SOUTH AFRICAN GROUND SQUIRREL) Fr. Écureuil foisseur du Cap; Ger. Kaperdhörnchen Xerus inauris (Zimmermann, 1780). Geogr. Gesch. Mensch. Vierf. Thiere 2: 344. 160 km north of Cape of Good Hope, South Africa.

Taxonomy Originally described in the genus Sciurus. Previously allocated to the genera Sciurus, Myoxus, Geosciurus and Xerus. Subgenus Geosciurus. Xerus inauris is considered to be a separate species from X. princeps based on minor chromosomal differences and morphological studies (Robinson et al. 1986, Herzig-Straschil et al. 1991). For a general account, see Skurski &Waterman (2005). Synonyms: africanus, albovittatus, capensis, dschinshicus, gininianus, levaillantii, namaquensis, setosus. Subspecies: none. Chromosome number: 2n = 38 (Robinson et al. 1986). Description Large terrestrial squirrel with white side-stripes and very small ears (hence the name inauris). Hairs sparse and short; dark skin underlying hairs easily visible. Dorsal pelage pale cinnamonbrown; hairs sandy with small white tip; some longer hairs with black tip. Conspicuous white side-stripe from shoulders to hips. Ventral pelage off-white or pale yellowish-white. Eyes large, with dull white stripe above and below each eye, extending anteriorly to nostrils. Testes on !! large, approximately 19.8% of HB. Forelimbs short, sandy above, off-white below; forefoot with four digits, each with long dark claw. Hindlimbs sandy; hindfoot off-white above, naked below; five digits, each with long dark claw. Tail moderately long (ca. 85% of HB), dorsoventrally flattened; hairs long (ca. 50–60 mm), banded, each hair white with two black bands (short blackish-brown band near base, long black band near tip) and long white tip. Skull: cheekteeth 5/4; posterior end of bony palate considerably posterior

to M3; masseteric tubercle prominent; outer surface of incisor teeth white. Nipples: 0 + 0 + 1 + 1 = 4. Geographic Variation

None recorded.

Similar Species X. princeps. Incisor teeth orange; tail hairs white with three black bands; tail comparatively longer; marginally sympatric in W Namibia. Distribution Endemic to Africa.Widespread in semi-arid regions of southern Africa. South-West Arid BZ (Kalahari Desert, Namib Desert and Karoo) and western part of Highveld BZ. Recorded from C South Africa, Botswana, W Lesotho and Namibia. Not recorded from coastal Namibia. Habitat Open semi-arid regions where mean annual rainfall is 125–500 mm. Preferred habitat is hard ground with some scrub cover along the edges of pans, river beds and open sandy veld; also recorded from short grasslands, overgrazed areas and cultivated fields. In Namibia, areas with short annual grass (Schmidtia kalahariensis) are preferred, and areas with longer perennial grasses are avoided. Abundance Common. In the Kalahari region of SE Namibia, density is 3–4/ha, with approximately 1.2 adult ""/ha, 1.3 adult !!/ha and 1.3 subadults/ha (Waterman 1995). In drought years,

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Xerus inauris.

density declines significantly (Waterman & Fenton 2000). Probably live in higher densities in areas with higher rainfall, but no comparable data available. Adaptations Diurnal and terrestrial. Cape Ground Squirrels dig their own complex burrows, which may be 60–80 cm deep, with 60 or more burrow openings within a cluster of burrows and more than one nest chamber in a burrow system (Herzig-Straschil 1978). When digging, the nostrils can be closed. During the heat of the day, the tail is orientated towards the direction of the sun while held up over the body, providing shade on the back and head. Cape Ground Squirrels also use their burrows as thermal retreats (Bennett et al. 1984). Their kidneys have a thick medulla, and urine concentrations are high. Hence squirrels can survive largely on metabolic water, without the need for free drinking water (Marsh et al. 1978). Foraging and Food Primarily herbivorous, also omnivorous. The diet consists of leaves, sheaths and roots of grasses, as well as seeds, fruit and insects. Preferred foods in central South Africa include the grasses Cynodon dactylon and Enneapogon brachystachyus, while in SC Namibia and the Kalahari National Park their diet includes the grasses Schmidtia kalahariensis and Eragrostis lehmanniana (Herzig-Straschil 1978, Knight 1991, Waterman 1996). They will also feed on the berries of Boscia albitrunca, Grewia flava and the fruits of Citrullus lanatus. Insects consumed include termites, beetles, locusts and caterpillars. No evidence of food hoarding has been found when the contents of burrows were examined. They have been blamed for damage to maize crops in southern Africa, but Zumpt (1970) attributed the damage by squirrels as only 0.2–0.4%. Social and Reproductive Behaviour Social. Females live in matrilineal groups of 1–4 adult "" and up to nine subadults of either sex (Herzig-Straschil 1978, Waterman 1995). Colonies can be as large as 30 individuals (Smithers 1971) and several groups of ""

Xerus inauris

may inhabit a colony or cluster of burrows (Herzig-Straschil 1978). In Namibia, only a single group of "" inhabits a burrow area. Some studies have indicated that a dominance hierarchy exists amongst "" (Herzig-Straschil 1978, Knight 1991), but no such hierarchy was observed in a study in Namibia (Waterman 1995). Adult !! form permanent, non-aggressive groups of up to 19 individuals that live independently of groups of "" (Waterman 1995, 1997). On a daily basis, !! form temporary subgroups (4–5 individuals), the size and individual composition of which are constantly changing. While in these subgroups, !! forage, sleep and roam their homerange together. Interactions between members of the same group of "" are primarily amicable, with frequent approaching, greeting and allogrooming. Agonistic interactions within a group are rare (Herzig-Straschil 1978, Waterman 1996). Juveniles interact as amicably with other members of the group as they do with their mothers. Interactions between different groups of "" are rare and usually agonistic. Agonistic interactions amongst !! in an all-male groups are also rare, and injury has never been observed. There is a linear dominance hierarchy in the group, which is correlated with age (Waterman 1995, 1997) and determined by non-aggressive displacements, rather than fighting. Interactions between different groups of !! have not be recorded. Females in the same social group share sleeping burrows and homeranges. Home-range area during a normal rainfall year was 4 ha; however during drought, ranges more than doubled in area (Waterman 1995, Waterman & Fenton 2000). There is some overlap of home-ranges of adjacent female social groups but no overlap of core areas between social groups. Females will defend core areas from neighbouring groups of "" (Herzig-Straschil 1978). Homeranges of male groups encompass a number of female groups but are not defended against other male groups, and new !! are accepted into the band without aggression (Waterman 1995). Males forage and roam their home-range together in smaller subgroups, sleeping 97

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together in vacant burrow clusters away from groups of "", and may join and leave a subgroup on a daily basis, resulting in continuous fission and fusion of subgroups (Waterman 1995, 1997). Average range size of radio-collared !! was 12.5 ha (Waterman 1995). Herzig-Straschil (1978) found smaller home-ranges for both !! and "" in a region with high rainfall. Herzig-Straschil (1978) described single !! associating with groups of "" for a few weeks during receptivity. Observations in Namibia and central South Africa suggest members of male subgroups travel together from one female group to another. Male subgroups sleep or stay in an occupied burrow area if a " is coming into oestrus, otherwise they sleep in vacant burrow areas (Waterman 1995, 1997, unpubl.). The majority of !! from the male group will attempt to mate with the " on her day of oestrus. Oestrus and parturition were asynchronous within and between groups of "". The operational sex ratio on a day of oestrus (oestrus lasts about 3 h) averaged ten !! to a ". Mating occurs both above and below ground, and the most dominant ! usually has preferential access to the oestrous ". Both !! and "" mate promiscuously. The large size of the testes of !! suggests that sperm competition could be an important element of their mating system (Waterman 1998). All members of the group (mother, other adult "" and subadult "" and subadult !!) provide care of juveniles through allogrooming, play and possibly predator detection and deterrence (Waterman 1995, unpubl. data). Knight (1991) reported seeing communal nursing. There is no male parental care. Cape Ground Squirrels have a number of vocalizations, including a high pitched alarm call that is given when the animal detects a potential threat. They will also respond to the alarm calls of other species, particularly Crowned Plovers and White-browed Sparrowweavers. Other vocalizations include an aggressive growl (used in encounters with conspecifics), a scream (used when released from a trap), a play call (used by young animals), nest-chirping (used in infants) and a protest squeak (Herzig-Straschil 1978). Adult !! and "" use secretions from the anal-genital region and the circumoral area for marking. Secretions are deposited by rubbing the body on the sand or by rubbing the snout on an object (Straschil 1975). Marking is most common on emergence from the burrow in the morning. Cape Ground Squirrels frequently share their burrow systems with Yellow Mongooses Cynictis penicillata and Suricates Suricata suricatta. The three species usually ignore each other and rarely interact (J. Waterman unpubl.). Reproduction and Population Structure Reproduction occurs throughout the year, with peaks of mating in the dry winter months. Litter loss is high, with some 70% of all oestrus cycles failing to produce successful litters (Waterman 1996). Females can have 1–4 litters per year. Gestation: 7 weeks. Litter-size: 1–4. Body weight at 7 days: 23.5 g (Herzig-Straschil 1978). Mothers lactate for

7.5 weeks. Sex ratio of litters at emergence is 1 : 1. Sexual maturity: 8 months (!), 10 months ("). Female maturity is delayed within larger groups of "" (Waterman 2002). Dispersal is male-biased. Males disperse at 8–10 months of age, whereas "" usually remain in their natal group. Adult sex ratio is 1 : 1 and the annual survival of !! is slightly higher than that of "" (78% vs. 70%). In captivity, individuals may live up to 13 years but the life-span in the wild is most likely to be 4–5 years. Survival of young to six months of age is negatively influenced by the number in the social group (Waterman 2002). During drought, the social structure remains intact; however, densities drop significantly and all reproductive activity ceases (Waterman & Fenton 2000). Predators, Parasites and Diseases Major predators include hawks, eagles, snakes, wild cats and jackals. Individuals of a group may mob potential predators such as Cape Cobras Naja nivea, Puff Adders Bitis arietans and Monitor Lizards Varanus exanthematicus (Waterman 1996, unpubl.). Ectoparasites include many species of fleas, ticks and mites (De Graaff 1981). Endoparasites include various helminth worms. Some of these parasites are vectors of human diseases such as plague, enterobiasis, biliary fever, East Coast fever and tick-bite fever (De Graaff 1981 and references therein). Conservation

IUCN Category: Least Concern.

Measurements Xerus inauris HB (!!): 246 (195–293) mm, n = 88 HB (""): 239 (181–300) mm, n = 134 T (!!): 209 (181–282) mm, n = 88 T (""): 206 (160–255) mm, n = 134 HF (!!): 69 (63–75) mm, n = 72 HF (""): 67 (45–74) mm, n = 114 E (!!): 12 (9–14) mm, n = 28 E (""): 11 (12–14) mm, n = 49 WT (!!): 591 (312–822) g, n = 71 WT (""): 565 (367–907) g, n = 84 GLS (!!): 56.7 (49.4–60.8) mm, n = 66 GLS (""): 55.3 (43.9–59.5) mm, n = 108 GWS (!!): 34.0 (29–38.3) mm, n = 66 GWS (""): 33.8 (26.5–37) mm, n = 106 P3–M3: 11.4 (10.0–12.6) mm, n = 157 Throughout geographic range (USNM, NMN, Waterman 1996, Herzig-Straschil et al. 1991) Key References Skurski & Waterman 2005; Herzig-Straschil 1978; Smithers 1983; Waterman 1995, 1996, 2002. Jane M. Waterman

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Xerus princeps

Xerus princeps DAMARA GROUND SQUIRREL (MOUNTAIN GROUND SQUIRREL, KAOKOVELD GROUND SQUIRREL) Fr. Écureuil fouisseur du Damara; Ger. Damara-Erdhörnchen Xerus princeps (Thomas, 1929). Proc. Zool. Soc., Lond. 1929: 106. Otjitundua, central Kaokoveld, N Namibia.

Xerus princeps.

Taxonomy Originally described in the genus Geosciurus. Minor chromosomal and morphological differences of the skull support Xerus princeps as a separate species from X. inauris (Robinson et al. 1986, Herzig-Straschil et al. 1991). For general account, seeWaterman & Herron (2004). Synonyms: none. Chromosome number: 2n = 38 (Robinson et al. 1986). Description Large terrestrial squirrel similar in appearance to X. inauris. Hairs sparse, coarse and short; dark skin underlying hairs easily visible. Dorsal pelage cinnamon-brown flecked with white; hairs sandy or dark brown at base, white at tip, which gives the species a more ‘grizzled’ appearance than X. inauris. Conspicuous white side-stripe from shoulders to hips.Ventral pelage white. Eyes large, with dull white stripe above and below each eye extending anteriorly to nostrils. Ears small. Tail long (ca. 99% of HB), dorsoventrally flattened; hairs long (ca. 70 mm), banded, each hair white with three black bands (two short blackish-brown bands near base, long black band near tip) and long white tip. Skull: cheekteeth 5/4; posterior end of bony palate considerably posterior to M3; masseteric tubercle prominent; outer surface of incisor teeth white or pale orange. Nipples: 0 + 0 + 1 + 1 = 4. Geographic Variation None recorded. Similar Species X. inauris. Incisor teeth white; tail hairs white with two black bands; tail comparatively shorter; distribution more extensive; very social.

Xerus princeps

Abundance Herzig-Straschil & Herzig (1989) suggest that densities are very low. No detailed information. Adaptations Diurnal and mainly terrestrial. Burrow systems are simple when compared to X. inauris, with only 2–5 openings and a single nest chamber (Herzig-Straschil & Herzig 1989). Burrow entrances are usually found under piles of rocks or boulders. Damara Ground Squirrels use the tail as a thermal shade (as described for X. inauris) and they are better adapted to survive high temperatures than X. inauris (Haim et al. 1987). The thermoneutral zone of X. princeps is 32–35 °C, and hyperthermy occurs at 35 °C (Haim et al. 1987). Salivation was observed at 38 °C. Thermal conductance was high (0.084 ± 0.005 ml O2/g/h/ °C), which results in high heat dissipation and water conservation. In comparison with X. inauris, the faeces of X. princeps are significantly drier (14.21 ± 4.2% moisture content) (Haim et al. 1987).

Distribution Endemic to Africa. South-West Arid BZ, mainly in the Namib Desert. Recorded from only the narrow band of the western escarpment that runs from SW Namibia north to SW Angola.

Foraging and Food Mainly herbivorous. Feeds on the base of grass stems and roots. In mopane savanna, animals have been observed climbing mopane trees (Colophospermum mopane) to feed on plant lice (Copaifera mopane) and on mopane leaves (Herzig-Straschil & Herzig 1989).

Habitat Rocky, hilly ground in arid areas where mean annual rainfall is ca. 125–250 mm (Herzig-Straschil & Herzig 1989). Also recorded on open plains, but prefers habitats with gravel and rocks in areas with single trees or sparse bush cover.

Social and Reproductive Behaviour Solitary, or small family groups (mother and young) of 2–4 individuals (Haim et al. 1987, Herzig-Straschil & Herzig 1989). Adult !! are associated with some groups but otherwise live solitarily. No allogrooming, playing or other 99

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Family SCIURIDAE

cohesive behaviours have been documented during the short period of time they have been observed (Herzig-Straschil & Herzig 1989). Reproduction and Population Structure Little information. Breeding most likely occurs during the winter months. Gestation: ca. 7 weeks. Litter-size: 1–3 (Herzig-Straschil & Herzig 1989). Predators, Parasites and Diseases Nothing is known about potential predators. The only recorded ectoparasite is a flea (Ctnochephalides connatus) (De Graaff 1981). Conservation IUCN Category: Least Concern. The limited geographical range and comparative rarity of the species may be cause for concern in the future (Waterman & Herron 2004). Measurements Xerus princeps HB (!!): 244 (230–280) mm, n = 15 HB (""): 239 (235–290) mm, n = 13

T (!!): 241 (220–260) mm, n = 15 T (""): 239 (205–282) mm, n = 13 HF (!!): 70 (65–75) mm, n = 15 HF (""): 71 (68–73) mm, n = 13 E (!!): 13 (12.5–14) mm, n = 15 E (""): 14 (13–15) mm, n = 13 WT (!!): 572, 716 g, n = 2 WT (""): 610, 700 g, n = 2 GLS: 58.2 (54.6–61.4) mm, n = 22 GWS: 35.4 (33.4–37.1) mm, n = 22 P3–M3: 11.2 (10.3–11.9) mm, n = 22 Throughout geographic range Body measurements and weight: NMN, Skinner & Smithers 1990 Skull measurements: Herzig-Straschil et al. 1991 Key References Herzig-Straschil & Herzig 1989; Skinner & Smithers 1990; Waterman & Herron 2004. Jane M. Waterman

Xerus rutilus UNSTRIPED GROUND SQUIRREL (PALLID GROUND SQUIRREL) Fr. Écureuil fouisseur unicolore; Ger. Streifenloses Erdhörnchen Xerus rutilus (Cretzschmar, 1828). In: Rüppell’s Atlas Reise Nordl. Afr., Zool., Säugeth. p. 59. Eastern slope of Abyssinia, Ethiopia (probably Massawa, Eritrea – see Mertens 1925: 26).

Geographic Variation The validity of the eight subspecies is dubious (Amtmann 1975). Differences in pelage colouration perhaps only reflect differences in soil colouration rather than taxonomic differences. Populations in drier areas tend to be paler. Xerus rutilus.

Taxonomy Originally described in the genus Sciurus. Polytypic with up to eight subspecies (Amtmann 1975, O’Shea 1991). For a general account, see O’Shea (1991). Synonyms: abessinicus, brachyotis, dabagala, dorsalis, fuscus, intensus, massaicus, rufifrons, saturatus, stephanicus. Subspecies: eight (validity uncertain). Chromosome number: 2n = 38 (Nadler & Hoffmann 1974). Description Medium-sized terrestrial squirrel with bristly coarse pelage and without side-stripe. Dorsal pelage pale tan to rich reddish-brown with variable amounts of pale and dark speckling; hairs short (ca. 5–6 mm), dark reddish-brown at base, with buff or black tip. Individuals in drier areas tend to be pale. No side-stripe (cf. all other species in genus). Ventral pelage whitish, and paler than dorsal pelage; hairs sparse. Head with conspicuous white or off-white eyering. Ear small. Upper surface of fore- and hindfeet white (reddishbrown in some individuals).Tail of moderate length (ca. 86% of HB); hairs long (ca. 40 mm), each hair banded, off-white at base, blackishbrown in middle, with reddish-brown (e.g. Tanzania) or white (e.g. Somalia) at tip. Skull: cheekteeth 4/4; posterior end of bony palate considerably posterior to M3; masseteric tubercle prominent; outer surface of incisor teeth orange. Nipples: 0 + 0 +1+ 1 = 4.

X. r. rutilus: N Ethiopia and NE Sudan. Dorsal pelage reddish brown. X. r. dabagala: Somalia and southern Ethiopia. Dorsal pelage dull to bright rufous or tawny. X. r. intensus: Gerlogubi Wells; red sandy country of C Somalia. Hairs of crown and dorsal region tipped with white. Tail rufous at base above and white below. X. r. stephanicus: L. Stephanie, Ethiopia. Dorsal pelage grizzled yellowish-grey. X. r. rufifrons: Guaso Nyiro, N Kenya. Rufous marking on muzzle extends onto dorsal region of head. X. r. saturatus: SE Kenya and NE Tanzania. Dorsal pelage with dark reddish tinge. X. r. dorsalis: L. Baringo, Kenya. Dorsal pelage dark olive, speckled with yellow and white. X. r. massaicus: Masai Reserve, Kenya. Rufous tinge on forehead, yellow on throat and underparts; skull appears to be larger than in other subspecies. Similar Species X. erythropus. Larger body size; side-stripe present; extra premolar (P3) present in some individuals; sympatric in some parts of geographic range. Distribution Endemic to Africa. Somalia–Masai Bushland BZ. Recorded in NE Sudan, E and S Ethiopia, Djibouti, Eritrea, Somalia,

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includes lunges by the dominant individual, but biting and fighting have not been observed (O’Shea 1976). A number of vocalizations have been noted, including churring vocalizations used by "" and juveniles when approached by !!, and a scolding chatter given by dominant individuals to subordinates (O’Shea 1976). Home-ranges of !! (7 ha) are larger than those of "" (1.4 ha); home-ranges overlap considerably within and between the sexes. There is no evidence of territoriality but individuals with overlapping home-ranges appear to form a linear dominance hierarchy. Males court "" by pilo-erecting the hairs of the tail, arching the tail over the head and approaching the "". Females usually move away, making quiet vocalizations, flicking the tail rapidly and slightly dragging the perineal region of the body on the ground. When ! approaches ", she takes a prostrate posture and roles onto her side allowing ! to sniff her anogenital region. The ! then mounts the prostrate " (O’Shea 1976). Young are born in burrows, which are usually located at the edge of a female’s home-range. Three to four weeks after the juveniles emerge, " returns to the centre of her home-range, away from the juveniles (O’Shea 1976). Xerus rutilus

Kenya, NE Uganda and NE Tanzania. The geographic range appears to be contracting in some areas of Kenya, possibly as a result of competition with the larger more aggressive X. erythropus (Key 1990). Habitat Semi-arid dense thornbush interspersed with open grassland (O’Shea 1991). In South Turkana, Kenya, lives in alluvial flats and gullies and in thickets of Salvadora persica (Coe 1972). Abundance Common. In South Turkana, Kenya, estimated densities may reach 848 individuals/km2 or approximately 8.5 individuals/ha (Coe 1972). Adaptations Diurnal and terrestrial. Burrows are often located at the base of bush stems or in termite mounds, and usually have 2–6 entrances. Unstriped Ground Squirrels may also shelter in the burrows of other mammals (including X. erythropus [O’Shea 1976]). In order to prevent overheating, the ventral surface of the body is pressed onto shaded or cool sand (Coe 1972). Foraging and Food Herbivorous and omnivorous. Feeds on herbaceous plants including seeds, leaves, flowers, soft fruits and the large fruits of baobab trees, as well as insects (Coe 1972, O’Shea 1991). Greater than 50% of stomach contents were found to be small dry seeds and leaves (Coe 1972). Unstriped Ground Squirrels are scatter-hoarders, storing food in many caches (O’Shea 1976). Social and Reproductive Behaviour Primarily a solitary species, although small family groups (mother and young), and !! living with one or two "" are also common (O’Shea 1976). Numbers of individuals sharing a burrow system can range from 1 to 6 (Coe 1972; O’Shea 1976). Individuals of different sexes tend to avoid each other; however, at feeding sites !! appeared to be dominant over "", as indicated by chases and displacements. Threat behaviour

Reproduction and Population Structure Almost nothing is known about reproduction and population dynamics. Breeding appears to occur throughout the year. Litter-size: one or two (Kingdon 1974, O’Shea 1991). All young eventually disperse from their natal area (O’Shea 1976). Sex ratio is probably 1 : 1 (Coe 1972). In captivity, a single ! has been documented to live for over six years (O’Shea 1991). Predators, Parasites and Diseases No information on potential predators. Ectoparasites include a tick that is specific to the species (Haemaphysalis calarata), and a flea (Synosternus somalicus). One cestode (Catenotaenia geosciuri) also recorded (O’Shea 1991). Conservation IUCN Category: Least Concern. The species does not appear to be threatened at the present time. Measurements Xerus rutilus HB (!!): 212 (161–230) mm, n = 6 HB (""): 215 (168–237) mm, n = 11 T (!!): 185 (153–195) mm, n = 6 T (""): 185 (145–205) mm, n = 11 HF (!!): 54 (50–60) mm, n = 6 HF (""): 54 (51–59) mm, n = 11 E (!!): 14 (12–16) mm, n = 3 E (""): 14 (12–17) mm, n = 4 WT (!!): 325 (135–440) g, n = 3 WT (""): 313 (155–420) g, n = 3 GLS: 50.7 (47.1–53.7) mm, n = 17 GWS: 29.4 (27.0–32.3) mm, n = 17 P3–M3: 9.3 (8.7–10.0) mm, n = 17 Body measurements and weight: throughout geographic range (USNM) Skull measurements: Somaliland (BMNH) Key References

O’Shea 1976, 1991. Jane M. Waterman 101

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Family GLIRIDAE

Family GLIRIDAE DORMICE

Gliridae Muirhead, 1819. Glirini Muirhead, 1819: 433. Mazology [sic]. pp. 393–480, pls. 353–358, in Edinburgh Encyclopedia, Vol. 13 (D. Brewster, ed.). Eliomys (2 species) Graphiurus (14 species)

Garden Dormice Dormice

p. 104 p. 109

The nine genera and 28 living species of Gliridae (commonly known as dormice) are currently arranged in three subfamilies and occur in tropical and temperate forests, savannas, steppes and deserts in Africa and the Palaearctic region (Holden 2005). The name Myoxidae is the original name for this family, but the Commission on Zoological Nomenclature (see Holden 2005) has ruled that the name is invalid, and that Gliridae is the valid family name for dormice. The family is divided into three subfamilies: Glirinae, Leithiinae and Graphiurinae (Wahlert et al. 1993, Montgelard et al. 2003, Holden 2005). The Glirinae contain Glis, indigenous to Europe, N Turkey, Caucasus, N Iran and SW Turkestan; and Glirulus, endemic to the Japanese islands of Honshu, Dogo, Shikoku and Kyushu. The Leithiinae contains the genera Chaetocauda, Dryomys, Eliomys, Muscardinus, Myomimus and Selevinia, and is distributed throughout Europe, the Middle East, C and W Asia; the geographic range of Eliomys also includes North Africa. The subfamily Graphiurinae contains only Graphiurus, and is endemic to sub-Saharan Africa. The family is represented in Africa by Eliomys (2 spp.) and Graphiurus (14 spp.). Living glirids range in body size from small to moderately large (HB 60–190 mm) although the maximum HB for nine of the 16 African glirid species is ca. 100 mm. The tail (40–165 mm) is usually moderately long relative to body size, and well-covered with hair throughout its length (in a similar way to the tail of squirrels). Most species have thick, soft dorsal pelage, are predominantly arboreal and resemble small squirrels in body form. Two exceptions are

the non-African Myomimus and Selevinia, which are terrestrial and have thinly haired tails. Colour of dorsal pelage ranges from pale grey to dark brown. Many species have a narrow or broad dark stripe (eye-mask) extending from the muzzle to the ears; some species exhibit only a thin dark line (eye-ring) encircling the eyes. Colour of ventral pelage is usually grey or white. In several species, especially in Graphiurus and the non-African Glis, the chin, chest and sometimes forelimbs are discoloured in many individuals; this discolouration is often reddish-brown, but is sometimes pale yellow or yellowish-green (Channing 1984, Nowak 1999, M. E. Holden unpubl.). This discolouration may result from staining that occurs when the dormice ingest insects (such as earwigs) and certain fruits (Chapin [in Hatt 1940a], Rosevear 1969, Nowak 1999), or it may be genetically induced (Channing 1984, B. Kryštufek & R. M. Baxter pers. comm.). All species have rounded ears, large eyes and short fore- and hindlimbs. The forefeet have four digits, the hindfeet have five and the palmar and plantar surfaces are naked. Claws are short and sharp, and those of arboreal species are recurved. The skull is smooth, usually without postorbital processes or supraorbital and temporal ridging. The zygomatic arches are prominent, the anterior palatal foramina are generally short, their posterior margins usually anterior to or even with the front margins of the ventral maxillary zygomatic processes, and the bony palate is long. Auditory bullae are usually large, often appear inflated relative to skull size and are divided internally by bony septae.The infraorbital foramen is moderately tall, ovate and is penetrated by a portion of the anterior medial masseter. The zygomatic plate, from which originates part of the anterior lateral masseter, tilts upwards in all genera (modified or convergent

Table 14. Species in the family Gliridae. Arranged in order of increasing head and body length. (n. d. = no data.)

a

Species

HB mean (range) (mm)

T mean (range) (mm)

GLS mean (range) (mm)

Upper toothrow length mean (range) (mm)a

Anterior palatal foramina length mean (range) (mm)

Graphiurus johnstoni Graphiurus kelleni Graphiurus lorraineus Graphiurus murinus

74.3 (69–84) 82.4 (75–92) 83.0 (72–93) 91.5 (81–103)

68.5 (65–75.5) 68.3 (54–81) 65.7 (54–74) 76.6 (69–85)

23.6 (23.3–23.9) 24.0 (23.1–24.5) 24.5 (22.7–26.1) 26.4 (25.2–28.8)

3.4 (3.3–3.5) 2.9 (2.8–3.0) 3.1 (2.8–3.4) 3.1 (3.0–3.3)

2.7 (2.6–2.9) 2.9 (2.6–3.2) 2.6 (2.0–3.0) 3.1 (2.7–3.7)

Graphiurus crassicaudatus

92.6 (83–98)

59.4 (55–70)

26.6 (25.2–27.8)

3.8 (3.4–4.2)

2.5 (2.3–2.8)

Graphiurus christyi Graphiurus microtis Graphiurus angolensis Graphiurus surdus Graphiurus platyops Graphiurus rupicola Eliomys munbyanus Eliomys melanurus Graphiurus ocularis Graphiurus nagtglasii Graphiurus monardi

97.6 (86–107) 98.8 (75–115) 98.8 (79–112) 99.0 (87–110) 107.1 (95–122) 110 (105–119) 117 (100–140) 128 (111–144) 134.3 (117–145) 138.5 (120–155) 160

79.8 (73–95) 75.2 (62–86) 79.2 (70–96) 72.3 (65–82) 78.7 (65–98) 104.2 (96–118) 108 (96–118) 122 (100–136) 114.5 (103–150) 105 (65–122) 130

28.0 (26.7–29.7) 27.4 (25.5–29.1) 28.2 (26.3–30.8) 27.6 (26.5–29.4) 30.4 (28.6–32) 31.3 (30.5–32.3) 33.6 (31.7–35.6) 35.9 (34.2–37.0) 35.8 (34.2–37.5) 36.8 (34.9–39.1) 34.1 (32.5–36.6)

3.2 (3.0–3.3) 3.0 (2.9–3.4) 3.2 (2.9–3.5) 3.2 (2.9–3.5) 3.1 (2.8–3.5) 3.4 (3.3–3.7) 4.7 (n. d.) 5.3 (n. d.) 3.3 (3.0–3.5) 5.1 (4.6–5.7) 3.9 (3.6–4.3)

3.0 (2.4–3.3) 3.4 (3.0–3.8) 3.4 (3.0–3.9) 2.8 (2.5–3.2) 3.2 (2.7–4.1) 3.4 (3.1–3.6) 4.3 (4.0–4.9) 4.3 (3.8–4.8) 3.5 (3.0–4.1) 3.7 (2.9–4.3) 4.1 (3.8–4.7)

P4–M3

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myopmorphous configuration) except in Graphiurus where it is below the infraorbital foramen (modified hystricomorphous condition, except no separate infraorbital neurovascular foramen is present, which is a feature of true hystricomorphy). The zygomatic plate is narrow, and its anterior margin does not project forward beyond the dorsal maxillary zygomatic process, so no zygomatic notch is present. The mandible has laterally directed angular processes that are usually perforated. Dental formula: I 1/1, C 0/0, P 1/1, M 3/3 = 32). Incisor tips form V-shaped cutting edges (forming a V-shaped notch from anterior view) in most species of Graphiurus, but straight cutting edges in species of Eliomys and other glirids. Premolars are absent in the non-African Selevinia, and markedly reduced in Graphiurus ocularis. Cheekteeth are low-crowned (brachydont), and their occlusal patterns are variations of four transverse lophs into which cusps are subsumed, forming a range of patterns defined by basins, transverse ridges and accessory crests. Glirids do not have a caecum, and lack a noticeable morphological boundary between large and small intestines. The arterial circulation to the brain is provided by the vertebral artery (the same pattern as in squirrels) rather than the internal carotid (the primitive condition). Females of all glirid species usually have four pairs of nipples (pectoral, brachial, and two inguinal) although some variation has been documented in the non-African Glis (Kryštufek 2004). These and other morphological attributes of glirids are described, illustrated and reviewed by Ellerman (1940), Klingener (1984), Bugge (1985), Wahlert et al. (1993), Holden (1996), Nowak (1999), Rossolimo et al. (2001), Potapova (2001) and Holden (2005). Glirids are one of the oldest families of living rodents.Their fossils first appear in the early Eocene deposits (Daams & De Bruijn 1995, Daams 1999), suggesting a late Palaeocene-early Eocene origin (Hartenberger 1994, 1998). This is concordant with the most recent molecular dating estimate based upon combined markers (Huchon et al. 2002, Adkins et al. 2003, Montgelard et al. 2003). Most extant glirid genera were clearly differentiated and exhibited their greatest

species diversity by the early to middle Miocene (Hartenberger 1994, Daams & De Bruijn 1995, Daams 1999). The Graphiurinae are an exception; definite examples of Graphiurus are known only as far back as the Pliocene (Pocock 1976, Hendey 1981), although late Miocene graphiurines have been recorded from South Africa and Namibia (Denys 1990b, Mein et al. 2000b). The living Palaearctic genera are relicts of a rich adaptive radiation of up to 15 genera. Modern glirids (excluding species of Graphiurus) have been characterized as myomorphous, and several authors have included Gliridae within the suborder Myomorpha (Simpson 1945, Chaline & Mein 1979, Wahlert et al. 1993, McKenna & Bell 1997). Wahlert et al. (1993) placed them within Myomorpha based on derived cranial characters; their phylogenetic reconstruction indicated that the hystricomorphous-like Graphiurus is the most primitive extant glirid, and the myomorphy exhibited by all other extant glirids is convergent to that of true myomorphs. The terms ‘pseudomyomorphy’ (VianeyLiaud 1985, Maier et al. 2002) or ‘pseudosciuromorphy’ (Landry 1999) have been employed to distinguish the zygomasseteric muscular arrangement found in glirids from that of muroids, and Maier et al. (2002) noted that pseudomyomorphy (the convergent myomorphous musculature of glirids) is one of the derived diagnostic characters of Gliridae. The origin of glirids, and the evolutionary relationship between glirids and other rodent groups, are subjects of historical debate and controversy (Holden 2005). Glirids have alternately been placed in the rodent suborder Myomorpha (Simpson 1945, Wahlert et al. 1993 and references therein), in the infraorder Sciurida (that also contains Sciuridae; see Meng 1990, McKenna & Bell 1997) and in the suborder Sciuromorpha (in subfamily Microparamyinae; VianeyLiaud 1994, Vianey-Liaud & Jaeger 1996). There is an overwhelming lack of support for inclusion of glirids in Myomorpha based upon recent morphological and molecular research (see Holden 2005, and references therein), and they are currently placed in the suborder Sciuromorpha (Carleton & Musser 2005). Most researchers

Auditory bullae length mean (range) (mm)

White tip to tail

Geographic distribution

Notes

6.8 (6.6–7.1) 7.8 (7.3–8.2) 7.2 (6.6–7.8) 7.1 (6.7–7.7)

Yes Yes (faint in some) No No (faint in some)

Malawi (limits uncertain) West, eastern and south-central Africa West and central Africa Ethiopia to South Africa

6.7 (6.5–6.9)

Yes

Liberia to Cameroon

7.4 (6.7–7.9) 8.1 (7.3–8.6) 9.0 (8.3–9.7) 7.3 (6.9–7.7) 8.4 (7.8–8.9) 9.4 (9.2–10.2) 10.2 (9.6–10.5) 11.7 (11.2–12.2) 9.8 (9.2–10.5) 7.9 (7.4–8.4) 10.3 (9.8–11.1)

No Yes Yes No Yes Yes Yes No (faint on some) Yes No Yes

N DR Congo, SW Cameroon Chad/Sudan to South Africa C and S Angola, Zambia Cameroon, Equatorial Guinea, Gabon, Congo Southern Africa, Zambia Angola, Namibia, South Africa Western Sahara/Morocco to Libya (coastal) Libya, Egypt (coastal) South Africa Sierra Leone to Central African Republic/Gabon NE Angola, S DR Congo, NW Zambia

Forest and woodland savanna habitats Woodland savanna habitats Rainforest habitats Forest habitats Rainforest habitats; broad interorbital constriction; supraorbital ridge present. Rainforest habitats Woodland savanna habitats Woodland savanna habitats; commensal Rainforest habitats Rocky habitats; skull flattened Rocky habitats; skull flattened Forests, plantations, bushes, cultivations, rocky habitats Trees, bushes, gardens, rocky habitats Rocky habitats; skull flattened Rainforest habitats Woodland savanna habitats

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currently agree with Wahlert et al. (1993) that the myomorphouslike zygomasseteric structure exhibited by most living glirid species is convergent with that of muroids, and most advocate glirid ‘pseudomyomorphy’ as being derived from a protrogomorphous ancestor. Vianey-Liaud & Jaeger (1996) proposed that Gliridae are paraphyletic, hypothesizing that graphiurines and anomalurids are closely related and should possibly be placed in the same family because, in their view, both groups are descended from African Eocene zegdoumyids. However, results from all other morphological inquiries, as well as molecular information, do not support this hypothesis, but rather support the monophyly of Gliridae (Meng 1990, Wahlert et al. 1993, Hartenberger 1994, 1998, Catzeflis et al. 1995, Daams & De Bruijn 1995, Hänni et al. 1995, Robinson et al. 1997, Suzuki et al. 1997, Debry & Sagel 2001, Montgelard et al. 2002, 2003). Results of molecular analyses by Bentz & Montgelard (1999) and Montgelard et al. (2002, 2003) explicitly refute paraphyly and phylogenetic alliance with anomalurids. Ecological information on African dormice is rather sparse, and only a few species have been studied in a moderate amount of detail. Most species of dormice are primarily arboreal, and three species are mainly found in rocky habitats. The digits of the fore- and hindfeet grip small twigs and branches, and the long claws allow for climbing up slightly uneven surfaces such as rock faces, boulders, tree trunks and walls. Dormice live in many arboreal habitats, such as forests, scrublands, plantations, orchards and woodland savannas where there are suitable hollows and crevices for nesting (see also below). They have been recorded in holes of trees, woodpiles, human houses, thatched rondavels, gardens, kitchens and food stores, banana plantations, date palms, abandoned nests of weaverbirds and beehives. Rock-living species (such as Eliomys spp. and G. ocularis) are found on limestone cliffs, in caves, under boulders and in rock crevices. In all these habitats, dormice build spherical nests of vegetation (shredded bark, leaves), which are lined with hair or wool; entrance holes for nests are hard to find, perhaps because it is important that dormice need to be totally surrounded by warm

nesting material to ensure adequate thermoregulation (see below). Dormice do not dig (or live in) burrows and hence are rarely trapped on the ground. Most studies suggest that dormice are vegetarians or omnivores. Details on the diet vary according to species and location, but there are few detailed studies. Vegetable foods include nuts, seeds, berries, cocoa pods, oil palm nuts, paw-paw, bananas and other fruits. Animal foods include eggs and a wide variety of insects (e.g. grubs, moths and earwigs) and other invertebrates (millipedes). One species (Eliomys munbyanus) has been observed catching butterflies. Dormice are mostly nocturnal, although there are a few records of daytime activity. When active, dormice are agile and swift, running swiftly along twigs and branches and up and over walls and rocks. However, activity is temperature dependent. In cooler weather, dormice become lethargic and torpid (hence the vernacular name ‘dormouse’ from the Latin dormire = to sleep). In those species that have been studied, Tb drops dramatically when Ta is low and/or when food is limited. Social organizations, and how they may change during the year, are virtually unknown. Most species appear to be solitary except when mothers are with their young, and when a male associates with his mate and young. Some species (e.g. Graphiurus ocularis, G. platyops, G. kelleni) are known to emit a variety of specific vocalizations and displays, which indicate, for example, agonistic behaviour and nonaggressive communication. Some species (e.g. G. ocularis) may have territories, but nothing is known in this regard for most species. Reproductive behaviour is also poorly known: there are typically 2–3 young (maximum about six); reproduction in the temperate parts of Africa appears to be mainly during summer, but information from tropical Africa is too scanty to make any generalizations. The two African genera are distinguished by the length of the rostrum, form of the zygomatic plate, zygomasseteric musculature arrangement, relative position of certain cranial foramina, dental morphology and facial colour pattern (Table 14). Mary Ellen Holden

GENUS Eliomys Garden Dormice Eliomys Wagner, 1840. Gelehrte Anz. I. K. Bayer. Akad. Wiss., München 8 (37):297. (See Kryštufek & Kraft (1997) for clarification of the publication date for Eliomys Wagner [1839 or 1840]). Type species: Eliomys melanurus Wagner, 1840.

There are three living species in the genus Eliomys: E. quercinus is a European endemic that occurs fromWestern Europe east to the Urals, and on numerous Mediterranean islands; E. melanurus is distributed in southern Turkey, the Middle East (Sinai to Iraq) and eastern North Africa; and E. munbyanus is endemic to western North Africa (Holden 2005). The habitats of the two African species include desert scrub, rocky escarpments, limestone cliffs and cultivated areas. The genus is characterized externally by a conspicuous striking facial colour pattern (dark, broad eye-mask that extends posteriorly under ears, and which contrasts with pale (white or cream) cheeks and postauricular patches), and by the contrasting colours of tail and tail tip. The skull is distinguished by a long and narrow rostrum, an upward tilted zygomatic plate in which the anterior margin extends

slightly anteriorly to the infraorbital foramen, pseudomyomorphous (convergent myomorphous) zygomasseteric musculature arrangement, position of certain cranial foramina, the presence of four clearly defined transverse lophs or ridges on cheekteeth, and differences in incisor enamel microstructure (Ellerman 1940,Wahlert et al. 1993, von Koenigswald 1993, 1995, Nowak 1999) (Figure 18). All species of Eliomys are predominantly arboreal and partly terrestrial, and are nocturnal.They are agile climbers.The two species in Africa are omnivorous, and consume fruits, seeds, invertebrates and small vertebrates. Individuals of both species become torpid at low ambient temperatures. The closest living relative of Eliomys is probably Dryomys, which is distributed in Europe, the Middle East and central Asia. This

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Eliomys melanurus

Eliomys melanurus.

relationship is supported by cranial and dental characters (Wahlert et al. 1993, von Koenigswald 1993, 1995, Daams & De Bruijn 1995, Potapova 2001, Rossolimo et al. 2001), and by molecular data (Bentz & Montgelard 1999, Montgelard et al. 2003). The evolutionary history of Eliomys dates from the middle Miocene of Europe, and from the early Pliocene of western Asia and the Mediterranean region (Daams & De Bruijn 1995, Nadachowski & Daoud 1995). In Africa, it is known only from Pleistocene sediments in North Africa (Mein & Pickford 1992, McKenna & Bell 1997). Eliomys possibly reached Africa at least twice, once during the Messinian (6.7–5.2 mya) from Iberia, and later during the late Pleistocene from the eastern Mediterranean. Multiple colonizations may account for the lack of morphometric cohesion among certain populations of North African endemic E. munbyanus.

Figure 18. Skull and mandible of Eliomys melanurus (RMCA 91-090-M-45).

The two species are distinguished by body size, absolute lengths of ear and tail, degree of conspicuousness of postauricular patches, bushiness of tail, colour of the tail tip and by skull characters. The two species are mostly allopatric, although they are marginally parapatric in Libya. Mary Ellen Holden

Eliomys melanurus LARGE-EARED GARDEN DORMOUSE Fr. Lérot du Sud-Est Asiatique; Ger. Großohr Löffelbilch Eliomys melanurus (Wagner, 1840). Gelehrte Anz. I. K. Bayer. Akad. Wiss., München 8 (37): 299. Sinai (restricted to vicinity of Mt Sinai by Nader et al. 1983), Egypt.

Taxonomy Originally described in genus Myoxus. Eliomys melanurus has historically been listed as a synonym of E. quercinus (Niethammer 1959, Corbet 1978), but more recently recognized by most researchers as a separate species (Ellerman & Morrison-Scott 1966, Niethammer 1987, Holden 1993, 2005, Filippucci et al. 1988b, c, Harrison & Bates 1991, Kryštufek & Kraft 1997). The geographic distribution and taxonomy of E. melanurus remains unresolved (Holden 2005). Based on morphometric analyses, many authors (e.g. Kahmann & Thoms 1981, Niethammer 1987, Kryštufek & Kraft 1997 and references therein) assign the western North African populations to E. quercinus, and the eastern North African and Middle Eastern populations to E. melanurus. Filippucci et al. (1988b, c), Filippucci & Kotsakis (1995) and Filippucci & Capanna (1996) analysed allozymic and karyological characters and identified all North African and Middle Eastern populations as E. melanurus. Holden (2005) agreed with Kryštufek & Kraft (1997) that only eastern North African and Middle Eastern populations represent E. melanurus, but suggested (as

Delibes et al. 1980 did) that the western North African populations should be recognized as a separate species, E. munbyanus, and that it is probably closely related to E. melanurus. Synonyms: cyrenaicus. Subspecies: none. Chromosome number 2n = 48 (Filippucci et al. 1988b, c, 1990, Zima et al. 1995). Description Medium-sized dormouse. Dorsal pelage ranges from reddish- and yellowish-brown to yellowish-grey. Pelage soft, sometimes woolly, and moderately long (rump hairs 10–12 mm, guard hairs up to 17 mm). Ventral pelage white or cream slightly suffused with grey. Dorsal and ventral pelage colours clearly delineated. Head colour matches that of dorsal pelage; paler towards muzzle. Eyes large; dark eye-mask conspicuous. Cheeks cream or white, forming part of a pale lateral stripe that extends from cheeks to shoulders. Ears large, brown, oval-shaped. White or cream postauricular patches usually present, though sometimes inconspicuous. Hindfeet white. Tail long (ca 95% of HB), hairs 105

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shorter at base (3–4 mm) and longer at tip (up to 23 mm); dorsally black, or black with faint white tip, except for reddish- and yellowishbrown (similar to back) at base; undersurface similar to uppersurface. Skull long (35.9 mm), angular and broad (20.9 mm), with moderately long rostrum (15.6 mm) and anterior palatal foramina (4.3 mm). Upper toothrow (5.3 mm) relatively long. Auditory bullae long (11.7 mm) and inflated (mastoid breadth 17.3 mm) relative to skull length (measurements listed are mean values from Libya [Kahmann & Thoms 1981]). Nipples: 1 + 1 + 2 = 8. Geographic Variation None recorded. Similar Species (size comparisons refer to mean values only) Eliomys munbyanus. Smaller average head and body length (117 mm). Tail (108 mm), hindfeet (25 mm) and ears (24 mm) shorter. Dorsal pelage similar, postauricular patches usually more conspicuous. Tail not bushy; black with conspicuous white at tip. Ventral tail colour uniformly pale grey or brownish-white from base to tip in some populations. Skull slightly shorter (33.6 mm) with shorter rostrum (14.7 mm). Upper toothrow shorter (4.7 mm), and auditory bullae shorter (10.2 mm) and less inflated (mastoid breadth 16.8 mm) both absolutely and relatively (measurements listed are mean values from Morocco [Kahmann & Thoms 1981]). Not sympatric, although both species occur in Libya. Distribution Mediterranean Coastal BZ and northern margins of Sahara Arid BZ. Recorded from Libya, eastwards from Barqah (Cyrenaica) and Egypt including the Sinai Peninsula (Holden 2005). Extralimitally known from the Middle East, Iraq and southern Turkey. Habitat Coastal dunes and adjacent inland plateaux, Mediterranean scrubland, escarpments, steppe-deserts, rocky areas, mountains, gardens and human dwellings (Flower 1932,Wassif & Hoogstraal 1953, Ranck 1968, Osborn & Helmy 1980, Harrison & Bates 1991). Other

habitat records include limestone cliffs in coastal desert (Osborn & Helmy 1980), on upper slopes of a wadi near coastal escarpment under a large evergreen bush (Setzer 1957), in a small mountainside garden at 1700 m, near a spring in a garden, and in a Bedouin tent (Wassif & Hoogstraal 1953). In eastern North Africa, captured at elevations from sea level to 1700 m (Harrison & Bates 1991). Abundance Little information. Probably uncommon in eastern North Africa, based on the low numbers of specimens obtained at individual localities, as well as collectors’ notes (Setzer 1957, Ranck 1968). In Israel, trap success during one night in spring (Mar–May) was 11.7%; in other seasons at the same localities the animals were uncommon (Haim & Rubal 1995). Adaptations Arboreal and terrestrial; nocturnal (Haim & Rubal 1995, Qumsiyeh 1996). Resting metabolic rate is relatively low. Daily energy expenditure is conserved by entering torpor even when Ta is as high as 25 °C (Haim & Rubal 1995). In the Negev Highlands, Ta ranges from above 30 °C to below 0 °C during the winter; dormice trapped in winter at ambient temperatures close to 0 °C were found in traps in torpor with a Tb of 12 °C; torpor can last up to several days (Haim & Rubal 1995). Foraging and Food Omnivorous, predominantly insectivorous and carnivorous. Stomach contents have included insects and other invertebrates, and occasionally small mammals and other small vertebrates (Atallah 1978, Nader et al. 1983, Qumsiyeh 1996). These dormice readily enter live-traps (Osborn & Helmy 1980). Social and Reproductive Behaviour Little information. Osborn and Helmy (1980) characterized wild caught animals as extremely wild and aggressive. Reproduction and Population Structure Apparently solitary. Mean litter-size in captivity: 2.8 (Kahmann & Thoms 1981, Kahmann 1987); no information for wild individuals. Gestation: ca. 22 days (in captivity; Kahmann & Thoms 1981). In Israel, a pregnant ! was captured in Apr, and a lactating ! was captured in May (Kahmann 1981). Sex ratio 40% "", 60% !! (Egypt; Osborn & Helmy 1980). Predators, Parasites and Diseases Ectoparasites include the siphonapteran fleas Myoxopsylla laverani, Nesophyllus henleyi and Xenopsylla ramesis (Hoogstraal & Traub 1965b, Krasnov et al. 1999). Mites (not identified) were collected from some individuals collected in Saudi Arabia (Nader et al. 1983). In Israel, skeletal remains were found in pellets of Barn Owls Tyto alba; in Syria, skeletal elements were identified in pellets of Barn Owls and Long-eared Owls Asio otus (Obuch 2001). Conservation

Eliomys melanurus

IUCN Category: Least Concern.

Measurements Eliomys melanurus HB: 128.0 (111–144) mm, n = 10 T: 122.0 (100–136) mm, n = 10 HF: 26.7 (26–27) mm, n = 10 E: 27 (26–29) mm, n = 10 WT: 51.8 (38.4–63.0) g, n = 16*

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GLS: 35.9 (34.2–37.0) mm, n = 7 GWS: 20.9 (19.8–22.0) mm, n = 7 P4–M3: 5.3 mm, n = 8** Libya (Kahmann & Thoms 1981) *Western Mediterranean Coastal Desert, Egypt (Osborn & Helmy 1980)

**Mean value only Key References Kahmann & Thoms 1981; Harrison & Bates 1991; Osborn & Helmy 1980; Ranck 1968. Mary Ellen Holden

Eliomys munbyanus MAGHREB GARDEN DORMOUSE Fr. Lérot Nord-africain; Ger. Nordafrikanischer Löffelbilch Eliomys munbyanus (Pomel 1856). Comptes Rendus de l’Academie des Sciences, Paris 42: 653. Region d’Oran (Province of Oran), Algeria.

Taxonomy Originally described in genus Myoxus. Eliomys munbyanus has historically been considered a synonym of E. quercinus (Niethammer 1959, Kryštufek & Kraft 1997), although Filippucci et al. (1988a, b), Filippucci & Kotsakis (1995) and Filippucci & Capanna (1996) considered munbyanus to be synonymous with the Middle Eastern E. melanurus, an arrangement tentatively followed by Holden (1993).A few authors have suggested recognizing certain North African populations as a separate species, utilizing the names E. munbyanus (Delibes et al. 1980) or E. tunetae (Tranier & Petter 1978). Holden (2005) argues that recent morphometric analyses (Kryštufek & Kraft 1997), considered together with karyologic and allozymic analyses (Delibes et al. 1980, Filippucci et al. 1988a, b, Filippucci & Kotsakis 1995, Filippucci & Capanna 1996), support the recognition of E. munbyanus as a separate species that is probably closely related to E. melanurus. The assignment of synonyms and geographic distributions are based primarily on the results of Kryštufek & Kraft’s (1997) morphological study. Additional genetic and karyological sampling of E. munbyanus and North African E. melanurus populations is needed. Only the Moroccan population of E. munbyanus has been sampled for allozyme variation, and karyological data have only been reported from Moroccan and Tunisian populations. The hypothesized distribution of E. munbyanus is concordant with that of other mammalian endemics of the Maghreb (see Carleton & Van der Straeten 1997). Synonyms: denticulatus, lerotina, occidentalis, tunetae. Subspecies: none. Chromosome number: 2n = 46 (Tranier & Petter 1978, Delibes et al. 1980, Moreno & Delibes 1982, Filippucci et al. 1988a, Zima et al. 1995). Description Medium-sized dormouse. Dorsal pelage reddish- or yellowish-brown suffused with grey. Pelage soft, sometimes woolly and moderately long (rump hairs 10–11 mm, guard hairs up to 16 mm). Ventral pelage white slightly suffused with grey. Dorsal and ventral pelage colours clearly delineated. Head colour matches that of dorsal pelage; paler towards muzzle. Eyes large; dark eye-mask conspicuous. Cheeks cream or white, forming part of a pale lateral stripe that extends from cheeks to shoulders. Ears moderately large, brown, oval-shaped. White or reddish postauricular patches usually present. Hindfeet white. Tail long (ca. 92% of HB), tail hairs shorter at base (3–4 mm) and longer at tip (up to 19 mm); dorsally black with white at tip, except for reddish- and yellowish-brown (similar to back) at base; ventrally either uniformly pale grey or brownishwhite, or pale grey or brownish-white at base with a black middle section and white tip. Skull moderately long (33.6 mm), angular and broad (19.5 mm), with moderately short rostrum (14.7 mm) but comparatively long anterior palatal foramina (4.3 mm). Upper

toothrow (4.7 mm) relatively short. Auditory bullae moderately long (10.2 mm) and moderately inflated (mastoid breadth 16.8 mm) relative to skull length (measurements listed are mean values from Morocco [Kahmann & Thoms 1981]). Nipples: 1 + 1 + 2 = 8. Geographic Variation Ventral tail colour and pattern varies geographically. In Western Sahara and SW Morocco, the ventral tail colour is white or grey proximally, changing to solid black in the middle section, with a conspicuous white tip. The black region of the tail is sometimes fringed in white. Specimens collected from other regions of Morocco and Algeria have a totally greyish-white or white ventral tail colour (similar to European E. quercinus). Individuals from Tunisia and W Libya exhibit the same pattern as those from Western Sahara and SW Morocco (Kahmann & Thoms 1981). Tail length and body size also appear to vary among some populations. Populations from N Morocco have a longer tail length (tail length exceeds that of head and body) than surrounding populations (Moreno & Delibes 1982). This population also exhibits smaller body size (reflected in shorter HB and GLS lengths), and less inflated auditory bullae compared with populations from southern Morocco and Algeria (Cabrera 1932, Saint-Girons & Petter 1965, Moreno & Delibes 1982, Aulagnier & Thévenot 1986). Similar Species (size comparisons refer to mean values only) Eliomys melanurus. Larger average head and body length (128.0 mm). Tail (122.0 mm), hindfeet (26.7 mm) and ears (27.0 mm) longer. Dorsal pelage similar, postauricular patches usually less conspicuous. Tail bushy; distal dorsal colour uniformly black, or black with faint white tip.Ventral tail colour not paler, and does not vary significantly geographically. Skull slightly longer (35.9 mm) and longer rostrum (15.6 mm). Upper toothrow longer (5.3 mm) and auditory bullae longer (11.7 mm) and more inflated (mastoid breadth 17.3 mm), both absolutely and relatively (measurements listed are mean values from Libya – see Measurements). Not sympatric, although both species occur in Libya. Distribution Endemic to Africa. Mediterranean Coastal BZ. Recorded from Western Sahara, Morocco, Algeria, Tunisia and Libya (as far east as the Tarābulus region of Tripolitania) (Holden 2005). One population in the Sahara Arid BZ in the Fezzan. Recorded from sea level up to 3800 m. Habitat Captured in thick Mediterranean scrubland comprised of heath (Arbutus, Calluna, Erica), mock privet Phillyrea, pistachio 107

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1856, Ranck 1968, Kahman & Thoms 1981). Kahmann & Thoms (1981) stated that the primary physical requirement of a nest is to provide all-round body contact. Foraging and Food Probably omnivorous, consuming fruit, insects, seeds and sometimes eggs. In Tunisia, reported to be a pest in fruit plantations (favouring pomegranate) and in vegetable gardens (legumes, paprika, eggplant); may also eat chicken eggs (Kahmann & Thoms 1981). Kahmann & Thoms (1981) attempted stomach analyses of 11 individuals from Cap Bon, Tunisia, but were unsuccessful. Remains of insects and land snails were found near nest entrances in rocky fields, and individuals were observed capturing butterflies by springing into the air with both forelimbs stretched out in front (Kahmann & Thoms 1981). Bait made from bread dipped in cooking oil (Morocco; Moreno & Delibes 1982) and moistened oatmeal (Libya; Ranck 1968) attracted these dormice. Social and Reproductive Behaviour

Eliomys munbyanus

Pistacia, myrtle Myrtus, Mediterranean fan palm Chamaerops; young cork oak Quercus suber; pine plantations (Pinus halepensis, P. insignis); mixed oak forests (Quercus canariensis, Q. pyrenaica and Q. suber); dry, overgrazed habitats with isolated Acacia, pistachio and Mediterranean fan palm (Saint-Girons & Petter 1965, Moreno & Delibes 1982); large oases and adjoining areas in or near date palms Phoenix and tamarisk Tamarix (Ranck 1968); potato fields (Khidas 1993); and occasionally in prickly pear cactus Opuntia (Kahmann & Thoms 1981). Other habitats include coastal dunes, montane cedar forests, montane boulder fields and cultivated areas (Ranck 1968, Kahmann & Thoms 1981, Moreno & Delibes 1982, Aulagnier & Thévenot 1986, Kowalski & Rzebik-Kowalska 1991). In the Fezzan, captures were in or adjacent to oases. Abundance Uncommon (Ranck 1968, Moreno & Delibes 1982, Aulagnier & Thevenot 1986, Khidas 1993). Adaptations Predominantly arboreal, partly terrestrial; nocturnal (Kahmann & Thoms 1981, Aulagnier & Thévenot 1986). Often enter torpor in winter in response to prolonged cooler ambient temperatures (Kahmann & Thoms 1981). In Libya, individuals were infrequently captured during the winter months when ambient temperatures at night often dropped below –1 °C; Ranck (1968) suggested that the seemingly low abundance might be explained by inactivity of torpid individuals. In Morocco, one individual was caught during a night during which the ambient temperature dropped below 0 °C (Moreno & Delibes 1982). Nests have been found in many situations: in holes in trees (including tamarisk, olive, willow, poplar and several species of palm), in shrubs (small palms, and rarely prickly pear cactus), in rock crevices, caves and at the bases of large rocks, and in thatched roofs, alcoves, attics and conduits of huts (Ranck 1968, Kahmann & Thoms 1981). Materials used to construct nests include grass, barley stems, palm fibre, goat hair, sheep and possibly dromedary wool, and even flower clusters of Acacia (Pomel

No information.

Reproduction and Population Structure Solitary. Reproductively active in spring and, at lower altitudes, autumn (Kahmann & Thoms 1981, Moreno & Delibes 1982). Litter-size: probably 4–6, although one pregnant ! contained eight embryos (Kahmann & Thoms 1981, Moreno & Delibes 1982). Young stay in the nest for approximately seven weeks (Kahmann & Thoms 1981). In N Morocco, lactating !! were captured in Nov (Moreno & Delibes 1982). In Tunisia, two pregnant !! were collected in Mar, and one pregnant ! was captured in Apr (Kahmann & Thoms 1981). In Tunisia, sex ratio was found to be male-biased, but this may reflect a sampling artefact due to seasonal differences in activity between sexes (Kahmann & Thoms 1981). Predators, Parasites and Diseases Principal host for the hoplopleurid louse Schizophthirus pleurophaeus (Durden & Musser 1994). In Algeria, skeletal remains were identified in owl pellets (Kowalski & Rzebik-Kowalska 1991) and in the scat of a jackal (Khidas 1986). Conservation

IUCN Category: Least Concern.

Measurements Eliomys munbyanus HB: 117 (100–140) mm, n = 26 T: 108 (96–118) mm, n = 26 HF: 25 (22–27) mm, n = 26 E: 24 (20–27) mm, n = 26 WT: 52 (42–62) g, n = 14 GLS: 33.6 (31.7–35.6) mm, n = 8 GWS: 19.5 (18.6–20.1) mm, n = 8 P4–M3: 4.7 mm* Morocco (Kahmann & Thoms 1981) *Mean value only Key References Kahmann & Thoms 1981; Kryštufek & Kraft 1997; Moreno & Delibes 1982; Niethammer 1959; Ranck 1968. Mary Ellen Holden

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GENUS Graphiurus African Dormice Graphiurus Smuts, 1832. Enumer. Mamm. Capensium, pp. 32–33. Type species: Sciurus ocularis Smith, 1829.

Graphiurus murinus.

Figure 19. Skull and mandible of Graphiurus nagtglasii (RMCA RG 5556).

The 14 living species of the genus Graphiurus occur throughout subSaharan Africa in tropical lowland rainforest, montane evergreen and semi-deciduous forest, moist and arid savanna woodlands and grasslands, as well as the Nama- and succulent-Karoo biomes and fynbos in southern Africa. Treeless deserts, such as the Namib, are devoid of dormice, except near oases. Primary habitat requirements for African dormice are trees, woody vines, bushes and weathering granitic outcrops (kopjes) that provide shelter and nesting sites. The genus is characterized externally by a generally plain-coloured face without obvious markings in most species (except G. ocularis), and the colour of the tail and tail tip. The skull is distinguished by a short, often broad rostrum, very narrow zygomatic plate that is situated entirely beneath the infraorbital foramen, modified hystricomorphous zygomasseteric musculature arrangement, differences in position of certain cranial foramina, V-shaped cutting edges of incisor tips (forming a V-shaped notch from anterior view) in most species, and indistinct, usually incomplete lophs or ridges on cheekteeth (Ellerman 1940, Wahlert et al. 1993, Nowak 1999) (Figure 19). Many closely related species of Graphiurus are difficult to distinguish, although a few species are easy to identify because of their unique prominent characters: G. crassicaudatus has a broad interorbital constriction and pronounced supraorbital ridging, and G. nagtglasii is the largest species and has a distichous tail. Species that typically dwell in rock crevices, such as G. platyops and G. ocularis, have a flattened cranium that is easily distinguished from the vaulted cranium found in all other species. Graphiurus ocularis is also easily identified by its striking facial colour pattern and its reduced, circular premolar. In their morphology and habits, all sub-Saharan African dormice resemble small-bodied arboreal squirrels, and like these animals all species of Graphiurus are arboreal; even those using crevices in rocky habitats, such as kopjes, for nesting sites are excellent climbers. All species are nocturnal, although at least one species (G. platyops) has also been documented to be active at dawn (Wilson 1975), and another species (G. angolensis) was reportedly active during the day (field notes; specimen labels). Most species are omnivorous, and consume fruits, insects, seeds and nuts; G. ocularis is predominantly insectivorous (Channing 1984). Some species, such as G. lorraineus and G. murinus, enter torpor during periods of low ambient temperature and inadequate food supply (Lachiver & Petter 1969, Webb & Skinner 1996b). Hartenberger (1994) considered many aspects of graphiurine morphology to be primitive, and speculated that the group has been in Africa since the Miocene; late Miocene graphiurines are recorded from Namibia (Mein et al. 2000b) and South Africa (Denys 1990b). Mein et al. (2000b) described the late Miocene Otaviglis from Namibia as the oldest known graphiurine (10–11 mya), possibly derived from the middle Miocene Microdryomys found in North Africa, and suggested it could be ancestral to Graphiurus. Molecularclock calibrations derived from phylogenetic analyses of nuclear and mitochondrial gene sequences indicate that the adaptive radiation within Graphiurus occurred 8–10 mya, which pre-dates the oldest 109

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fossil representative of the genus (early Pliocene, ca. 5 mya), but is consistent with the late Miocene graphiurine records (Montgelard et al. 2003). The time frame also brackets a period of low sea levels and extensive interchange of faunas between Europe and Africa. Following the colonization of Africa by a late Miocene ancestor, graphiurines underwent an adaptive radiation resulting in a modern fauna that is richer in species than those in all the other genera of dormice combined (Montgelard et al. 2003). The taxonomic revision of extant Graphiurus by Genest-Villard (1978a) was based mostly on size grades and underestimated species diversity, particularly in G. murinus and closely related species. Subsequently, certain species were defined in reports covering different geographical regions (e.g. Robbins & Schlitter 1981, Ansell & Dowsett 1988, Holden 1996). Profiles of the 14 species in this volume are based on literature sources, examination of museum specimens and preliminary multivariate analyses of cranial and dental measurements (M. E. Holden unpubl.). It is likely that future studies that incorporate molecular data will show that some species recognized here actually contain two or more separate species. Graphiurus has been divided into as many as four separate genera (e.g. Allen 1939, Holden 1996, Pavlinov & Potapova 2003): Aethoglis, containing the largest African dormouse G. nagtglasii (sometimes erroneously including G. monardi); Graphiurus comprising G. ocularis, with its reduced, simple upper premolar; Gliriscus consisting of the rupicolous G. platyops and G. rupicola; and Claviglis the so-called

‘tree dormice’, to which the remaining species of Graphiurus were assigned. Two of these, Graphiurus and Claviglis, have often been retained as subgenera (e.g. Ellerman et al. 1953, Rosevear 1969). No published studies based upon a broad sample of species have addressed the validity of these subgeneric boundaries as used by past authorities, nor have they presented hypotheses of relationships among species. However, phylogenetic analyses of cranial and middle ear morphology by Pavlinov & Potapova (2003) identified three monophyletic groups (subgenera) within Graphiurus: Aethoglis (containing G. nagtglasii), Claviglis (containing G. crassicaudatus) and Graphiurus (containing all other Graphiurus species). Their study suggested that first G. nagtglasii, then G. crassicaudatus, diverged early in the evolution of African Dormice, and that the remaining taxa that they sampled (angolensis, christyi, kelleni, lorraineus, murinus, ocularis, parvus and surdus) form a separate monophyletic group. Pavlinov & Potapova’s (2003) subgeneric arrangement is followed by Holden (2005). Here, the species are listed alphabetically. The 14 species are distinguished externally by body size, length of tail, ear and hindfoot, degree of conspicuousness of eye-mask, and colour of the tail tip. The skulls are differentiated by the overall shape of the skull, the shape of the zygomatic arch, presence/absence of supraorbital ridges, length and relative inflation of auditory bullae, differences in toothrow measurements and relative size of the upper premolar, and several skull dimensions. Mary Ellen Holden

Graphiurus angolensis ANGOLAN AFRICAN DORMOUSE Fr. Graphiure d’Angola; Ger. Angolischer Bilch Graphiurus angolensis de Winton, 1897). Ann. Mag. Nat. Hist., ser. 6, 20: 320. Caconda, Angola.

Taxonomy Originally described in the genus Gliriscus. Allen (1939) recognized G. angolensis as a species distinct from G. platyops, G. rupicola and parvulus. Ellerman et al. (1953) included G. angolensis, G. rupicola and parvulus as subspecies of G. platyops. Genest-Villard (1978a) placed G. rupicola as a subspecies of G. platyops, but synonymized G. angolensis and parvulus under G. murinus. Ansell (1974, 1978) recognized that the north-western Zambian population (identified by him as G. platyops parvulus) is morphologically and ecologically different from G. platyops. Holden (1993) provisionally listed populations from Angola, S DR Congo and NW Zambia under G. platyops. Populations in Angola and NW Zambia exhibit a distinctive skull morphology that is consistently separable from that of G. platyops and G. rupicola (M. E. Holden unpubl.). Ansell (1974, 1978) had correctly surmised that these populations are probably aligned with G. microtis. Here, following Allen (1939), G. angolensis is considered as a valid species and distinct from G. platyops and G. rupicola.The form parvulus (as described by Monard 1933) is probably a junior synonym of G. angolensis (Holden 2005). Synonyms: dasilvai, parvulus. Subspecies: none. Chromosome number: not known.

Ventral pelage white or cream slightly suffused with grey. Dorsal and ventral pelage colours clearly delineated. Head colour matches that of dorsal pelage, slightly paler towards muzzle. Eyes large; eye-mask conspicuous. Ears brown, large, rounded. Cheeks cream or white, forming part of a pale lateral stripe that extends from cheeks to shoulders. Cream postauricular patches usually present. Hindfeet white, or white with dark metatarsal streak. Tail moderately long (ca. 80% of HB), tail hairs shorter at base (5–10 mm) and longer at tip (up to 33 mm). Tail colour generally matches dorsal pelage. White hairs are mixed throughout tail; tip white. Skull long, robust and moderately broad (15.5 mm), with a relatively vaulted braincase (height of braincase 7.7 mm). The appearance of a vaulted braincase is augmented by the concave curvature of the braincase (lateral view) and large auditory bullae. Interorbital constriction moderately narrow (4.2 mm), anterior palatal foramina comparatively long (3.4 mm) and wide (2.1 mm), and auditory bullae long (9.0 mm) and inflated relative to skull length (mean values from Kabompo and Zambezi [formerly Balovale], Zambia; M. E. Holden unpubl.). Nipples: 1 + 1 + 2 = 8.

Description Small dormouse. Dorsal pelage dark brown, rufous, golden, or drab brown, with darkening of pelage towards the mid-line of head and back in some individuals. Dorsal pelage soft, sleek, thick and moderately long (rump hairs 8 mm, guard hairs up to 12 mm).

Geographic Variation Individuals from C Angola are notably darker and usually exhibit a dark metatarsal streak on the hindfeet, whereas individuals from S and C Angola (type locality of parvulus) and Zambia are drab or golden brown, usually with white hindfeet.

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Graphiurus angolensis

Ear pinnae shorter (on average) in Angolan specimens (13.5 mm) than in Zambian specimens (15.9 mm) (mean values from Ondjiva and Caconda, Angola, and from Kabompo and Zambezi [formerly Balovale], Zambia; M. E. Holden unpubl.).

Habitat Little information. Woodland savanna. Most collecting localities in Angola are in or near wetter miombo woodland, and in wetter miombo woodland and Zambezian dry evergreen forest in Zambia. Individuals have also been captured in human dwellings.

Similar Species (size comparisons refer to mean values only) Graphiurus microtis. Similar mean head and body length (98.8 mm). Tail (mean 75.2 mm) and hindfeet (mean 16.9 mm) absolutely and relatively shorter. Dorsal pelage often similar in colour, but grey in some populations. Skull similar in length (27.4 mm), but slightly narrower (15.0 mm). Interorbital constriction (3.9 mm) slightly narrower. Anterior palatal foramina 3.4 mm long, 2.1 mm wide. Mean upper toothrow length (3.0 mm) shorter, and mean upper premolar breadth (0.8 mm) narrower. Auditory bullae (8.1 mm) absolutely and relatively shorter and less inflated (measurements listed are mean values from Zimbabwe; M. E. Holden unpubl.). Parapatric in NW Zambia (Ansell 1978). Graphiurus microtis occurs in savannas throughout most of subSaharan Africa. Graphiurus rupicola. Slightly larger head and body length (mean 110 mm). Tail (mean 104.2 mm) and hindfeet (mean 21.5 mm) absolutely and relatively longer. Dorsal pelage grey. Mean skull length slightly longer (31.3 mm). Interorbital constriction broader (5.0 mm) and palate longer (10.4 mm) both absolutely and relatively. Braincase absolutely more vaulted (8.0 mm), but flatter relative to skull length. Anterior palatal foramina 3.4 mm long, 2.3 mm wide. Upper toothrow (3.4 mm) similar in absolute length, but shorter relative to skull length (measurements listed are mean values from Erongo, Karibib and Mt Brukkaros, Namibia; M. E. Holden unpubl.). Parapatric in Angolan highlands; also occurs in Namibia and NW South Africa.

Abundance Common in the Kabompo and Zambezi districts of Zambia (Ansell 1978). The species is probably common in the interior plateaux region of Angola (Bocage 1890, Hill & Carter 1941).

Distribution Endemic to Africa. Zambezian Woodland BZ. Recorded only from C and S Angola and NW Zambia (Holden 2005).

Adaptations Arboreal, probably nocturnal. (Most species of African Dormice are nocturnal.) Some individuals of this species were noted to be active during the day in C Angola (field notes; specimen labels). These dormice have most often been caught in trees; in Angola, they were also found in abandoned beehives (field notes; specimen labels). In Zambia, they have sometimes been captured in buildings; one ! with young was caught in the roof of an African hut, another solitary ! was caught among some planks in a carpenter’s shop, and one " was obtained in a store (Ansell 1978; specimen labels). Foraging and Food Probably omnivorous. Little is known regarding the diet. An individual captured in Angola was recorded to have eaten tree grubs and the fruit of a ‘parasitic growth on trees’ (field note, Phipps-Bradley Angola Expedition). One individual was caught in a trap baited with meat (Chubb 1909). Social and Reproductive Behaviour Little information. Lactating !! are often caught with young. One ! was recorded as being caught ‘with four half-grown young, but apparently no longer lactating’ (specimen label). Monard (1935) states that Angolan Dormice are aggressive. Reproduction and Population Structure Litter-size: probably 3–5 (Ansell 1963; specimen labels). In Zambia, a ! captured in late Oct gave birth the following day to three young (Ansell 1963). Predators, Parasites and Diseases No information. Conservation

IUCN Category: Data Deficient.

Measurements Graphiurus angolensis HB: 98.8 (79–112) mm, n = 49 T: 79.2 (70–96) mm, n = 45 HF: 18.4 (17–20) mm, n = 50 E: 15.9 (14.5–18) mm, n = 50 WT: n. d. GLS: 28.2 (26.3–30.8) mm, n = 36 GWS: 15.5 (14.4–16.6) mm, n = 21 P4–M3: 3.2 (2.9–3.5) mm, n = 39 Kabompo and Zambezi, Zambia (M. E. Holden unpubl.) Key References Ansell 1978; Hill & Carter 1941; Monard 1935. Graphiurus angolensis

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Graphiurus christyi CHRISTY’S AFRICAN DORMOUSE Fr. Graphiure de Christy; Ger. Christys Bilch Graphiurus christyi Dollman, 1914. Revue Zoologique Africaine 4 (1): 80. Mambaka, DR Congo.

Taxonomy Morphologically similar to some named forms currently synonymized under G. murinus, some of which occur in areas adjacent to this species, e.g. vulcanicus from the Virunga Mts. Other named forms synonymized under G. murinus that occur in adjacent areas are morphologically distinct, e.g. soleatus, from the Rwenzori Mts. Synonyms: none. Chromosome number: not known. Description Small dormouse. Dorsal pelage medium brown, rufous-brown or rufous golden-brown. Dorsal pelage soft, silky and moderately thick (rump hairs 6–8 mm, guard hairs up to 11 mm). Ventral pelage grey washed with white. Dorsal and ventral pelage colours not clearly delineated. Head colour matches that of dorsal pelage. Eyes large; eye-mask usually conspicuous. Ears brown, large, rounded. Cheeks usually white. Postauricular patches not present. Hindfeet white with dark metatarsal streak. Tail moderately long (ca. 82% of HB), hairs shorter at base (3–5 mm) and longer at tip (up to 21 mm). Tail colour generally matches that of dorsal pelage and does not exhibit white tip. Skull medium length (28.0 mm), moderately narrow (15.1 mm) and moderately vaulted (height of braincase 8.1 mm). Interorbital constriction (4.7 mm) narrow. Supraorbital ridges present. Premaxilla and nasal bones often extend farther beyond the anterior face of the incisors than in similar species. Anterior palatal foramina moderately long (mean 3.0 mm) and wide (mean 2.2 mm), palate moderately long (8.5 mm), auditory bullae short (7.4 mm) and not inflated relative to skull length. Anterior chamber of auditory bullae markedly less inflated than posterior chambers in some individuals (measurements listed are mean values from DR Congo; M. E. Holden unpubl.). Nipples: 1 + 1 + 2 = 8. Geographic Variation None recorded. Similar Species (size comparisons refer to mean values only) Graphiurus lorraineus. Smaller head and body length (mean 83 mm), ears shorter (mean 12.1 mm). Dorsal pelage usually more rufous. Skull (24.5 mm) shorter and anterior palatal foramina narrower (1.7 mm) relative to skull length. Nasals and premaxilla usually do not extend as far beyond incisors. Palate absolutely shorter (7.8 mm) but relatively similar. Anterior palatal foramina 2.7 mm long, 1.7 mm wide. Auditory bullae similar in length (7.2 mm), but longer relative to skull length (measurements listed are mean values from DR Congo; M. E. Holden unpubl.). Sympatric in SW Cameroon and NE DR Congo. Occurs in West and central Africa. Graphiurus surdus. Similar mean head and body length (99.0 mm), ears shorter (mean 12.3 mm). Dorsal pelage usually greyishbrown, not rufous. Tail (mean 72 mm) absolutely and relatively shorter (ca. 73% of HB). Zygoma robust, and the anterior, superior margins are relatively straight when viewed from side (see Holden 1996). Palate longer (9.3 mm); anterior palatine foramina absolutely and relatively shorter (2.8 mm) and narrower (1.8 mm) (measurements listed are mean values from Cameroon, Equatorial Guinea and Gabon; Holden 1996). Not sympatric,

Graphiurus christyi

although both species have been collected in SW Cameroon and N DR Congo. Graphiurus crassicaudatus. Smaller head and body length (mean 92.6 mm). Dorsal pelage rufous-brown, similar to some individuals of G. christyi. Skull shorter (26.6 mm). Breadth of skull (16.1 mm) and interorbital constriction (4.9 mm) markedly broader relative to skull length. Supraorbital ridges present. Anterior palatal foramina absolutely and relatively shorter (2.5 mm) and narrower (1.6 mm). Palate (9.4 mm) and upper toothrow (3.8 mm) absolutely and relatively longer (measurements listed are mean values from S Cameroon; M. E. Holden unpubl.). Not sympatric, though their distributions overlap in SW Cameroon; occurs in West and west-central Africa. Distribution Endemic to Africa. Recorded from Rainforest BZ (mainly East Central Region) with one outlier in West Central Region. Recorded from NE DR Congo (many localities, all north of the Congo and Lualaba rivers) and SW Cameroon (one locality). A specimen from Inkongo, C DR Congo, identified as this species (Hatt 1940a) is, in fact, a specimen of G. surdus (Holden 1996). Habitat Rainforest. According to Hatt (1940a), these dormice occur in high forest. Abundance Little information and rarely encountered. The Lang–Chapin Congo Expedition collected 29 specimens at Medje, DR Congo, but caught only one specimen at each of the three other localities where they encountered this species. Robbins & Schlitter

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(1981) obtained four individuals from Lolodorf, SW Cameroon, and Schlitter et al. (1985) collected eight individuals from Yalosemba, DR Congo.

Predators, Parasites and Diseases Little information. Some specimens in museum collections have several louse exoskeletons attached to pelage and tail hairs

Adaptations Arboreal and probably nocturnal. Nests in hollow trees (Hatt 1940a).

Conservation

Foraging and Food Probably omnivorous, eating fruit, insects, seeds and nuts. Notes taken by the Lang–Chapin Congo Expedition indicated that the stomachs of four individuals contained ‘a whitish or somewhat greenish paste-like vegetable matter’ (Hatt 1940a). Social and Reproductive Behaviour In DR Congo, five individuals were collected from a hollow tree that contained no nest: two "", two !! and one escaped (Hatt 1940a). Reproduction and Population Structure Litter-size: probably 2–3 (Hatt 1940a). In DR Congo, a ! was found in Jan with two young (eyes closed); two additional young (eyes closed) were collected from a separate nest; and one pregnant ! contained three embryos (Hatt 1940a).

IUCN Category: Least Concern.

Measurements Graphiurus christyi HB: 97.6 (86–107) mm, n = 27 T: 79.8 (73–95) mm, n = 25 HF: 18.0 (16–20) mm, n = 28 E: 14.2 (12–17) mm, n = 27 WT: 29.0 (25–33) g, n = 6 GLS: 28.0 (26.7–29.7) mm, n = 23 GWS: 15.1 (13.3–16.7) mm, n = 22 P4–M3: 3.2 (3–3.3) mm, n = 29 DR Congo (M. E. Holden unpubl.) Key References Hatt 1940a; Holden 1996; Robbins & Schlitter 1981; Schlitter et al. 1985. Mary Ellen Holden

Graphiurus crassicaudatus THICK-TAILED AFRICAN DORMOUSE Fr. Graphiure à grosse queue; Ger. Dickschwanz-Bilch Graphiurus crassicaudatus (Jentink, 1888). Notes from the Leyden Museum 10: 41–42. Hill Town, Du Queah River, Liberia.

Taxonomy Originally described in the genus Claviglis. Rosevear (1969) and Holden (1996) hypothesized that the morphological similarity between G. crassicaudatus and G. nagtglasii (formerly G. hueti) indicated a close phylogenetic relationship between the two species. In contrast, recent cladistic analysis of African dormice based on cranial and middle ear characters does not support this conclusion (Pavlinov & Potapova 1993, see also Holden 2005). Synonyms: dorotheae (see Allen 1939, Rosevear 1969). Subspecies: none. Chromosome number: not known. Description Small dormouse. Dorsal pelage usually rufousbrown. Pelage soft and short (rump hairs 4–5 mm, guard hairs up to 10 mm). Ventral pelage grey washed with ochre, cream or white. Dorsal and ventral pelage colours clearly delineated. Head colour matches that of dorsal pelage; muzzle short. Eyes large; eye-mask conspicuous in some individuals. Ears brown, short and rounded. Postauricular patches usually not present. Hindfeet white, or white with a dark metatarsal streak. Tail short (ca. 65% of HB), hairs shorter at base (3–4 mm) and longer at tip (up to 27 mm).Tail colour generally matches that of dorsal pelage. A few white hairs are mixed throughout the tail, but tip is not white. Skull broad (16.1 mm), with vaulted braincase (height of braincase 7.9 mm) and conspicuously wide interorbital constriction (4.9 mm) with supraorbital ridges. Anterior chamber of auditory bullae usually markedly less inflated than posterior chambers. Zygomatic arches flare out at a 90 degree angle from the rostrum. Rostrum short and narrow, with short nasal bones (8.9 mm). Anterior palatal foramina comparatively short (mean 2.5 mm) and very narrow (mean 1.6 mm) relative to skull

length. Palate (9.4 mm) and upper toothrow (3.8 mm) relatively long (measurements listed are mean values from southern Cameroon; M. E. Holden unpubl.). Nipples: 1 + 1 + 2 = 8. Geographic Variation

None recorded.

Similar Species (size comparisons refer to mean values only) Graphiurus lorraineus. Smaller head and body length (mean 83.0 mm). Dorsal pelage similar in colour. Skull shorter (24.5 mm) and narrower (13.8 mm), with much narrower interorbital constriction (4.3 mm) and without supraorbital ridges. Anterior palatal foramina 2.6 mm long, 1.7 mm wide. Palate (7.8 mm) and upper toothrow (3.1 mm) absolutely and relatively shorter, auditory bullae similar in length (7.2 mm), but relatively longer (measurements listed are mean values from DR Congo; M. E. Holden unpubl.). Sympatric at several localities in Liberia, Côte d’Ivoire and S Cameroon; occurs in West and central Africa. Graphiurus surdus. Larger head and body length (mean 99.0 mm), with longer hindfeet (mean 20.8 mm). Dorsal pelage greyish-brown, with no rufous hue. Skull narrower (14.6 mm), with narrower interorbital constriction (4.5 mm) and without supraorbital ridges. Anterior palatal foramina 2.8 mm long, 1.8 mm wide. Upper toothrow (3.2 mm) absolutely and relatively shorter (measurements listed are mean values from Cameroon, Equatorial Guinea and Gabon; Holden 1996). Sympatric in SW Cameroon, also occurs in DR Congo. Graphiurus christyi. Larger head and body length (mean 97.6 mm). Dorsal pelage sometimes similar in colour. Skull slightly longer 113

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and in grass near houses (Sanderson 1940, Rosevear 1969, Robbins & Schlitter 1981; specimen labels), and even in a hole in a concrete culvert under railway tracks (Schlitter et al. 1985). In Nigeria, Sanderson (1940) found large, spherical nests a few feet above the ground in dense vegetation and, in Cameroon, he found five individuals living in a nest in a hollow tree. This nest was made of dead leaves, and lined with fibre from a species of nut that lay in large quantities on the surrounding ground. (Rosevear 1969 correctly noted that the specimens Sanderson (1940) identified as Claviglis spurrelli and Claviglis haedulus actually represent G. crassicaudatus.) Captive animals adapted the nests of weaver-birds for their use (Rosevear 1969). Foraging and Food Probably omnivorous, consuming nuts (Rosevear 1969, Robbins & Schlitter 1981), insects (Rosevear 1969) and probably fruit. Social and Reproductive Behaviour In Cameroon, Sanderson (1940) reported five adults in the same nest in a hollow tree. Two !! were caught, the other three escaped. This species bred readily in captivity (Rosevear 1969). Graphiurus crassicaudatus

Reproduction and Population Structure (28.0 mm). Breadth of skull (15.1 mm) and interorbital constriction (4.7 mm) narrower relative to skull length, and without supraorbital ridges. Anterior palatal foramina absolutely and relatively longer (3.0 mm) and wider (2.2 mm). Palate (8.5 mm) and upper toothrow (3.2 mm) absolutely and relatively shorter (measurements listed are mean values from DR Congo; M. E. Holden unpubl.). No records of sympatry, though geographic range overlaps in SW Cameroon; also occurs in DR Congo. Distribution Endemic to Africa. Rainforest BZ (Western and West Central Regions) and Northern Rainforest–Savanna Mosaic. Recorded from Liberia and Guinea east to SW Cameroon (excluding Benin) and Bioko I. (see Rosevear 1969, Eisentraut 1973). Habitat

In or near primary and secondary rainforest.

Abundance Uncommon. The species is represented by a total of approximately 50 museum specimens, and it is considered rare or at least difficult to trap (Heim de Balsac 1967a, Robbins & Schlitter 1981, Happold 1987, Grubb et al. 1998).

No information.

Predators, Parasites and Diseases An ectoparasitic hoplopleurid louse, Schizophthiris sp., has been recorded on this species in Liberia (Kuhn & Ludwig 1965). Conservation

IUCN Category: Data Deficient.

Measurements Graphiurus crassicaudatus HB: 92.6 (83–98) mm, n = 11 T: 59.4 (55–70) mm, n = 9 HF: 17.7 (16–19) mm, n = 12 E: 13 (11–14) mm, n = 9 WT: 24.8 (20–29) g, n = 6 GLS: 26.6 (25.2–27.8) mm, n = 8 GWS: 16.1 (15.7–16.6) mm, n = 6 P4–M3: 3.8 (3.4–4.2) mm, n = 14 S Cameroon (M. E. Holden unpubl.) Key References Holden 1996; Robbins & Schlitter 1981; Rosevear 1969.

Adaptations Arboreal. Individuals have been collected from hollow trees, on horizontal branches and vines, among bushes, among rocks

Mary Ellen Holden

Graphiurus johnstoni JOHNSTON’S AFRICAN DORMOUSE Fr. Graphiure de Johnston; Ger. Johnstons Bilch Graphiurus johnstoni Thomas, 1898. Proc. Zool. Soc. Lond. 1897: 934. Zomba, Malawi.

Taxonomy Ansell & Dowsett (1988), Ansell (1989b), Happold & Happold (1989a) and Holden (1993) synonymized johnstoni under G. kelleni. Recent re-examination and comparisons of museum specimens (including all holotypes), and preliminary multivariate

analyses, indicate that G. johnstoni is a separate valid species (M. E. Holden unpubl.). Morphologically, it is closely related to G. lorraineus and distinct from G. kelleni. Here, G. johnstoni is retained as a valid species, pending further revision of the genus (see also Holden

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Graphiurus johnstoni

2005). Few specimens exist that can be attributed to G. johnstoni with certainty. If future research indicates that G. johnstoni and G. lorraineus are conspecific, the latter would be a junior synonym of G. johnstoni; this would substantially modify and enlarge the geographic range of what is now considered G. johnstoni. Synonyms: none. Chromosome number: not known. Description Small dormouse. Dorsal pelage reddish-brown. Dorsal pelage soft and short (rump hairs 5–6 mm, guard hairs up to 8–9 mm). Ventral pelage grey, moderately suffused with buff or cream. Dorsal and ventral pelage colours not clearly delineated. Head colour matches that of dorsal pelage. Eyes large; eye-mask inconspicuous. Ears brown, short and rounded. Cheeks cream or grey suffused with cream. Postauricular patches not present or inconspicuous. Hindfeet cream with dark metatarsal streak. Tail long (ca. 92% of HB), hairs shorter at base (2–3 mm) and longer at tip (up to 19 mm). Tail appears splayed because the hairs project laterally. Tail colour generally matches that of dorsal pelage and is uniform in colour, with sparse or no white hairs, and without white at tip. Skull short (23.6 mm), moderately vaulted (height of braincase 7.3 mm) and broad (13.9 mm) with relatively short rostrum (length of nasal bones 8.4 mm). Interorbital constriction broad (4.0 mm), toothrow long (3.4 mm) and upper premolar wide (1.0 mm) relative to skull length. Anterior palatal foramina moderately long (2.7 mm) and moderately narrow (mean 1.7 mm) relative to skull length. Auditory bullae relatively short (6.8 mm) and moderately inflated. Nipples: 1 + 1 + 2 = 8. Geographic Variation None recorded. Similar Species (size comparisons refer to mean values only) Graphiurus lorraineus. Larger head and body length (mean 83.0 mm). Ear (12.6 mm) slightly larger. Tail relatively shorter (65.7 mm) with similar colouration. Dorsal pelage colour similar. Skull slightly longer (24.5 mm). Rostrum relatively short as in G. johnstoni, but length of nasal bones absolutely longer (9.1 mm). Anterior palatal foramina 2.6 mm long, 1.7 mm wide. Upper toothrow slightly shorter (3.1 mm) and upper premolar narrower (0.8 mm). Not sympatric; G. lorraineus occurs in West and central Africa. Graphiurus microtis. Larger head and body length (98.8 mm). Tail relatively shorter (75.2 mm), usually with white hairs mixed throughout, and with conspicuous white tip. Dorsal pelage colour sometimes similar, but beige or grey in some populations. Skull longer (27.4 mm) and wider (15.0 mm), auditory bullae longer (8.1 mm) and more inflated. Interorbital constriction (3.9 mm) similar in breadth, but relatively narrower. Anterior palatal foramina absolutely and relatively longer (3.4 mm) and wider (2.1 mm). Tooth row shorter (3.0 mm) both absolutely and relative to skull length. Sympatric in Thyolo, Malawi, but probably not syntopic; widespread in savannas of eastern and central Africa. Distribution Endemic to Africa. Zambezian Woodland BZ. Recorded only from S Malawi; limits of geographic range unknown.

Graphiurus johnstoni

Habitat Little information. Habitats on the Shire Highlands (ca. 900–1500 m) include sub-montane forests, miombo woodlands, farmlands, tobacco fields and secondary growth (Happold & Happold 1989a, b, 1997, 1998). Several specimens have been found in houses surrounded by ornamental gardens. Mean annual rainfall is ca. 1300 mm, with considerable annual variation. Abundance No information. Rarely encountered. Remarks Little information. Probably arboreal and nocturnal. Happold & Happold (1997) captured an individual in a farmhouse, indicating that this species may nest in human dwellings. Conservation

IUCN Category: Data Deficient.

Measurements Graphiurus johnstoni HB: 74.3 (69–84) mm, n = 3 T: 68.5 (65–75.5) mm, n = 3 HF: 16 (15–17) mm, n = 4 E: 11.8 (11–12) mm, n = 4 WT: n. d. GLS: 23.3, 23.9 mm, n = 2 GWS: 13.6, 14.1 mm, n = 2 P4–M3: 3.4 (3.3–3.5) mm, n = 3 S Malawi (M. E. Holden unpubl.). Key References Ansell 1989b; Ansell & Dowsett 1988; Happold & Happold 1989a, 1997. Mary Ellen Holden

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Family GLIRIDAE

Graphiurus kelleni KELLEN’S AFRICAN DORMOUSE Fr. Graphiure nain (Graphiure de Kellen); Ger. Kellens Bilch Graphiurus kelleni (Reuvens, 1890). Notes from the Leyden Museum 13: 74. Reuvens originally listed Damaraland, Namibia, as the type locality; this was later emended to ‘Damara-land’, Mossamedes district, Angola (Hill and Carter 1941).

Taxonomy Originally described in genus Eliomys. Comparisons of museum specimens (including holotypes) and multivariate analyses of cranial morphology of G. kelleni, G. parvus and G. olga (M. E. Holden unpubl.) support the recognition of only one species (G. kelleni), and hence olga and parvus, previously considered as separate species (Holden 1993), are now placed as synonyms. These and other synonyms listed below have either been treated as separate species or included in species other than G. kelleni (see summary in Holden 1996, 2005). Ansell (1978) listed the species as G. johnstoni (recognized here as a separate valid species), but later (Ansell & Dowsett 1988, Ansell 1989b) referred to it as G. kelleni. The species requires taxonomic revision. Future studies will need to include larger samples, specimens from poorly collected localities and genetic data; such studies may well show that two or more separate species are contained within G. kelleni. No specimens of G. kelleni have been identified from South Africa (M. E. Holden unpubl.); however, some South African populations that are now placed in G. murinus are small in size. Future study and comparison of those populations may show that they are G. kelleni or an as yet undescribed species. Schlitter et al. (1985) discussed taxonomic problems and historical treatment of this species, and pointed out that G. kelleni is the oldest available scientific name. Synonyms: ansorgei, brockmani, cuanzensis, dollmani, Graphiurus kelleni foxi, internus, nanus, olga, parvus, personatus, tasmani. Subspecies: none. Chromosome number: 2n = 70 (Dobigny et al. 2002b). Geographic Variation Certain populations of G. kelleni exhibit variations in the colour and texture of dorsal pelage and in skull Description Small dormouse. Dorsal pelage various shades of morphology that are consistent within these populations; e.g. in brown, beige or grey, sometimes with golden or reddish hue, with Somalia, dorsal pelage is sleek and pale reddish-tan; in montane darkening of pelage towards the mid-line of the head and back in some populations in Kenya, dorsal pelage is thick and medium goldenindividuals. Dorsal pelage silky, sleek in some populations, thick in brown. Some skulls in certain populations are very delicately built, others (rump hairs 6–7 mm, guard hairs up to 11 mm). Ventral pelage others are robust; some are comparatively flat, others moderately usually white or cream, lightly or moderately suffused with grey. vaulted; some have long and inflated auditory bullae, others exhibit Dorsal and ventral pelage colours clearly delineated. Head colour short and only moderately inflated auditory bullae relative to skull usually matches that of dorsal pelage, sometimes paler towards muzzle. length. On the basis of predominantly small sample sizes, and Eyes large; eye-mask conspicuous. Ears brown, medium or large, inadequate sampling over the vast geographic range of this species, rounded. Cheeks cream or white, forming part of a pale lateral stripe preliminary morphometric analyses support the recognition of just a that extends from cheeks to shoulders. Cream or white postauricular single species without subspecies. patches usually present. Hindfeet white, or white with dark metatarsal streak. Tail moderately long (ca. 82% of HB), tail hairs shorter at base Similar Species (size comparisons refer to mean values only) (2–3 mm) and longer at tip (up to 20 mm). Tail appears splayed in G. microtis. Larger head and body length (mean 98.8 mm); ears some populations (particularly in Angola, Zimbabwe and Zambia) longer (mean 15.5 mm). Dorsal pelage colour sometimes similar. because hairs project laterally. Dorsal tail colour matches that of dorsal Tail usually has white hairs mixed throughout, and conspicuous pelage, often laterally fringed with white hairs and with faint or white tip. Skull longer (27.4 mm) and broader (15.0 mm). conspicuous white at tip. Ventral tail colour usually paler than dorsal Anterior palatal foramina 3.4 mm long, 2.1 mm wide. Breadth of tail colour. Skull short (24.0 mm), moderately vaulted (height of interorbital constriction (3.9 mm) and length of upper toothrow braincase 7.0 mm) and moderately broad (13.5 mm), sometimes (3.0 mm) absolutely similar so relatively smaller in G. microtis. gracile. Interorbital constriction moderately narrow (4.0 mm), Auditory bullae absolutely slightly longer (8.1 mm), or of similar anterior palatal foramina relatively long (2.9 mm) and auditory bullae length in some populations, but relatively shorter. Sympatric in long (7.8 mm) and inflated relative to skull length (measurements savannas throughout much of sub-Saharan Africa. listed are mean values from Zambezi [formerly Balovale], Zambia ; M. G. murinus. Larger head and body length (mean 91.5 mm). Hindfeet E. Holden unpubl.). Nipples: 1 + 1 + 2 = 8. (18.5 mm) longer, ears (13.3 mm) absolutely and relatively 116

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shorter. Dorsal pelage colour sometimes similar. Skull longer (26.4 mm), more vaulted (height of braincase 8.1 mm) and broader (14.2 mm). Interorbital constriction slightly broader (4.5 mm) and upper toothrow slightly longer (3.1 mm). Anterior palatal foramina 3.1 mm long, 1.9 mm wide. Auditory bullae shorter (7.1 mm) and less inflated relative to skull length (measurements listed are mean values from Mt Kenya; M. E. Holden unpubl.). Generally not sympatric, although they have been captured in close proximity at several Kenyan localities. Occurs in forests throughout much of sub-Saharan Africa. Distribution Endemic to Africa. Recorded from parts of Sahel Savanna, Sudan Savanna, Guinea Savanna and Somalia–Masai Bushland BZs; also Zambezian Woodland BZ as far south as Angola, Malawi, Zimbabwe and Mozambique at altitudes up to 1524 m (Holden, 2005). Not recorded from Namibia, Botswana and South Africa. Habitat Woodland savanna, riverine woodland, rocky areas including caves, disturbed areas and human dwellings. Specimens have been captured in or near dom palms (Hyphaene thebaica), thorn trees (Acacia spp.) and in miombo (Brachystegia) trees. Also occurs on mountains in East Africa (up to at least 1524 m), in rocky areas, and in caves (Osgood 1910, Dollman 1912, Hollister 1919; specimen labels). Less commonly found in disturbed areas, such as woodpiles, corn fields and in human dwellings (de Winton 1896, Stanley et al. 2002; specimen labels). Abundance Little information. Comprised 1.7% of muroid and glirid rodents captured in savanna at Foro, Côte d’Ivoire, and 3.5% of muroid and glirid rodents captured in Guinea savanna in C Côte d’Ivoire (Gautun et al. 1991). Specimen labels from most localities indicate that the species is uncommon, but very large series from certain localities (e.g. C Angola, NE Zambia) indicate that it is common in at least certain parts of its range. No estimates of population density. Adaptations Arboreal and nocturnal. These dormice frequently nest in crevices under bark, or in holes in savanna trees. Two nest holes were 0.5 m and 1 m above ground (specimen labels), and one nest was made of leaves and grass (Hill 1941). Several individuals were caught in nests of weaver-birds on Acacia trees and in the mud nests of swallows under roofs of caves or on undersides of rocks (Hollister 1919; specimen labels). Reported to utilize abandoned spider (Stegodyphys sp.) nests (Roberts 1951; specimen label) and abandoned beehives (Hill 1941). A few individuals have been caught in wood piles, in roofs of African huts and in pantries (Lawrence & Loveridge 1953; specimen labels).

Foraging and Food Probably omnivorous. In Somalia, one individual was caught in a trap baited with fresh meat (specimen label). Social and Reproductive Behaviour Little information. Males are apparently solitary. Lactating !! are often caught with young (specimen labels). Vocalizations of this dormouse have frequency components that range from ca. 1 kHz to well into the ultrasonic range above 20 kHz. Most vocalizations (termed ‘kecker/ shrieks’ and recorded from both sexes) seem to occur during agonistic behaviour (Hutterer & Peters 2002). Vocalizations characterized as ‘twitters’ also recorded from both sexes in non-aggressive closerange situations. Reproduction and Population Structure Litter-size: 2–4 (Hollister 1919, Hill 1941; specimen labels). Young individuals and lactating !! found in many months of the year throughout the range; however, paucity of information does not allow any conclusions on reproductive seasons or reproductive strategy. The scattered information includes: Senegal, young in Jul; one young in Aug (Côte d’Ivoire); young in Apr (Benin); pregnant !! in Nov (Hollister 1919) and Dec, and young in Nov and Apr (Kenya); lactating !! in Dec and young in Apr (Zimbabwe); lactating !! in Sep and Oct, and young in Jan (NW Zambia); and young in Oct (Angola) (specimen labels). Predators, Parasites and Diseases Principal host for the hoplopleurid louse Schizophthirus graphiuri (Durden & Musser 1994, Pajot 2000). Conservation

IUCN Category: Least Concern.

Measurements Graphiurus kelleni HB: 82.4 (75–92) mm, n = 14 T: 68.3 (54–81) mm, n = 13 HF: 16.0 (15.3–16.5) mm, n = 14 E: 14.8 (14–16) mm, n = 14 WT: 19.1 (10.9–23.5) g, n = 8* GLS: 24.0 (23.1–24.5) mm, n = 13 GWS: 13.5 (12.9–14.1) mm, n = 14 P4–M3: 2.9 (2.8–3.0) mm, n = 13 Body and skull measurements: Zambezi (Balovale), Zambia (M. E. Holden unpubl.) *Zimbabwe (M. E. Holden unpubl.) Key References

Ansell 1989b; Holden 2005; Hollister 1919. Mary Ellen Holden

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Graphiurus lorraineus LORRAINE’S AFRICAN DORMOUSE Fr. Graphiure de Lorrain; Ger. Lorraines Bilch Graphiurus lorraineus Dollman, 1910. Ann. Mag. Nat. Hist., ser. 8, 5: 285. Molegbwe, south of the Setema Rapids, Welle (Uele) River, DR Congo.

Taxonomy The named form lorraineus, originally described as a valid species, has been considered historically as a subspecies or synonym of G. murinus (Rosevear 1969, Eisentraut 1973, Genest-Villard 1978a), or as a valid species (Hatt 1940a, Robbins & Schlitter 1981, Holden 1993, 1996, 2005). Populations that are here placed in G. lorraineus (haedulus from Cameroon, spurrelli from Ghana) have never been thoroughly analysed and compared to the population of G. lorraineus from E DR Congo (see Geographic Variation). Future studies may show that one or more populations of what now comprises G. lorraineus merits recognition as one or more separate valid species.A comparative study of museum specimens and preliminary multivariate analyses (M. E. Holden unpubl.) showed that G. lorraineus appears to represent a valid species distinct from but closely related to G. johnstoni. If future research indicates that the two species are conspecific, G. lorraineus would become a junior synonym of G. johnstoni. Synonyms: haedulus, spurrelli. Subspecies: none. Chromosome number: 2n = 70 (an individual from Côte d’Ivoire identified as G. murinus (Tranier & Dosso 1979) but probably represents G. lorraineus). Description Small dormouse. Dorsal pelage reddish-brown, occasionally sandy or golden-brown. Dorsal pelage soft and short (rump hairs 5–6 mm, guard hairs up to 9 mm). Ventral pelage dark grey washed with cream or ochre, or mostly cream. Dorsal and ventral pelage colours usually not clearly delineated. Head colour matches that of dorsal pelage. Eyes large; eye-mask conspicuous in some individuals. Ears brown, short and rounded. Cheeks dark grey washed with cream or ochre, or predominantly cream. Postauricular patches usually not present, but white postauricular patches exhibited by some individuals from Cameroon. Hindfeet usually white with dark metatarsal streak. Tail moderately long (ca. 79% of HB), tail hairs shorter at base (2–3 mm) and longer at tip (up to 21 mm). Tail appears splayed because the hairs project laterally. Tail colour generally matches that of dorsal pelage and is uniform in colour, with sparse or no white hairs, and usually without white tip. Tail may be shorter than normal due to injury, ending in a thick, brush-like tuft of hairs (white in colour). Skull short (24.5 mm), broad (13.8 mm) and moderately vaulted (height of braincase 7.2 mm). Interorbital constriction (4.3 mm) relatively broad. Anterior chamber of auditory bullae markedly less inflated than posterior chambers in some individuals. Rostrum relatively short with short nasal bones (9.1 mm). Anterior palatal foramina comparatively short (mean 2.6 mm ) and narrow (mean 1.7 mm), interorbital constriction moderately broad (4.3 mm), palate moderately long (7.8 mm), upper toothrow moderately long (3.1 mm) and auditory bullae long relative to skull length (measurements listed are mean values from DR Congo; M. E. Holden unpubl.). Nipples: 1 + 1 + 2 = 8. Geographic Variation The populations of G. lorraineus in E DR Congo (lorraineus) and in S Cameroon (haedulus) are smaller than those from Ghana (spurrelli). Montane populations in Cameroon (haedulus) exhibit morphological and ecological differences compared with lorraineus and spurrelli (see Habitat).

Similar Species (size comparisons refer to mean values only) Graphiurus christyi. Larger head and body length (mean 97.6 mm), with longer hindfeet (mean 18.0 mm), and longer ears (mean 14.2 mm). Dorsal pelage usually golden-brown or greyishbrown. Skull (28.0 mm) longer, with elongate, higher rostrum and longer nasal bones (10.5 mm); nasal bones and premaxilla extend well beyond incisors from lateral and ventral view. Anterior palatal foramina 3.0 mm long, 2.2 mm wide. Palate absolutely and relatively longer (8.5 mm). Interorbital constriction absolutely broader (4.7 mm), but narrower relative to skull length. Upper toothrow similar in absolute length (3.2 mm), but relatively shorter. Auditory bullae average slightly longer (7.4 mm), but are relatively shorter (measurements listed are mean values from DR Congo; M. E. Holden unpubl.). Sympatric in NE DR Congo and SW Cameroon. Occurs in NE DR Congo and SW Cameroon. Graphiurus crassicaudatus. Larger head and body length (mean 92.6 mm). Dorsal pelage similar in colour. Skull longer (26.6 mm) and broader (16.1 mm) with absolutely and relatively broader interorbital constriction (4.9 mm). Anterior palatal foramina absolutely similar in length (2.5 mm) and breadth (1.6 mm), but relatively shorter and narrower. Auditory bullae shorter (6.7 mm) relative to skull length. Palate (9.4 mm) and upper toothrow (3.8 mm) absolutely and relatively longer (measurements listed are mean values from S Cameroon; M. E. Holden unpubl.). Sympatric at several localities in Liberia, Côte d’Ivoire and SW Cameroon. Occurs in W and WC Africa. Graphiurus surdus. Larger head and body length (mean 99.0 mm), hindfeet (mean 20.8 mm) longer but relatively similar. Ears similar in absolute length (12.3 mm), but relatively shorter. Dorsal pelage greyish-brown. Skull (27.6 mm) longer; interorbital constriction absolutely broader (4.5 mm), but narrower relative to skull length. Zygomatic arch straight in lateral view (figured in Holden 1996). Palate absolutely and relatively longer (9.3 mm). Anterior palatal foramina similar in length (2.8 mm) and breadth (1.8 mm). Upper toothrow (3.2 mm) and auditory bullae (7.3 mm) similar in absolute length, but relatively shorter (measurements listed are mean values from Cameroon, Equatorial Guinea and Gabon; Holden 1996). Sympatric in SW Cameroon, Equatorial Guinea and DR Congo. Occurs in C Africa. Graphiurus johnstoni: Smaller head and body length (mean 74.3 mm), ear (mean 11.8 mm) slightly smaller. Tail relatively longer (68.5 mm) with similar colouration. Dorsal pelage colour similar. Skull slightly shorter (23.6 mm). Rostrum relatively short as in G. lorraineus, but length of nasal bones absolutely shorter (8.4 mm). Anterior palatal foramina absolutely similar in length (2.7 mm) and breadth (1.7 mm), but relatively longer. Upper toothrow slightly longer (3.4 mm) and upper premolar broader (1.0 mm). Not sympatric. Occurs in S Malawi.

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Graphiurus lorraineus

Distribution Endemic to Africa. Rainforest, Guinea Savanna and northern part of Zambezian Woodland BZs, and Rainforest– Savanna Mosaics. Recorded from Guinea to Cameroon, Central African Republic, Congo, E DR Congo, NE Angola and N Zambia; also Bioko I. (see Holden 2005). Not recorded from Togo, Benin and Nigeria west of Niger R. Specimens recorded from Gambia, originally thought to be this species (Schlitter et al. 1985) are now considered as G. kelleni (Grubb et al. 1998, Holden 2005). The western distributional limit for this species is Sierra Leone (Holden 1993, Grubb et al. 1998); specimens identified as ‘Graphiurus murinus’ (presumably spurrelli) from wooded savanna in Senegal (Hubert et. al. 1973) cannot be substantiated, and are not included here. Habitat Gallery forests, forest margins, woodland savanna and disturbed areas (including banana, cocoa, palmyra (Borassus) and pawpaw farms, and occupied and abandoned buildings) (Hatt 1940a, Heim de Balsac 1967a, Rosevear 1969, Jeffrey 1973, Robbins & Schlitter 1981, Schlitter et al. 1985, Grubb et al. 1998). In Côte d’Ivoire, Dosso (1975a) noted that the plant Microdesmis, commonly found in secondary and disturbed forest, was associated with areas where G. lorraineus was caught. These dormice were never trapped in, and seemed to avoid, primary forest (Hatt 1940a, Jeffrey 1973). In Cameroon, Eisentraut (1963; and specimen labels) trapped individuals in montane forest at altitudes of 1700–2100 m, but these populations (identified as haedulus) may represent a different species (see discussion under Taxonomy). Abundance Relatively common compared with other forest dormice such as G. crassicaudatus and G. surdus (Hatt 1940a, Heim de Balsac & Lamotte 1958, Heim de Balsac 1967a, Rosevear 1969, Gautun et al. 1986). Comprised 0.77% of muroid and glirid rodents captured at Lamto, Côte d’Ivoire (Dosso 1975a), although in a later survey at Lamto they comprised 7.6% of captures (Traore et al. 1980). At Foro, Côte d’Ivoire, comprised 2.1% of rodent captures (Traore et al. 1980).

Adaptations Arboreal, although some individuals seem to spend much time on the ground (Rosevear 1969). Nocturnal. Distribution seems to depend on the presence of suitable nesting sites, such as cavities in trees in gallery forest and isolated savanna trees near forest, even in disturbed areas (Verheyen & Verschuren 1966, Rosevear 1969). Many individuals have been caught in or near occupied or abandoned buildings (Verheyen & Verschuren 1966, Rosevear 1969, Jeffrey 1973, Robbins & Schlitter 1981; specimen labels). In C Africa, some individuals have been found nesting in rocky caves (Verheyen & Verschuren 1966). Others were found in abandoned nests of swallows (Hatt 1940a, Verheyen & Verschuren 1966). One such nest (Hatt 1940a) was occupied by an active nest of paper wasps, and the dormice had to crawl upside down on a nearly horizontal stone surface to enter it. Nests have also been found amongst epiphytic ferns (Verheyen & Verschuren 1966) and in a cocoa pod (Schouteden 1946). In Sierra Leone, individuals were caught in spherical nests constructed of pappus. Eisentraut (1963) caught specimens of montane populations in Cameroon in traps set 6–10 m high on large diagonal or horizontal branches near holes in trees; individuals were never trapped on the ground, another indication that this population may represent a different species. These dormice enter torpor under certain conditions. Lachiver & Petter (1969) found that individuals from Central African Republic became lethargic when experiencing sudden shifts from high to low ambient temperature, or when deprived of food at low temperature. Eisentraut (1962) could not induce torpor in individuals from Cameroon. Foraging and Food Probably omnivorous, consuming fruit, insects, seeds and nuts. In Liberia, Central African Republic and DR Congo, individuals have been caught in banana plantations, where they reportedly ate the fruit (Hatt 1940a, Chippaux & Pujol 1964, Coe 1975). They have also been caught in areas where palmyra (Borassus) (Heim de Balsac 1967a), pawpaw (Jeffrey 1973), Microdesmis (Dosso 1975a), cassava, cocoa, oil palm, plantains, Raphia and yams are common (specimen labels). The type specimen of haedulus was noted to have been ‘caught in bushes eating seeds of Piper subpellatum’, a species of pepper (specimen label). Four specimens were taken from a nest containing the remains of several hundred earwigs (Hatt 1940). Verheyen & Verschuren (1966) saw an individual running and jumping after termites, finally capturing them mostly in mid-air. Social and Reproductive Behaviour Lactating !! are often caught with young. In DR Congo, one adult ! was nesting in an old double nest of a swallow with her three ‘well-grown young’ (Hatt 1940a), indicating that offspring may stay in the nest past weaning. An adult ", in captivity, was observed to be very lively and aggressive. It moved its tail up and down with the hairs spread wide, and when excited would chatter‘gak gak’ repeated four or five times in succession. These dormice reportedly bite ‘furiously’ (H. Lang in Hatt 1940a). Reproduction and Population Structure Litter-size: 2–7. Mostly 2–4 nestlings or embryos are reported (Hatt 1940a, Eisentraut 1963, Jeffrey 1973). In Côte d’Ivoire, Heim de Balsac (1967a) captured one ! with six naked young, and another with a litter of seven (months of capture not given). These two litters are the largest recorded for this species. In Ghana and Cameroon, 119

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pregnant !! have been collected in Jan, Mar and Jul (Eisentraut 1963, Jeffrey 1973). In Ghana, a lactating ! with three placental scars was captured in Nov (Jeffrey 1973). Predators, Parasites and Diseases Predators include snakes and owls. Remains have been found in the stomach of a green mamba Dendraspis viridis (Jeffrey 1973) and in the pellet of a Barn Owl Tyto alba (Heim de Balsac & Lamotte 1958). Conservation

IUCN Category: Least Concern.

T: 65.7 (54–74) mm, n = 16 HF: 16.5 (14–19) mm, n = 20 E: 12.6 (9–15) mm, n = 18 WT: 16.8 (12–24) g, n = 5 GLS: 24.5 (22.7–26.1) mm, n = 21 GWS: 13.8 (12.2–14.9) mm, n = 15 P4–M3: 3.1 (2.8–3.4) mm, n = 29 DR Congo (M. E. Holden unpubl.) Key References Eisentraut 1963; Grubb et al. 1998; Hatt 1940a; Robbins & Schlitter 1981; Rosevear 1969.

Measurements Graphiurus lorraineus HB: 83.0 (72–93) mm, n = 19

Mary Ellen Holden

Graphiurus microtis NOACK’S AFRICAN DORMOUSE Fr. Graphiure de Noack; Ger. Noack’s Bilch Graphiurus microtis (Noack, 1887). Zoologische Jahrbücher 2: 248. Qua Mpala, Marungu, DR Congo.

Taxonomy Originally described in genus Eliomys. The type specimen of G. microtis has been lost, but Noack’s (1887) figure depicts a dormouse whose skull morphology agrees with specimens of G. microtis from regions near the type locality (Holden 2005).The taxon microtis has historically been listed as a synonym or valid subspecies of G. murinus (Allen 1939, Ellerman et al. 1953, Genest-Villard 1978a). Genest-Villard (1978a) separated what she considered to be the‘savanna subspecies’ of G. murinus from the ‘forest subspecies’, and considered G. m. microtis to be one of several valid savanna subspecies. Ansell (1989a) agreed, but considered that the three savanna subspecies comprised a single valid species, G. microtis, on the basis of morphological and ecological differences, a position endorsed by Holden (1993, 2005). Holden (1996) summarized the historical taxonomic arrangements of G. murinus and other species of Graphiurus. Many of the 76 specific and subspecific names proposed for African Dormice are synonyms of G. murinus or G. microtis. Because G. microtis is now recognized as a valid species (Ansell & Dowsett 1988, Ansell 1989a, Holden 1993, 2005), synonyms associated with it have traditionally been listed as synonyms of G. murinus. Holden (1993) did not separate the synonyms associated with G. murinus and G. microtis, but the taxonomy presented here results from further examination of specimens, data and multivariate analyses (M. E. Holden unpubl.). Historically, many authors have not recognized G. microtis as a valid species, and the data given in their papers are thus composite for both species; when such publications are cited here, only the sections relevant to G. microtis as outlined in this account are pertinent. The species requires taxonomic revision; significant geographic variation exists, and it is likely that two or more separate species are contained within G. microtis. Synonyms: albolineata, butleri, etoschae, griselda, littoralis, marrensis, ?orobinus, pretoriae, schneideri, smithii, streeteri, sudanensis, tzaneenensis, vandami, woosnami (Holden 2005). Subspecies: none. Chromosome number: 2n = 46 (Transvaal, South Africa; D. N. MacFadyen pers. comm.; see species profile for G. murinus for discussion of karyotypes). Description Small dormouse. Dorsal pelage various shades of brown, beige or grey, sometimes with golden or reddish hue, with

darkening towards the mid-line of head and back in some individuals. Dorsal pelage usually sleek, but moderately thick in some populations (rump hairs 6–8 mm, guard hairs up to 13 mm). Ventral pelage usually white or cream, slightly or moderately suffused with grey. Dorsal and ventral pelage colours clearly delineated. Head colour usually matches that of dorsal pelage, sometimes becoming paler towards muzzle. Eyes large; eye-mask conspicuous. Ears brown, medium or large, rounded. Cheeks cream or white, forming part of a pale lateral stripe that extends from cheeks to shoulders. Cream or white postauricular patches usually present. Hindfeet white, or white with dark metatarsal streak. Tail moderately long (ca. 76% of HB), hairs shorter at base (5–8 mm) and longer at tip (up to 26 mm). Tail colour generally matches that of dorsal pelage.White hairs are usually mixed throughout tail; tip white. Skull moderately long (27.4 mm) with slightly to moderately vaulted braincase (height of braincase 7.5 mm), and relatively long anterior palatal foramina (3.4 mm). Anterior palatal foramina moderately long (mean 3.4 mm) and wide (mean 2.1 mm). Interorbital constriction narrow (3.9 mm) and auditory bullae usually long (8.1 mm) and inflated relative to skull length. Nipples: 1 + 1 + 2 = 8. Geographic Variation Certain populations of this dormouse exhibit variations in the colour of the dorsal pelage and in skull morphologies that are consistent within these populations; e.g. in C Botswana, dorsal pelage colour is ash-grey and skulls tend to be long and comparatively flat, with a narrow interorbital constriction. In nearby Namibia, dorsal pelage colour is also grey, but skulls are shorter and more vaulted, with a broader interorbital constriction. Other populations, e.g. certain populations in Uganda and Sudan, have sandy- to medium-brown dorsal pelage, a long and comparatively vaulted skull with long anterior palatal foramina, broad interorbital constriction, and greatly inflated auditory bullae. Differences like these are found throughout the range of G. microtis. Some populations have such distinctive morphologies that future studies may show that one or more of them is a separate valid species.

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Graphiurus microtis

Similar Species (size comparisons refer to mean values only) Graphiurus angolensis: Similar head and body length (mean 98.8 mm). Tail (mean 79.2 mm) and hindfeet (mean 18.4 mm) absolutely and relatively longer. Dorsal pelage sometimes similar in colour. Skull similar in length (28.2 mm), but slightly broader (15.5 mm) and more vaulted (height of braincase 7.9 mm). Auditory bullae markedly longer (9.0 mm) and more inflated relative to skull length (measurements listed are mean values from Kabompo and Zambezi [formerly Balovale], Zambia; M. E. Holden unpubl.). Parapatric in NW Zambia (Ansell 1978). Occurs in Angola and NW Zambia. Graphiurus johnstoni: Smaller head and body length (mean 74.3 mm). Tail relatively longer (mean 68.5 mm). Dorsal pelage colour sometimes similar, tail colour usually uniform and without white tip. Skull shorter (23.6 mm) and narrower (13.9 mm). Interorbital constriction (4.0 mm) similar in breadth, but relatively broader. Anterior palatal foramina absolutely and relatively shorter. Upper toothrow (3.4 mm) longer and auditory bullae (6.8 mm) shorter both absolutely and relative to skull length (measurements listed are mean values from S Malawi; M. E. Holden unpubl.). Sympatric in Thyolo, Malawi, but probably not syntopic. Currently known to occur only in S Malawi. Graphiurus kelleni: Smaller head and body length (mean 82.4 mm); ears shorter (mean 14.8 mm). Dorsal pelage colour sometimes similar.Tail usually more uniform in colour, with inconspicuous white tip. Skull shorter (24.0 mm) and narrower (13.5 mm). Breadth of interorbital constriction (4.0 mm) and length of upper toothrow (2.9 mm) absolutely similar, so relatively larger in G. kelleni. Auditory bullae somewhat shorter (7.8 mm) both absolutely and relative to skull length (measurements listed are mean values from Zambezi, Zambia; M. E. Holden unpubl.). Sympatric in savannas throughout much of subSaharan Africa. Graphiurus murinus: Similar head and body length (mean 91.5 mm). Hindfeet (18.5 mm) longer, and ears (13.3 mm) shorter. Dorsal pelage colour sometimes similar. Postauricular patches inconspicuous or not present. Ventral pelage colour usually greyer and not clearly delineated from dorsal pelage. Hindfeet usually have dark metatarsal streak. Tail usually uniform in colour, white tip inconspicuous or absent. Skull similar in length (26.4 mm), but slightly narrower (14.2 mm) and more vaulted (height of braincase 8.1 mm). Anterior palatal foramina absolutely slightly shorter (3.1 mm), but relatively shorter. Interorbital constriction broader (4.5 mm), and auditory bullae shorter (7.1 mm) and less inflated relative to skull length (measurements listed are mean values from Mt Kenya; M. E. Holden unpubl.). Not sympatric. Occurs in forests throughout much of sub-Saharan Africa. Graphiurus platyops: Larger head and body length (mean HB: 107.1). Tail (mean 78.7 mm) absolutely longer, but relatively shorter. Hindfeet (21.1 mm) longer, and ears (15.2 mm) shorter, both absolutely and relatively. Dorsal pelage grey to greyish-brown. Inconspicuous white postauricular patches sometimes present. Tail tip white. Skull longer (30.4 mm) and broader (17.1 mm). Height of braincase (7.8 mm) similar, so relatively flattened in G. platyops. Interorbital constriction (4.8 mm) absolutely and relatively broader. Anterior palatal foramina absolutely similar in length (3.2 mm), but relatively shorter. Upper toothrow

(3.1 mm) and auditory bullae (8.4 mm) similar to slightly longer, so are relatively shorter (measurements listed are mean values from Zimbabwe and NE South Africa populations; M. E. Holden unpubl.). Generally sympatric throughout range of G. platyops, but seem to be segregated by habitat, and are probably not syntopic. Occurs in rocky habitats in eastern and southeastern Africa. Graphiurus rupicola: Slightly larger head and body length (mean 110 mm). Tail (104.2 mm). Hindfeet (21.5 mm) absolutely and relatively longer. Dorsal pelage colour sometimes similar in colour, especially in animals from Namibia. Small, cream supraauricular patches and faint white postauricular patches present. Tail tip white. Skull markedly longer (31.3 mm) and most skull measurements are accordingly larger. Upper toothrow (3.4 mm) absolutely longer, but relatively similar. Upper premolar (0.9 mm) absolutely slightly broader, but relatively narrower. Anterior palatal foramina absolutely similar in length (3.4 mm), but relatively shorter. Auditory bullae (9.5 mm) longer both absolutely and relative to skull length (measurements listed are mean values from Namibia; M. E. Holden unpubl.). Sympatric near Okahandja, Namibia. Occurs in rocky habitats in Angola, Namibia and NW South Africa. Distribution Endemic to Africa. Widespread in Zambezian Woodland BZ, with extensions into parts of Eastern Rainforest– Savanna Mosaic, Guinea Savanna and Sudan Savanna BZs on the eastern side of the continent. Recorded from savannas in Chad, Sudan, Eritrea and Ethiopia south to Namibia and NE South Africa (Holden 2005). Late Quaternary fossils of G. microtis have been recorded from C Zambia (Avery 1996). Habitat Woodland savanna, riverine woodland, rocky areas including caves, disturbed areas, and human dwellings. These dormice have been captured in or near aloes, willows, upaca trees

Graphiurus microtis

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(Upaca kirkiana), thorn trees (Acacia sp.), camel thorn (Acacia erioloba (giraffae)), Zambezian teak or mukusi (Baikiaea sp.), mopane (Colophospermum mopane), leadwood (Conbretum imberbe), seringa (Burkea sp.), palm (Hyphaene sp.) and buffalo thorn (Ziziphus mucronata) (Shortridge 1934, Misonne 1965a, Misonne & Verschuren 1966, Smithers 1971, Wilson 1975, Smithers & Wilson 1979, Ansell & Dowsett 1988; specimen labels). They have been captured in tall grass near shrubs and trees, and in piles of debris deposited by high floods near seasonally dry rivers (Smithers 1983). Individuals have been observed on a vertical rock face at the entrance of a cave situated on a rocky hillside (Ansell 1974), and captured among rocks in or near caves (Roberts 1917, Misonne 1965a, Misonne & Verschuren 1966). Common also in disturbed areas, including buildings, fields, gardens and near rubbish dumps (Wilson 1975, Smithers & Lobão Tello 1976, Smithers & Wilson 1979, Ansell & Dowsett 1988, Taylor et al. 1994). When evaluating the range of microhabitats tolerated by these dormice, it is important to consider that more than one valid species is probably contained within G. microtis (see Taxonomy). Abundance Common. Although published faunal surveys have not estimated population densities (due to the commonly mistaken inclusion of this species in G. murinus), the large number of museum specimens, and notes written on specimen labels, indicate that it is common throughout much of its range. In Malawi, Ansell & Dowsett (1988) found it to be the most frequently encountered dormouse, and stated that it is widely distributed. Adaptations Primarily arboreal, partly terrestrial (Smithers & Wilson 1979, Smithers 1983). Nocturnal (Smithers 1971, 1983, Wilson 1975, Smithers & Wilson 1979. These dormice frequently nest in crevices under bark, or in holes in savanna trees (Shortridge 1934, Smithers & Lobão Tello 1976, Smithers 1983). The entrance to most nesting holes is circular, and is commonly situated 1–3 m above ground, although some have been found at heights of up to 6 m (Shortridge 1934; specimen labels). Nests are composed of soft plant material or grass, and sometimes feathers (Roberts 1951; specimen labels). They also nest in rocky habitats. One individual was observed at a cave entrance easily negotiating a vertical rock face (Ansell 1974), another was captured among rocks near a cave (Misonne 1965a). Several individuals nested in aloes in rocky terrain (Smithers & Lobão Tello 1976). They also utilize the nests of birds; one individual made its nest inside the nest of a swallow Hirundo abyssinica under a rock; the dormouse’s nest contained feathers, wood debris, grass and scales from a snake (Misonne 1965a). Several adults and young have been found in nests of various species of weaverbirds (Roberts 1951). Nests may also be built in huts and houses, often in thatched roofs, sometimes in pantries or even in switch boxes of water pumps or transformers where they have caused short circuits in electrical supplies (Wilson 1975, Smithers & Lobão Tello 1976, Smithers 1983, specimen labels).

Foraging and Food Probably omnivorous, consuming fruit, insects, seeds, nuts and occasionally small vertebrates. Stomach contents have included dry outer skins of buffalo thorn fruit Ziziphus mucronata, seeds of Acacia sp., insects – including large moths, rose beetles, millipedes Doratogonus flavifilis, but not blister beetles nor Hemiptera (Misonne 1965a, Smithers 1971, 1983, Smithers & Wilson 1979, Pienaar et al. 1980). The stomach of one individual contained the remnants of a small bird (specimen label). Social and Reproductive Behaviour Little information. Males are apparently solitary. Lactating !! are often caught with young. Reproduction and Population Structure Litter-size: 3–7 (specimen labels). Most often 3–4 embryos or nestlings are reported (Ansell 1974, Wilson 1975, Smithers & Wilson 1979, Sheppe & Haas 1981, specimen labels). In Uganda, a pregnant ! was captured in Nov, and two lactating !! were captured in Aug. In Malawi, a pregnant ! was collected in Oct (Ansell 1974). In Botswana, a pregnant ! was obtained in Apr (Sheppe & Haas 1981). In Zimbabwe, pregnant !! have been collected in Feb, Apr, Jun, Nov and Dec (Wilson 1975, Smithers & Wilson 1979). Predators, Parasites and Diseases One individual was found in the stomach of a mamba Dendroaspis sp. (specimen label). Close to 20 individuals were identified in owl pellets collected in Kruger N. P., South Africa (M. E. Holden unpubl). These dormice are the type and a principal host for the trypanosome Trypanosoma graphiuri (Dekeyser 1955). They may host the same ectoparasites as G. murinus (see species profile). Conservation

IUCN Category: Least Concern.

Measurements Graphiurus microtis HB: 98.8 (75–115) mm, n = 33 T: 75.2 (62–86) mm, n = 28 HF: 16.9 (14–20) mm, n = 33 E: 15.5 (13–21) mm, n = 31 WT: 29.5 (17.6–42.5) g, n = 21 GLS: 27.4 (25.5–29.1) mm, n = 28 GWS: 15.0 (13.9–16.2) mm, n = 21 P4–M3: 3.0 (2.9–3.4) mm, n = 34 Zimbabwe (M. E. Holden unpubl.). Key References Ansell & Dowsett 1988; Holden 2005; Misonne 1965a; Smithers 1983; Smithers & Lobão Tello 1976; Smithers & Wilson 1979. Mary Ellen Holden

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Graphiurus monardi

Graphiurus monardi MONARD’S AFRICAN DORMOUSE Fr. Graphiure de Monard; Ger. Monards Bilch Graphiurus monardi (St Leger, 1936). Ann. Mag. Nat. Hist., ser. 10, 17: 465. Kioko, 15 km above Dala, Tyihumbwe (Chiumbe) River, Angola. 1250 m (see Hill & Carter 1941).

Taxonomy Originally described in the genus Claviglis. Allen (1939) included monardi as a subspecies of nagtglasii, and GenestVillard (1978a) followed this arrangement. Ellerman et al. (1953), Robbins & Schlitter (1981) and Holden (1993, 2005) agree that monardi has no close affinity with G. nagtglasii. Genest-Villard (1978a) included schoutedeni as a synonym of G. platyops, but Ansell (1989a) correctly arranged it as a synonym of G. monardi. Synonyms: schoutedeni (Ansell 1989a). Subspecies: none. Chromosome number: not known. Description Large dormouse. Dorsal pelage yellowish-brown, darkening towards mid-line due to a higher density of guard hairs in that region. Dorsal pelage sleek and long (rump hairs 11–13 mm, guard hairs up to 17 mm), with many conspicuous dark brown guard hairs projecting beyond the fur. Ventral pelage cream, sometimes lightly suffused with grey. Dorsal and ventral pelage colours clearly delineated. Head colour matches that of dorsal pelage. Eyes large; eye-mask usually conspicuous. Ears brown, relatively small, rounded. Cheeks cream. Cream postauricular patches sometimes present. Hindfeet cream, sometimes with a thin, dark metatarsal streak. Tail moderately long (ca. 81% of HB), hairs shorter at base (6–9 mm) and longer at tip (up to 33 mm). Dorsal tail colour generally matches that of dorsal pelage, but ventral tail colour paler brown. Many white hairs are mixed throughout tail; tip white or cream. Skull long (34.1 mm) and vaulted (height of braincase 9.4 mm). Interorbital constriction narrow (mean 5.0 mm), anterior palatal foramina very long (4.1 mm) and wide (2.6 mm), and palate short (mean 10.7 mm) relative to skull length. Upper toothrow moderately broad as exhibited by medium breadth of upper premolar (mean 1.1 mm). Auditory bullae (mean 10.3 mm) absolutely and relatively long and inflated Nipples: 1 + 1 + 2 = 8. Geographic Variation None recorded. Similar Species (size comparisons refer to mean values only) Graphiurus nagtglasii: This species is strikingly unlike G. monardi, but a comparison is given here due to previous inclusion of monardi in G. nagtglasii. Similar head and body length (mean 138.5 mm); most specimens of G. monardi have no data on head and body measurements, but based on specimen comparisons the size range of the two species is similar. Ear (mean 18.1 mm) and hindfoot (mean 26.5 mm) are absolutely and relatively longer. Dorsal pelage shorter and woolly, with inconspicuous guard hairs. Skull longer (mean 36.8 mm) with absolutely and relatively longer palate (mean 13.0 mm). Anterior palatal foramina shorter (3.7 mm) and narrower (2.3 mm) relative to skull length. Auditory bullae absolutely and relatively shorter (mean 7.9 mm) and much less inflated. Not sympatric. Graphiurus nagtglasii occurs in West Africa from Sierra Leone to Gabon.

Graphiurus monardi

Distribution Endemic to Africa. Zambezian Woodland BZ. Known from only seven localities in NE Angola, S DR Congo and NW Zambia. Habitat Central African savanna. No specific habitat information has been recorded for this dormouse. All specimens, except one from Katanga (DR Congo), have been taken on plateaux in wetter Miombo woodland. St Leger (1936) gives anecdotal information obtained from Dr Monard that this dormouse occurs in forest, as well as in cultivated fields and houses. Another species, G. angolensis, is also found in this region, and is known to frequent vacant and occupied buildings. It is likely that some, if not all, of the reports of this species in cultivated fields and buildings are attributable to G. angolensis. Abundance Little information. Only about a dozen museum specimens exist. Judging from the large numbers of specimens of dormice and other animals collected at some of the same localities, G. monardi appears to be rare. Remarks Probably predominantly arboreal. Hayman (1963b) includes photographs of a live animal climbing on a branch. Conservation

IUCN Category: Data Deficient.

Measurements Graphiurus monardi HB: 160 mm, n = 1* 123

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T: 130 mm, n = 1* HF: 21.9 (21.5–22) mm, n = 4 E: 15.5 mm (estimated from two dry specimens) WT: n. d. GLS: 34.1 (32.5–36.6) mm, n = 6 GWS: 19.7 (18.2–21.6) mm, n = 5 P4–M3: 3.9 (3.6–4.3) mm, n = 9

NE Angola, S DR Congo and NW Zambia (M. E. Holden unpubl.) *St Leger (1936) Key References Ellerman et al. 1953; Hayman 1963b; St Leger 1936. Mary Ellen Holden

Graphiurus murinus FOREST AFRICAN DORMOUSE Fr. Graphiure murin (Lérot de savanne); Ger. Busch-Bilch Graphiurus murinus (Desmarest, 1822). Mammalogie. In: Encyclop. Méth., 2 (Suppl.): 542. Cape of Good Hope, South Africa.

Taxonomy Originally described in genus Myoxus. Many of the 76 scientific names proposed for African Dormice are synonyms of G. murinus (which usually inhabits forests) or G. microtis (which usually inhabits savannas) (see Holden 1996). Historically, many authors have not recognized G. microtis as a valid species, and the data given in their papers are thus composite for both species; when such publications are cited here, only the sections relevant to G. murinus as outlined in this account are pertinent. The species needs taxonomic revision, requiring comprehensive comparisons with G. lorraineus, G. johnstoni, G. microtis and G. christyi (Holden 2005). Significant morphological geographic variation exists, and it is likely that two or more separate species are contained within G. murinus. Because the definition of this species is so uncertain, information on abundance, distribution and biology may be applicable to other species, as yet undescribed. Synonyms: alticola, cineraceus, cinerascens, collaris, erythrobronchus, griseus, isolatus, johnstoni Heller, 1912 (not Thomas, 1898), lalandianus, raptor, saturatus, selindensis, soleatus, ?subrufus, vulcanicus, zuluensis. Subspecies: none. Chromosome number: 2n = 46 (Natal, South Africa, D. N. MacFadyen pers. comm.; Hobbiton, South Africa, Kryštufek et al. 2004). Three different karyotypes were reported in the G. murinus species group in southern Africa (Dippenaar et al. 1983); however, G. microtis may have been included in this sample. Description Small dormouse. Dorsal pelage various shades of golden- or greyish-brown, sometimes with reddish or coppery hue, with darkening of pelage towards the mid-line of head and back in some individuals. Dorsal pelage soft, silky, sometimes thick (rump hairs 7–8 mm, guard hairs up to 13 mm). Ventral pelage grey, lightly suffused with white or cream. Dorsal and ventral pelage colours usually not clearly delineated. Head colour matches that of dorsal pelage. Eyes large; eye-mask conspicuous in some populations. Ears brown, medium-sized, rounded. Cheeks cream or white. Postauricular patches usually not present. Hindfeet usually white with dark metatarsal streak. Tail moderately long (ca. 84% of HB), tail hairs shorter at base (2–4 mm) and longer at tip (up to 21 mm). Tail colour generally uniform, matching that of dorsal pelage. White hairs are sometimes mixed inconspicuously throughout tail in some populations, tip usually not white although some populations exhibit very faint white tip. Skull moderately long (26.4 mm) with moderately vaulted braincase (height of braincase 8.1 mm). Anterior palatal foramina of moderate length (3.1 mm) and width (1.9 mm); anterior portion often narrower anteriorly, resulting in tear-drop

shape. Palate moderately long (4.4 mm) and auditory bullae usually short (7.1 mm) and uninflated or moderately inflated relative to skull length. Nipples: 1 + 1 + 2 = 8. Geographic Variation As with G. microtis, significant variation in external and skull morphology exists, and these differences are consistent within certain populations. Kryštufek et al. (2004) found that individuals from riverine forest near Grahamstown, South Africa, were significantly larger, and differed significantly in several cranial dimensions and proportions (weight, head and body, ear and length of PM4) than individuals from Afromontane forest at Hobbiton, South Africa (Kryštufek et al. 2004). Individuals from Ukinga Highlands, Tanzania, have smaller, broader skulls, with broader interorbital constriction and wider cheekteeth than individuals from nearby Uzungwa Mts and Rungwe Mts, Tanzania. Differences such as these are found throughout the range of G. murinus. Some populations have distinctive morphologies, and future studies may show that one (or more) of these populations is a separate species. Similar Species (size comparisons refer to mean values only) Graphiurus microtis. Similar head and body length (mean 98.8 mm). Hindfeet (mean 16.9 mm) shorter, ears (mean 15.5 mm) longer. Dorsal pelage colour similar to G. murinus in some populations. Postauricular patches often conspicuous. Ventral pelage colour usually paler and well delineated from dorsal pelage. Hindfeet usually white or cream. Tail usually with conspicuous white tip. Skull similar in length (27.4 mm), slightly broader (15.0 mm) and braincase less vaulted (height of braincase 7.5 mm). Anterior palatal foramina absolutely only slightly longer (3.4 mm), but relatively noticeably longer. Interorbital constriction narrower (3.9 mm) and auditory bullae longer (8.1 mm) and more inflated relative to skull length (measurements listed are mean values from Zimbabwe; M. E. Holden unpubl.). Not sympatric. Occurs in savannas throughout much of sub-Saharan Africa. Graphiurus kelleni. Smaller head and body length (mean 82.4 mm). Hindfeet (mean 16.0 mm) shorter, ears (mean 14.8 mm) absolutely and relatively larger. Dorsal pelage colour similar to G. murinus in some populations. Skull shorter (24.0 mm), narrower (13.5 mm) and braincase less vaulted (height of braincase 7.0 mm). Anterior palatal foramina 2.9 mm long, 1.7 mm wide. Interorbital constriction slightly narrower (4.0 mm), and auditory bullae longer (7.8 mm) and much more inflated relative to skull length (measurements listed are mean values from Zambia; M.

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Graphiurus murinus

Provinces, South Africa (Swanepoel 1988, Lynch 1983, 1989, Taylor 1998), but uncommon in Lesotho (Lynch 1994). Uncommon in four out of six montane forests in the Eastern Arc Mts, Tanzania (South Pare, West Usambara, East Usambara and Uluguru), and seemingly absent in two forests (Nguru and Udzungwa) (Stanley et al. 1998a, b); although Allen & Loveridge (1933) reported one individual from Udzungwa. At Chome Forest Reserve, Tanzania, comprised only 0.6% of small rodents (1 of 165) (Stanley et al. 1998a). Trap success (and assessment of abundance) is probably higher for traps set above ground than on the ground e.g. in Afromontane forest in South Africa, trap success was 2.3% above ground and 0.1% on the ground, and most individuals (94.5%) were captured in traps placed more than 0.5 m above ground (Kryštufek et al. 2004). Most published information derives from South African populations; more information is needed from populations throughout the range of the species to generally characterize its abundance.

Habitat Primarily a forest species. Recorded mostly at altitudes of 1000–4100 m, and occasionally at sea level in coastal forests. Very adaptable and found in many types of forest (Afromontane, plateau, riverine, coastal). Less commonly recorded from montane grassland with rocks, giant groundsel, or trees. Rarely recorded from dry scrub or thicket. When evaluating the range of microhabitats tolerated by these dormice, it is important to consider that more than one valid species is probably contained within G. murinus (see Taxonomy). For example, at Grahamstown, South Africa, these dormice were trapped only in riverine forest and not in nearby dry thicket (B. Kryštufek pers. comm.), whereas at other South African localities individuals were captured in dry thicket and among rocks (Taylor 1998). It may be that individuals trapped in habitats other than forest are members of populations that will later be considered separate species.

Adaptations Primarily arboreal, partly terrestrial; some populations rupicolous. Frequently nests in holes and crevices in forest trees (Roberts 1951; specimen labels). Nests have also been found among epiphytic ferns and mosses of giant forest trees (Allen & Loveridge 1933), in beehives (Kingdon 1974), in swallows’ nests, and less commonly, in human dwellings. Nests are often composed of strips of grass, bark and other material, which is finely shredded and formed into a round ball (Roberts 1951). A nest in NE South Africa was constructed of moss and lined with sheep’s wool (specimen label). On Mt Kilimanjaro, Tanzania, a globular nest was composed of grass and slips of banana fronds, and lined with fine grass; it was about 13 cm in diameter, with a hole in its side, and was situated about 1.5 m above ground in a bush (specimen label). Under certain conditions, will enter torpor (as do many species of glirids). If experimentally deprived of food at Ta = 25 °C, individuals initially decreased activity, but remained euthermic. At Ta = 10 °C, when deprived of food, the same individuals entered torpor with greater frequency during the day (Webb & Skinner 1996b). Cold temperatures (10 °C) and simulated winter photoperiod (10 hours light, 14 hours dark) also induced torpor; these periods of torpor exceeded 24 hours, suggesting hibernation or deep torpor occurs under these conditions (Ellison & Skinner 1991). The digestive system indicates a diet of mainly protein (see also below). Compared with 18 spp. of other rodents (mostly murids), the alimentary canal is short in relation to head and body length (3.7 times HB, cf. 4.8–5.6 for herbivores), the small intestine is relatively short and the large intestine is relatively long in relation to total hindgut length; the caecum is absent (the only species of those studied without a caecum) and there are no spiral folds in the large intestine (cf. all other 18 spp. except Cryptomys hottentotus). Most of these characters are primitive and indicative of a nutritious diet (Perrin & Curtis 1980).

Abundance Common in some regions, uncommon in others. In Giant’s Castle Game Reserve, South Africa, relative abundance (as assessed by trap success) was 0.3% in grouped tree woodland, 0.8% in scrub, 3.8% in forest, 0.6% in temperate grassland boulder bed and 0.4% in temperate grassland (Rowe-Rowe & Meester 1982a). The species is fairly common and widespread in SE South Africa from Western Cape Province to Mpumalanga and Limpopo

Foraging and Food Omnivorous, predominantly insectivorous and carnivorous. Stomach contents have included insects and other invertebrates, seeds, leaves, stems, fruit and occasionally small vertebrates (Kingdon 1974, Perrin & Curtis 1980, Rowe-Rowe 1986,Wirminghaus & Perrin 1992,Taylor 1998; specimen labels). In KwaZulu–Natal, South Africa, most stomachs contained seeds, and all contained arthropods (n = 11; Rowe-Rowe 1986). In another study

Graphiurus murinus

E. Holden unpubl.). Generally not sympatric, although they have been captured in close proximity at several localities in Kenya. Occurs in savannas throughout much of sub-Saharan Africa. Distribution Endemic to Africa. Afromontane–Afroalpine BZ in eastern and southern Africa, Highveld BZ, and Coastal Forest Mosaic BZ in southern part of range. Recorded from Ethiopia, Kenya, Uganda, Rwanda, Burundi (and possibly extreme NE DR Congo), Tanzania, N Malawi, Zimbabwe, South Africa, Swaziland and Lesotho (Holden 2005).

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in KwaZulu–Natal, South Africa, stomachs contained invertebrates (88.8%), fruits (11.04%), leaves and stems (0.16%), and flowers (0%); this population was thus predominantly insectivorous with more than 80% of the diet in any month comprising invertebrates (n = 23; Wirminghaus & Perrin 1992). One individual found in a beehive (Duff-MacKay 1965, quoted by Kingdon 1974) was extremely fat, and its stomach contained a brown sludge with white specks – presumably honey and wax.

G. murinus, and therefore the ectoparasites listed here could have been collected from either species. Ectoparasites include three families of mites (Laelaptidae [1 sp.], Ereynetidae [1 sp.] and Trombiculidae [8 spp.]), four families of fleas (Ceratopsyllidae [1 sp.], Chimaeropsyllidae [1 sp.], Hystricopsyllidae [7 spp.] and Pulicidae [5 spp.]), and the rhipicephalid tick Rhipicephalus simus (De Graaff 1981). Type (and principal) host for the hoplopleurid louse Schizophthirus graphiuri (Durden & Musser 1994, Pajot 2000).

Social and Reproductive Behaviour Little information. Lactating !! often caught with young. Allen & Loveridge (1933) observed subadult nestlings following an adult ! (up to eight individuals) as they climbed and leapt across tree branches. In two separate instances, adult !! were captured with two subadult "", indicating that offspring may stay in the nest past weaning (Allen & Loveridge 1933; specimen label).

Conservation IUCN Category: Least Concern. Further sampling is required to assess conservation status throughout geographic range; some historically recorded populations may be threatened or even extinct (e.g. the population in the Udzungwe Mts, Tanzania).

Reproduction and Population Structure Litter-size: 1–5. Most often 3–4 embryos or nestlings are reported (Hollister 1919, Ansell 1974, De Graaff 1981; specimen labels). In Kenya, pregnant !! were collected in Sep and Nov (specimen labels). In Zambia, a pregnant ! was captured in Jul (Ansell 1974). In South Africa, !! exhibiting placental scars were captured in Feb, and pregnant !! were collected in Oct, Dec and Feb (Lynch 1989; specimen labels). Taylor (1998) suggests that this species may breed mostly during summer in KwaZulu–Natal, South Africa. No information on gestation and development of young. Sex ratio has been found to be female-biased in some southern African populations (De Graaff 1981, Smithers 1983).

Measurements Graphiurus murinus HB: 91.5 (81–103) mm, n = 21 T: 76.6 (69–85) mm, n = 19 HF: 18.5 (16–20) mm, n = 21 E 13.3 (11.5–16) mm, n = 7 WT: 17 g, n = 1 GLS: 26.4 (25.2–28.8) mm, n = 19 GWS: 14.2 (13.2–15.9) mm, n = 11 P4–M3: 3.1 (3–3.3) mm, n = 21 Mt Kenya (1829–3353 m), Kenya (M. E. Holden unpubl.) Key References Allen & Loveridge 1933; Kryštufek et al. 2004; Roberts 1951; Taylor 1998. Mary Ellen Holden

Predators, Parasites and Diseases No information on predators. Most authors have included G. microtis as a synonym of

Graphiurus nagtglasii NAGTGLAS’S AFRICAN DORMOUSE Fr. Graphiure de Nagtglas (formerly Graphiure de Huet); Ger. Nagtglas Bilch Graphiurus nagtglasii Jentink, 1888. Notes from the Leyden Museum 10: 38–41. Hill Town, Du Queah River, Liberia (restricted locality).

Taxonomy Most previous publications on dormice in West Africa have referred to this species as G. hueti (Huet’s African Dormouse). Although originally described in genus Graphiurus, it was later placed in its own genus, Aethoglis (Allen 1936; see profile Genus Graphiurus). Grubb & Ansell (1996) recommended applying the name G. nagtglasii to the large West African dormouse, traditionally known as G. hueti, because of the dubious nature of the type locality of G. hueti given by de Rochebrune (1883), the lack of an available or likely holotype for G. hueti, and the existence of a holotype for nagtglasii. The animal used by de Rochebrune as a model to figure G. hueti is probably from Gabon, not Senegal (the type locality of G. hueti). Jentink (1888, and on his specimen labels) designated a series of five syntypes representing G. nagtglasii from Ghana and Liberia, and listed what are presumed to be the type localities as the Du Queah and Farmington rivers for the three Liberian specimens. As Rosevear (1969) noted, Jentink based his description on (and gave measurements for) an adult " in alcohol that he considered typical of the species. Allen (1939) and Genest-

Villard (1978a) included G. monardi as a subspecies of G. hueti (now G. nagtglasii), but Ellerman et al. (1953), Ansell (1978), Robbins & Schlitter (1981) and Holden (1993, 2005) observed that G. monardi is clearly distinct from and probably not closely related to G. nagtglasii. Synonyms: argenteus, hueti (see Allen 1939, Grubb & Ansell 1996). Subspecies: none. Chromosome number: 2n = 40 (Tranier & Dosso 1979). Description Large dormouse. Dorsal pelage brown, greyishbrown, or rufous. Dorsal pelage soft, woolly, dense and short (rump hairs 5–7 mm, guard hairs up to 12 mm). Ventral pelage dark grey washed with ochre, cream or white. Dorsal and ventral pelage colours not clearly delineated. Head colour matches that of dorsal pelage. Eyes large; eye-mask usually conspicuous. Ears brown, mediumlength, narrow and pointed. Cheeks greyish-white or ochraceouswhite. Postauricular patches not present. Hindfeet brown, or white with dark metatarsal streak. Tail moderately long (ca. 76% of HB), tail hairs shorter at base (14–18 mm) and longer at tip (up

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to 37 mm). Tail colour generally matches that of dorsal pelage, but has variegated frosted appearance. A few white hairs are occasionally mixed throughout tail; tip not white. Tail distichous underneath, and often nearly naked along mid-ventral line. Skull large, vaulted (height of braincase 10.2 mm) and somewhat broad (20.7 mm) relative to skull length. Anterior chamber of auditory bullae usually markedly less inflated than the posterior chambers. Interorbital constriction (5.5 mm) relatively narrow. Palate (13.0 mm) and upper toothrow (5.1 mm) relatively long, and wide as exhibited by breadth of upper premolar (1.5 mm). Anterior palatal foramina absolutely somewhat long (mean 3.7 mm) but relatively short. Auditory bullae short (7.9 mm) and uninflated relative to skull length (measurements listed are mean values from Ghana; M. E. Holden unpubl.). Nipples: 1 + 1 + 2 = 8. Geographic Variation None recorded. Similar Species (size comparisons refer to mean values only) Graphiurus monardi.This species is not at all similar to G. nagtglasii, but a comparison is given here due to previous inclusion of G. monardi in G. nagtglasii. Probably similar head and body length (mean 160 mm). (Only one adult specimen of G. monardi has information on head and body length associated with it; however, head and body length appears to be similar for other specimens.) Ear (15.5 mm) and hindfeet (21.9 mm) are shorter both absolutely and relative to skull length. Dorsal pelage long, straight and silky, with many conspicuous dark brown guard hairs. Skull shorter (34.1 mm) with absolutely and relatively shorter palate (10.7 mm). Anterior palatal foramina longer (4.1 mm), both absolutely and relative to skull length. Auditory bullae (10.3 mm) absolutely and relatively longer, and much more inflated (measurements listed are mean values from NE Angola, S DR Congo and NW Zambia populations; M. E. Holden unpubl.). Not sympatric. Occurs in S DR Congo, NE Angola and NW Zambia.

Distribution Endemic to West Africa. Rainforest BZ (Western and West Central Regions) and Northern Rainforest–Savanna Mosaic. Recorded in or near rainforest from S Sierra Leone to Cameroon (excluding Benin), SW Central African Republic and Gabon. The southern distributional limit of this species is not known, as the Gabon specimen has no specific locality. Although de Rochebrune (1883) listed the type locality of G. hueti as Senegal, and reported it from neighbouring Gambia, the occurrence of the species has never been substantiated in these countries (Grubb & Ansell 1996). Habitat Recorded in rainforest, secondary forest, abandoned farmlands, in cocoa plantations and other farms in forested areas (Rosevear 1969, Happold 1987, as G. hueti specimen labels). Most museum specimens were collected from hollow trees, one from an old hollow banana stem, and several from banana groves within cocoa plantations (Jeffrey 1973). They are often caught by farmers preparing new farms from secondary bush or forest (Jeffrey 1973). Several individuals were trapped on vines in secondary forest near hollow trees (Robbins & Schlitter 1981). Abundance Rosevear (1969) stated that these dormice are fairly common and widespread throughout the West African rainforest. In Côte d’Ivoire, Heim de Balsac (1967a) found them to be fairly common in Lamto. In contrast, Dosso (1975a) did not trap this dormouse during his faunal study at Adiopodoumé, Côte d’Ivoire. Happold (1987) reported its occurrence in Nigeria as rare compared with G. murinus (the ‘G. murinus’ of Happold (1987) is now known to represent G. lorraineus, G. kelleni and G. crassicaudatus). Based on numbers of specimens in museum collections from countries throughout its distribution, the abundance of this dormouse probably ranges from uncommon to common. Adaptations Arboreal and probably nocturnal. According to Jeffrey (1973), these dormice climb well, but move slowly on the ground. Nests are often made in hollow trees, and one ! made a nest of dry banana fibres inside an old banana plant stem (specimen label). Eight !! with young trapped in banana groves made their nests of dry banana leaves. Happold (1987) states that these dormice curl up in their nests with the tail folded over the head during the day. Aside from one individual said by the collector, G. L. Bates, to have been ‘shot with bow, coming out of hollow tree’ (specimen label), all other specimens for which such information is available were taken while sleeping during the day. Foraging and Food Probably omnivorous. In the wild, foods include cocoa pods, oil palm nuts, paw-paw, bananas and insects (Everard 1968, Happold 1987). Because these dormice are known to nest in hollow trees, and have been taken only among vines or in trees, they presumably do not forage on the ground. Social and Reproductive Behaviour Apparently solitary, except for lactating !!. Rosevear (1969) reported that this dormouse was less shy and not as easily scared as the smaller dormice occurring in West Africa.

Graphiurus nagtglasii

Reproduction and Population Structure Litter-size: probably 2–3. In Liberia, Coe (1975) collected a parous adult in Apr. 127

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In Ghana, G. S. Cansdale trapped several !! in Sep and Mar with three lactating young whose eyes were not yet opened (specimen labels). In W Ghana, Jeffrey (1973) found eight !!, each nesting with two or three young in Sep, Oct and Nov. In Cameroon, an adult ! containing two embryos was found in Feb, and lactating ! was caught at the same locality in Apr (Robbins & Schlitter 1981). These very limited data suggest that most young are born during the wetter months of the year, but this needs confirmation. Predators, Parasites and Diseases Heim de Balsac (1967a) and Jeffrey (1973) state that local people actively hunt Nagtglas’s Dormice for their prized meat. This dormouse is the type and principal host for the hoplopleurid louse Schizophthirus aethogliris (Durden & Musser 1994, Pajot 2000). Conservation

Measurements Graphiurus nagtglasii HB: 138.5 (120–155) mm, n = 48 T: 105 (65–122) mm, n = 43 HF: 26.5 (20–30) mm, n = 46 E: 18.1 (15–22) mm, n = 44 WT: 73.1 (48–98) g, n = 18 GLS: 36.8 (34.9–39.1) mm, n = 35 GWS: 20.7 (18.3–22) mm, n = 39 P4–M3: 5.1 (4.6–5.7) mm, n = 48 Measurements: Ghana (M. E. Holden unpubl.) Weight: Liberia, Côte d’Ivoire and Ghana (M. E. Holden unpubl.) Key References Grubb & Ansell 1996; Grubb et al. 1998; Happold 1987; Robbins & Schlitter 1981; Rosevear 1969.

IUCN Category: Least Concern. Mary Ellen Holden

Graphiurus ocularis SPECTACLED AFRICAN DORMOUSE (NAMTAP) Fr. Graphiure du Cap; Ger. Brillen-Bilch Graphiurus ocularis (Smith, 1829). Zool. J., 4: 439. Near Plettenberg Bay, Cape Province, South Africa.

Taxonomy Originally described in the genus Sciurus. In some regions of South Africa, the common name Namtap is used (see Channing 1987 for discussion of other common names applied to this species). Synonyms: capensis, elegans, typicus (see Allen 1939). Subspecies: none. Chromosome number: 2n = 46 (D. N. MacFadyen pers. comm.).

Auditory bullae (9.8 mm) of medium length and only moderately inflated relative to skull length (measurements listed are mean values from Northern and Western Cape Provinces, South Africa; M. E. Holden unpubl.). Nipples: 1 + 1 + 2 = 8.

Description Large dormouse. Dorsal pelage medium silverygrey. Dorsal pelage woolly, thick and moderately long (rump hairs 11–12 mm, guard hairs up to 16 mm). Ventral pelage dark grey washed with white. Dorsal and ventral pelage colours moderately delineated. Head silvery-grey, paler towards muzzle. Eyes large; eye-mask very conspicuous and broad, extending further posteriorly (to beneath the ear) than in other species of Graphiurus. Ears brown, moderately large, rounded. Cheeks white, forming part of a white sharply demarcated lateral stripe that extends from cheeks to shoulders. Conspicuous white supra-auricular patches. The combination of broader, more extensive face-mask, white cheeks, pale muzzle and larger, more conspicuous supra-auricular patches results in a striking black-and-white colour pattern on head and shoulders that allows for easy identification of this dormouse based on external characters alone. Hindfeet white and short (24.2 mm) relative to body size. Tail moderately long (ca. 85% of HB), tail hairs shorter at base (10–15 mm) and longer at tip (up to 35 mm). Dorsal tail colour generally matches that of dorsal pelage, but ventral tail colour solid brown-black medially, fringed with white laterally; tip white. Skull long (35.8 mm), moderately flattened (height of braincase 9.3 mm) and moderately broad (19.5 mm). Upper toothrow (3.3 mm) very short, upper premolar circular and very narrow (0.6 mm) both absolutely and relative to skull length; the reduced premolar allows for easy identification of skulls of this species. Anterior palatal foramina of moderate length (3.5 mm) and width (2.2 mm), but short and narrow relative to skull length.

Similar Species (size comparisons refer to mean values only) Graphiurus rupicola. Smaller body length (mean 110 mm). The combination of paler grey dorsal pelage colour, darker muzzle, smaller and less conspicuous white supra-auricular patches and shorter eye-mask (terminating posterior to eye) results in a less striking facial colour pattern. Tail (104.2 mm), ear (17.3 mm) and hindfoot (21.5 mm) absolutely shorter, but relatively longer. Ventral tail colour similar to dorsal tail colour; not darker as in G. ocularis. Skull shorter (31.3 mm), interorbital constriction (5.0 mm) absolutely narrower, but relatively broader. Upper toothrow (3.4 mm) similar in absolute length, but longer relative to skull length. Upper premolar broader (0.9 mm) and oval-shaped. Anterior palatal foramina absolutely similar in length (3.4 mm) and breadth (2.3 mm), but relatively longer and broader. Auditory bullae (9.4 mm) absolutely shorter, but relatively longer and more inflated (mean values from Erongo, Karibib, and Mt Brukkaros, Namibia; M. E. Holden unpubl.). Parapatric in Little Namaqualand, Western Cape Province, South Africa. Also occurs in Namibia and Angola.

Geographic Variation

None recorded.

Distribution Endemic to Africa. South-West Arid (Karoo) and South-West Cape BZs. Recorded in Eastern Cape, Northern and Western Cape Provinces, South Africa. Altitudinal range from near sea level to 1585 m. Other historical distributional records outside the range given here are questionable (Holden 2005).

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unnatural foods such as crackers, fruit, meat and honey, as well as dog food and rat pellets (Channing 1984, Perrin & Ridgard 1999). Social and Reproductive Behaviour Spectacled Dormice emit vocalizations and exhibit intimidation displays during disturbance and aggression (Channing 1987, Van Hensbergen & Channing 1989). Fluorescent powder tracking suggests that individuals lay scent trails, and that "" and !! follow common routes (Channing 1984, Van Hensbergen & Channing 1989). Home-range (as assessed by harmonic range analysis) is 1.1–2.3/ha for !! and 2.1–3.8/ha for "" (Van Hensbergen & Channing 1989). There is some evidence that these dormice may be territorial because pairs remain together in the same area for up to 11 months.

Graphiurus ocularis

Habitat Spectacled Dormice prefer rock piles, outcrops, crevices and stone kraals (Roberts 1951, Channing 1984, Fox et al. 1985, Skinner & Smithers 1990). They have also been captured in huts (Channing 1984) and within the hollow door of a farmhouse (De Graaff & Rautenbach 1983), and there is one report of an individual in a tree (Rautenbach 1982). Abundance Uncommon. Population densities ranged from 1.8/ ha to 3.1/ha on a 7.75 ha study site; densities vary according to the suitability of the habitats (Channing 1984). Smithers (1986a) and De Graaff & Rautenbach (1983) consider it rare. Adaptations Predominantly rupicolous, also terrestrial. Nocturnal. The flattened cranium allows animals to move through narrow rock crevices. These dormice prefer to travel along rocks above ground level, even if the distance would be considerably shortened by taking a ground-level path (Channing 1997). Fox et al. (1985) suggest that these dormice are an early successional species after fire. They remain active throughout the year, but if there is a drop in temperature, and/or a scarcity of food, they can enter torpor for up to a month (Channing 1997, Perrin & Ridgard 1999). In captivity, they were unable to tolerate ambient temperatures greater than 35 °C (Perrin & Ridgard 1999). Foraging and Food Predominantly insectivorous, occasionally carnivorous. Diet primarily insects and arthropods, although birds and lizards are also taken; probably no seasonal variation in diet (Channing 1984). In captivity, Spectacled Dormice consume

Reproduction and Population Structure Reproductively active in spring and summer. Litter-size: 4–6. Litter interval: 6–8 weeks.Young stay in the nest for 5–6 weeks (Channing 1997). Social structure is primarily male–female pairs with their young of the year. Each pair (with young) occupies the most favourable habitats. Other individuals, including young after leaving their parents, occupy less favourable habitats. Average life-span is thought to be at least four years (Channing 1984). Predators, Parasites and Diseases Ectoparasites include species of three families of fleas (Hystricopsyllidae, Listropsyllidae and Chimaeropsyllidae) and the tick Rhipicephalus simus (De Graaff 1981). Conservation IUCN Category: Least Concern. The categorization of this species should be changed to Near Threatened, or at least Data Deficient, due to its discontinuous distribution and poor representation in museum collections. Smithers (1986) categorized the species as rare. Measurements Graphiurus ocularis HB: 134.3 (117–145) mm, n = 19 T: 114.5 (103–150) mm, n = 20 HF: 24.2 (20–26) mm, n = 25 E: 19.5 (15–25) mm, n = 24 WT: 78 (72–85) g, n = 4 GLS: 35.8 (34.2–37.5) mm, n = 20 GWS: 19.5 (18.3–20.9) mm, n = 15 P4–M3: 3.28 (3–3.5) mm, n = 24 Northern and Western Cape Provinces, South Africa (M. E. Holden unpubl.) Key References Channing 1997; De Graaff 1981; Roberts 1951; Skinner & Smithers 1990. Mary Ellen Holden

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Graphiurus platyops FLAT-HEADED AFRICAN DORMOUSE Fr. Graphuire á tête plate; Ger. Flachkopf-Bilch Graphiurus platyops Thomas, 1897. Ann. Mag. Nat. Hist., ser. 6, 19: 388. Enkeldorn, Mashonaland, S Zimbabwe.

Taxonomy Ellerman et al. (1953) synonymized G. rupicola from NW South Africa and Namibia, and G. angolensis (including parvulus) from Angola and NW Zambia within this species. Their arrangement was generally followed by Genest-Villard (1978a), though she placed the Angolan populations (G. angolensis and parvulus) as junior synonyms of G. murinus. Allen (1939) recognized G. platyops, G. rupicola and G. angolensis, as well as parvulus (listed here as a junior synonym of G. angolensis) as separate species. Roberts (1951) also treated G. rupicola and G. platyops as distinct species, a position followed by Holden (1993, 2005) based on comparisons of museum specimens. Following Ansell (1978), Holden (1993) provisionally listed angolensis and parvulus as synonyms of G. platyops, but analyses of museum specimens (M. E. Holden unpubl.) support recognition of the Angola and NW Zambia population as a separate valid species, G. angolensis (including parvulus). Synonyms: eastwoodae, jordani (see Holden [2005] regarding allocation of albicaudatus, a name that apparently was never published). Subspecies: none. Chromosome number: 2n = 46 (D. N. MacFadyen pers. comm.). Description Medium-sized dormouse. Dorsal pelage grey, brownish-grey or greyish-brown. Pelage sleek and moderately long (rump hairs 10 mm, guard hairs up to 13 mm). Ventral pelage white or cream suffused with grey. Dorsal and ventral pelage colours clearly delineated. Head colour matches that of dorsal pelage, paler towards muzzle. Eyes large; eye-mask conspicuous. Ears brown, moderately large, rounded. Cheeks white, forming part of a pale lateral stripe that extends from cheeks to shoulders. Faint white postauricular patches sometimes present. Hindfeet usually white, sometimes with dark metatarsal streak. Tail moderately short (ca. 71% HB), similar in colour to dorsal pelage, with many scattered white hairs; hairs shorter at base (5–7 mm) and longer at tip (up to 30 mm); tip white. Skull gracile, broad (17.1 mm) and flat (height of braincase 7.8 mm). In lateral profile, dorsal outline of skull from rostrum to occiput is practically horizontal. Interorbital constriction moderately broad (4.8 mm), anterior palatal foramina moderately long (3.2 mm), but somewhat short relative to skull length. Upper toothrow relatively short (3.1 mm) and narrow as exhibited by narrow breadth of upper premolar (0.8 mm). Auditory bullae of medium length (8.4 mm) and only moderately inflated relative to skull length (measurements listed are mean values from Zimbabwe and NE South Africa; M. E. Holden unpubl.). Nipples: 1 + 1 + 2 = 8.

Graphiurus platyops

the height of braincase (7.5 mm) is absolutely shorter in G. microtis, in lateral view the braincase is markedly more vaulted and its dorsal outline convex. It is also relatively higher relative to skull length. Upper toothrow (3.0 mm), anterior palatal foramina (3.4 mm) and auditory bullae (8.1 mm) similar to slightly shorter in length, so relatively longer. Anterior palatal foramina 3.4 mm long, 2.1 mm wide. Interorbital constriction (3.9 mm) absolutely and relatively narrower (measurements listed are mean values from Zimbabwe; M. E. Holden unpubl.). Generally sympatric throughout range of G. platyops, but seems to be segregated by habitat, and is probably not syntopic. Graphiurus microtis occurs in savannas throughout most of sub-Saharan Africa. Distribution Endemic to Africa. Southern part of Zambezian Woodland BZ and northern part of Highveld BZ. Recorded from NE and S Zambia, Zimbabwe, S Malawi, E Botswana, S Mozambique and NE South Africa (Holden 2005). Previously thought to occur in C Botswana (De Graaff 1981); however, the museum specimen on which the record was based is G. microtis (Holden 2005).

Geographic Variation None recorded. Similar Species (size comparisons refer to mean values only) Graphiurus microtis. Smaller head and body length (mean 98.8 mm). Tail (75.2 mm) absolutely shorter, but relatively longer. Hindfeet (16.9 mm) shorter, and ears (15.5 mm) longer, both absolutely and relatively. Dorsal pelage greyish-brown to dark greyishbrown in populations sympatric with G. platyops. Skull shorter (27.4 mm), vaulted and relatively narrower (15.0 mm). Although

Habitat Flat-headed African Dormice are most often trapped in crevices in rock kopjes, krantzes and under exfoliating granite (Roberts 1951, Smithers 1971, 1983, Wilson 1975). They have sometimes been found in association with dassies (Heterohyrax and Procavia) (Roberts 1951, Smithers & Lobão Tello 1976). Three individuals were trapped in caves in South Africa (specimen labels). On the Save R., Mozambique, individuals were collected in dry Androstachys sp. scrub thickets in a dry river bed (Smithers & Lobão

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Tello 1976). In NE South Africa, one ! with three young was caught in a hollow tree branch (Roberts 1951). Rautenbach (1982) observed that this species generally occurs at altitudes higher than 609 m, and is not associated with mopane woodland. Abundance Little information. Uncommon in E Zambia (Ansell 1978). There are only ca. 50 museum specimens of G. platyops, and usually only one or two individuals have been found at each locality, suggesting that even in suitable habitat, densities are not high. Adaptations Predominantly rupicolous. The markedly flattened cranium allows movement of these animals through narrow rock crevices, where they are most commonly found. Most authors characterize this dormouse as being nocturnal (e.g. Smithers 1971), but five individuals trapped by Wilson (1975) in W Zimbabwe were taken between 06:00 and 09:00h, suggesting that at least some individuals are crepuscular or diurnal. Foraging and Food Omnivorous. Stomach contents of individuals from Zimbabwe and Botswana contained remains of well-masticated small seeds, traces of green vegetable matter and the chitinous remains of insects, including moths (Smithers 1983). One animal was trapped using a portion of a rat carcass for bait (specimen label). Social and Reproductive Behaviour Apparently solitary (Smithers 1983). Unbaited tunnel traps that have been entered by one individual seem to attract others, suggesting that, like Spectacled Dormice, these dormice use scent trails (Channing 1997). They are aggressive, flourishing and whipping their tails as a visual signal (Channing 1997).Vocalizations include a soft warning call, consisting of a number of short, low-pitched notes. An aggression call follows if the intruder does not leave. The aggression call consists of a series of brief spits, each consisting of a 0.1-second burst of high amplitude

white noise. The encounter escalates into a fight if the intruder remains (Channing 1997). Reproduction and Population Structure Little information. In Zimbabwe, a pregnant ! carrying two full-term embryos was obtained in Feb (specimen label). Predators, Parasites and Diseases Flat-headed African Dormice are hosts of the chimaeropsyllid flea Chiastopsylla nama, which also occurs on other rupicolous rodents including Aethomys namaquensis and Petromyscus collinus (Haeselbarth et al. 1966, De Graaff 1981). Conservation IUCN Category: Least Concern. Little information is known regarding distribution and abundance; a more appropriate categorization would be Data Deficient. Measurements Graphiurus platyops HB: 107.1 (95–122) mm, n = 18 T: 78.7 (65–98) mm, n = 15 HF: 21.1 (18–25) mm, n = 21 E: 15.2 (13–18) mm, n = 19 WT: 45.7 (30.4–52.8) g, n = 5 GLS: 30.4 (28.6–32) mm, n = 19 GWS: 17.1 (16.1–18.6) mm, n = 20 P4–M3: 3.1 (2.8–3.5) mm, n = 25 Zimbabwe and NE South Africa (M. E. Holden unpubl.) Key References Ansell 1978; Channing 1997; De Graaff 1981; Roberts 1951; Smithers 1983. Mary Ellen Holden

Graphiurus rupicola RUPICOLOUS AFRICAN DORMOUSE Fr. Graphiure des rochers; Ger. Felsen-Bilch Graphiurus rupicola (Thomas and Hinton, 1925). Proc. Zool. Soc. Lond. 1925: 232. Karibib, Namibia. 3842 ft (1170 m).

Taxonomy Originally described in the genus Gliriscus. Ellerman et al. (1953) and Genest-Villard (1978a) considered rupicola to be a subspecies of G. platyops. Roberts (1951) recognized the substantial morphological differences between rupicola and platyops, and listed G. rupicola as a separate species, an arrangement followed here. Synonyms: australis, kaokoensis, montosus. Subspecies: none. Chromosome number: not known. Description Medium-sized dormouse. Dorsal pelage silverygrey, drab grey or slate-grey. Dorsal pelage woolly, thick and moderately long (rump hairs 10–11 mm, guard hairs up to 17 mm). Ventral pelage white or cream lightly suffused with dark grey. Dorsal and ventral pelage colours clearly delineated. Head colour matches that of dorsal pelage, slightly paler towards muzzle. Eyes large; eye-mask conspicuous. Ears brown, large, oval-shaped. Cheeks cream or white, forming part of a pale lateral area that extends from cheeks to shoulders. Cream supra-auricular patches, and faint

postauricular patches present. Hindfeet white, or white with dark metatarsal streak. Tail long (ca. 95% HB), similar in colour to dorsal pelage, with many scattered white hairs; hairs shorter at base (9– 12 mm) and longer at tip (up to 43 mm); white at tip. Skull long and moderately flattened (height of braincase 8.0 mm). Palate long (10.4 mm), upper premolar breadth (0.88 mm) somewhat narrow relative to skull length, interorbital constriction (5.0 mm) broad, anterior palatal foramina moderately long (3.4 [3.1–3.6] mm) and wide (mean 2.3 mm), auditory bullae long (9.4 mm) and inflated relative to skull length (measurements are mean values from Erongo, Karibib and Mt Brukkaros, Namibia; M. E. Holden unpubl.). Nipples: 1 + 1 + 2 = 8. Geographic Variation

None recorded.

Similar Species (size comparisons refer to mean values only) Graphiurus angolensis. Slightly smaller body length (mean 98.8 mm). 131

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Family GLIRIDAE

Tail (79.2 mm) and hindfeet (18.4 mm) are absolutely and relatively shorter. Dorsal pelage is drab or dark brown. Skull averages slightly shorter (28.2 mm). Although height of braincase (7.7 mm) is absolutely shorter in G. angolensis, in lateral view the braincase appears more vaulted and its dorsal outline is more convex. Interorbital constriction (4.2 mm) absolutely narrower and auditory bullae (8.9 mm) absolutely shorter but relatively longer. Upper premolar breadth (0.90 mm) absolutely similar but broader relative to skull length. Anterior palatal foramina absolutely similar in length (3.4 mm) but relatively longer (measurements listed are mean values from Kabompo and Zambezi [formerly Balovale], Zambia; M. E. Holden unpubl.). Parapatric in C Angolan highlands, occurs in Angola and NW Zambia. Graphiurus microtis. Slightly smaller body length (mean 98.8 mm). Tail (75.2 mm) and hindfeet (16.9 mm) are absolutely and relatively shorter. Dorsal pelage sometimes similar in colour, especially in Namibia, Botswana and NE South Africa populations. Skull markedly shorter (27.4 mm). Palate (8.8 mm) shorter, interorbital constriction (3.9 mm) narrower, and auditory bullae (8.1 mm) shorter both absolutely and relatively. Upper toothrow (3.0 mm) absolutely shorter, but relatively similar. Upper premolar breadth (0.85 mm) absolutely narrower, but broader relative to skull length. Anterior palatal foramina 3.4 mm long, 2.1 mm wide, similar in length, so relatively longer (measurements listed are mean values from Zimbabwe; M. E. Holden unpubl.). Sympatric near Okahandja, Namibia; occurs in savannas throughout sub-Saharan Africa. Graphiurus ocularis. Larger body length (mean 134.3 mm). Tail (114.5 mm), ear (19.4 mm) and hindfeet (24.2 mm) are absolutely longer, but relatively shorter. Dorsal pelage darker grey with striking facial colour pattern. Dorsal tail darker than ventral tail colour. Skull (35.8 mm) longer. Palate (12.0 mm) absolutely longer, but relatively similar. Interorbital constriction (5.4 mm) absolutely broader, but relatively narrower. Auditory bullae

(9.8 mm) slightly longer and upper toothrow (3.3 mm) similar, but both shorter relative to skull length. Anterior palatal foramina absolutely similar in length (3.5 mm) and breadth (2.2 mm), but relatively shorter. Upper premolar circular and greatly reduced in size (breadth 0.6 mm) (measurements listed are mean values from Northern and Western Cape Provinces, South Africa; M. E. Holden unpubl.). Parapatric in Little Namaqualand,Western Cape Province, South Africa; also occurs in Eastern and Northern Cape Provinces. Distribution Endemic to Africa. South-West Arid BZ (with possible northern extension to Zambezian Woodland BZ in Angola). Occurs on central mountains and plateaux from Mt Soque, Angola, south to Port Nolloth and Eenriet in Little Namaqualand, South Africa (Holden 2005). The northern distributional limit for G. rupicola was previously thought to be Kamanjab, Namibia, but specimens from Mt Soque, Angola, are considered to represent this species (Holden 2005). A specimen from Dilolo, DR Congo, resembles G. rupicola in pelage characters, but resembles G. monardi in skull characters; at present the specimen cannot be allocated to either species (Holden 2005). Habitat Rock crevices in rocky outcrops and kopjes, from altitudes of 400 m to at least 1586 m. Most specimens were caught in bushy Karoo–Namib shrubland or Karoo transition vegetation zones; two specimens from Mt Soque, Angola, were captured in ‘evergreen wood at mountain top’ (specimen label). Abundance Little information. Roberts (1951) considered the species to be rare, an observation supported by the few specimens (ca. 20) in museums. Remarks The moderately flattened skull enables animals to squeeze through narrow rock crevices. The very few habitat notes (Shortridge 1934, Roberts 1951; specimen labels) suggest that G. rupicola nests only in rock crevices. A ! obtained on Mt Brukkaros, Namibia, in Sep was pregnant (specimen label). Conservation IUCN Category: Least Concern. So little is known about the species that a classification of Data Deficient may be more appropriate. Measurements Graphiurus rupicola HB: 110 (105–119) mm, n = 9 T: 104.2 (96–118) mm, n = 9 HF: 21.5 (21–22) mm, n = 8 E: 17.3 (16–20) mm, n = 9 WT: (subadult): 25 g, n = 1* GLS: 31.3 (30.5–32.3) mm, n = 7 GWS: 17.2 (16.7–17.9) mm, n = 6 P4–M3: 3.4 (3.3–3.7) mm, n = 8 Erongo, Karibib and Mt Brukkaros, Namibia (M. E. Holden unpubl.) *Pella Mission, south bank of the Orange River, South Africa (M. E. Holden unpubl.) Key References

Graphiurus rupicola

Roberts 1951; Shortridge 1934. Mary Ellen Holden

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Graphiurus surdus

Graphiurus surdus SHORT-EARED AFRICAN DORMOUSE Fr. Graphiure sourd; Ger. Kurzohr-Bilch Graphiurus surdus Dollman, 1912. Ann. Mag. Nat. Hist., ser. 8, 9: 314. Benito River, Rio Muni Province, Equatorial Guinea.

Taxonomy Although described and initially recognized as a species, G. surdus was later synonymized within a broadly defined G. murinus (Misonne 1974, Genest-Villard 1978a). Robbins & Schlitter (1981) and Holden (1996) provided morphological evidence for recognizing G. surdus as a valid species. Synonym: schwabi. Subspecies: none. Chromosome number: not known. Description Small to medium-sized dormouse. Dorsal pelage greyish-brown. Dorsal pelage silky and moderately long (rump hairs 5–7 mm, guard hairs up to 11 mm). Ventral pelage dark grey washed with whitish-buff. Dorsal and ventral pelage colours not clearly delineated. Head colour matches that of dorsal pelage. Eyes large; eye-mask inconspicuous. Ears brown, short and rounded. Cheeks grey washed with white. Postauricular patches not present. Hindfeet cream or white with dark metatarsal streak. Tail moderately long (ca. 73% of HB), similar in colour to dorsal pelage, with many scattered white hairs resulting in a frosted appearance; hairs shorter at base (3–8 mm) and longer at tip (up to 20 mm); tip not white. Skull medium length (27.6 mm), moderately vaulted (height of braincase 8.1 mm), with narrow greatest width of skull (14.6 mm), and comparatively straight conformation of the zygomatic arch in lateral view (figured in Robbins & Schlitter 1981 and Holden 1996). Interorbital constriction (4.5 mm) narrow. Anterior chamber of auditory bullae markedly less inflated than posterior chambers in most individuals. Anterior palatal foramina relatively short (2.8 mm) and narrow (1.8 mm). Upper premolar narrow (0.8 mm), palate long (9.3 mm) and auditory bullae short (7.3 mm) and uninflated relative to skull length (measurements listed are mean values from Cameroon, Equatorial Guinea and Gabon; Holden 1996). Nipples: 1 + 1 + 2 = 8.

(24.5 mm) much shorter, and palate absolutely and relatively shorter (7.8 mm). Similar average length of auditory bullae (7.1 mm) and breadth of upper premolar (0.8 mm), but bullae are longer and upper premolar is wider relative to skull length. Anterior palatal foramina absolutely similar in length (2.6 mm), but relatively somewhat longer (measurements listed are mean values from DR Congo; M. E. Holden unpubl.). Sympatric in SW Cameroon, Equatorial Guinea and DR Congo (Holden 1996). Graphiurus crassicaudatus. Smaller head and body length (92.6 mm). Hindfeet (17.7 mm) and tail (59.4 mm) shorter. Dorsal pelage usually rufous-brown. Skull shorter (26.6 mm). Greatest width of skull (16.1 mm) and interorbital constriction (4.9 mm) markedly broader relative to skull length, supraorbital ridges present. Palate (9.4 mm) similar in length but relatively longer. Upper toothrow (3.8 mm) absolutely and relatively longer (measurements listed are mean values from S Cameroon; M. E. Holden unpubl.). Sympatric in SW Cameroon; occurs in West and west-central Africa. Distribution Endemic to Africa. Rainforest BZ (West Central Region [Gabon Subregion], and marginally East Central and South Central Regions). Recorded from two disjunct areas in central Africa: (1) S Cameroon south to Equatorial Guinea and Gabon; (2) NE Congo (Masako) and SC Congo (Inkongo). Of the 22 known specimens, 15 are from S Cameroon. Limits of geographic range unknown.

Geographic Variation None recorded. Similar Species (size comparisons refer to mean values only) Graphiurus christyi. Similar head and body length (mean 97.6 mm). Ears longer (14.2 mm). Dorsal pelage colour sometimes similar, but most individuals have a rufous hue. Eye-mask usually more distinct. Tail absolutely and relatively longer (ca. 83% of HB). Skull averages slightly shorter (97.8 mm). Palate shorter (8.5 mm) both absolutely and relative to skull length. Anterior palatal foramina absolutely similar in length (3.0 mm), but absolutely and relatively wider in breadth posteriorly (2.2 mm). Auditory bullae similar in length (7.4 mm) and relative inflation. (Measurements listed are mean values from DR Congo; M. E. Holden unpubl.). No sympatry recorded, though both species have been collected in SW Cameroon and N DR Congo (Holden 1996). Graphiurus lorraineus. Smaller head and body length (mean 83 mm). Hindfeet (16.6 mm) shorter. Ears similar in size, but relatively larger. Dorsal pelage usually rufous, though sometimes brown. Anterior palatal foramina 2.6 mm long, 1.7 mm wide. Skull

Graphiurus surdus

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Family GLIRIDAE

Habitat Probably occurs in primary rainforest; may also be found in secondary forest (specimen labels). These dormice have been trapped on the forest floor (W.Verheyen pers. comm.), and in forests with vines (Robbins & Schlitter 1981). Abundance Little information. Most specimens are from localities that have been fairly well sampled. This suggests either low population densities, or that the species is trap-shy, or that traps have not been set in appropriate habitats. Remarks Arboreal. One individual collected by G. L. Bates from Bitye, Cameroon was ‘smoked out of a hollow tree’ (specimen label). This suggests that, like G. crassicaudatus, G. nagtglasii and G. lorraineus, some individuals nest in hollow trees.

Conservation

IUCN Category: Data Deficient.

Measurements Graphiurus surdus HB: 99.0 (87–110) mm, n = 8 T: 72.3 (65–82) mm, n = 6 HF: 20.8 (18–22) mm, n = 12 E: 12.3 (9–14) mm, n = 11 WT: 24.8 (18–34) g, n = 6 GLS: 27.6 (26.5–29.4) mm, n = 10 GWS: 14.6 (13.4–15.7) mm, n = 11 P4–M3: 3.2 (2.9–3.5) mm, n = 16 Cameroon, Equatorial Guinea, Gabon (Holden 1996) Key References

Holden 1996; Robbins & Schlitter 1981. Mary Ellen Holden

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Family DIPODIDAE

Family DIPODIDAE JERBOAS

Dipodidae Fischer de Waldheim, 1817. Mem. Soc. Imp. Nat., Moscow 5: 372. Allactaga (1 species) Jaculus (2 species)

Four-toed Jerboa Jerboas

p. 136 p. 137

the forelimbs are very short, have five digits and are rarely used for locomotion. The very long tail (ca. 140–170% of HB in African species) has a tuft of hairs at the tip and is used as a counterbalance during rapid running or hopping locomotion.When the long-legged species (Jaculus, Dipus, Allactaga) are resting on the hindlimbs, the end of the tail near the tip provides a tripod-like support. There are several modes of locomotion; the most typical is a fast bipedal jump using only the two hindlimbs (‘richochetal locomotion’). The skull is broad, and has an enlarged jugal plate, well-developed auditory bullae and laminate molars. The mandibles are weak without a bony inflection at the angle of the mandible, and perforations (fossae) on the angular process. Dental formula is: I 1/1, C 0/0, P 0/0, M 3/3 = 16 (P is 0/1 in some non-African species). Size categories of species in the family (based on mean head and body length) are given in the order Rodentia profile. All species of the Dipodidae live in open, usually arid, habitats where vegetation is sparse. The climate where dipodids live is extremely varied: species that live the cold deserts of Asia hibernate for 6–7 months during the coldest season of the year, and some species (Cardiocranius, Salpingotus, Pygeretmus) store fat in their tails. Species living in hot deserts are lethargic, and exhibit less activity, during the coolest part of the year. During the day they rest in burrows, which may be up to 3 m underground and extend for many metres; when inside the burrow, some species block the entrance with a plug of sand. Individuals emerge at night to forage on the ground. Species with long legs hop and run over the substrate with great speed and agility. The diet is principally seeds, bulbs and grass stalks; some species may feed occasionally on insects. Species that live in arid habitats (e.g. Jaculus spp.) possess kidneys that have special anatomical features to minimize water loss. Little information is available on the social behaviour in dipodids; they are usually found singly or in small groups; during cold weather, several individuals often huddle together in a nest in the burrow. Reproduction is seasonal, litter-size is mostly 2–6, and !! of some species may have more than one litter each breeding season. Fossil remains of dipodids are uncommon. Fossil jumping mice and jerboas are known from the Pliocene of Europe and from the Miocene and Pliocene of Asia. There are very few fossil remains of jerboas from Africa (Holden & Musser 2005). In Africa, there are two subfamilies and two genera:

The Dipodidae contains 51 species arranged in six subfamilies and 16 genera (Holden & Musser 2005).The family is distributed throughout the Palaearctic region, the Middle East and North Africa in forests, grasslands and steppe, and in hot and cold deserts. Only two genera and three species of this widespread family occur in Africa. The taxonomy of the family is controversial and uncertain. The family appears to be monophyletic (Holden & Musser 2005 and references therein) and to have diversified greatly in recent times.The family is divided into six subfamilies: Sicistinae (birch mice) with the single genus Sicista, Zapodinae (jumping mice) with Zapus, Eozapus and Napeozapus, Cardiocraniinae with Cardiocranius, Salpingotus and Salpingotulus, Euchoreutinae with Euchoreutes, Allactaginae with Allactaga, Allactodipus and Pygeretmus, and Dipodinae with Dipus, Eremodipus, Jaculus, Paradipus and Stylodipus. Only the Allactaginae and Dipodinae occur in Africa. The family as a whole is characterized by enlarged masseter muscles, which penetrate the infraorbital foramen (a character normally associated with hystricomorph rodents), an enlargement of the jugal to form a plate anterior to the orbit, which protects the eye, and the lack of a bony inflection on the angle of the mandible (which is characteristic of most murid rodents). Two of the subfamilies, the Sicistinae and Zapodinae, differ from the other four in many respects and some authorities place these subfamilies in a separate family, the Zapodidae; however, there is considerable overlap in characters between all the subfamilies and no clear dividing lines separating each subfamily. Because, on balance, Sicistinae and Zapodinae are similar to each other, with marked differences to the remaining subfamilies, the Dipodidae are here regarded as separate from the Zapodidae (Corbet & Hill 1986); this arrangement reduces the family Dipodidae to four subfamilies, 12 genera and 33 species. See Holden & Musser (2005) for further details. The best-known characters of Dipodidae (as understood here) are large auditory bullae, non-cuspidate low-crowned cheekteeth, long hindlimbs (ca. 50–56% of HB in African species), reduction in the number of digits, elongation of the metatarsal bones and partial or total fusion of these bones to a form a single ‘cannon bone’, which supports the weight of the body, a flattened nasal region reminiscent of that of a pig, large (or very large) ears and large dark eyes. The neck is short and the vertebrae are small and fused in most species. The metatarsal bones and the phalanges of the hindfeet Allactaginae: five digits on hindfoot (four in A. tetradactyla), three metatarsals (Digits 2, 3 and 4) fused to form cannon bone, are very specialized and varied in structure. In Cardiocranius there undersurface of hind digits with small hairs, three molars and one are five hind digits (Digit 1 being the shortest) and the metatarsal small premolar in each upper jaw; Allactaga (1 sp.). bones are not fused; in Salpingotus Digits 1 and 5 are lost and the metatarsals are not fused; in Allactaga the metatarsals of Digits 2, 3 Dipodinae: three digits on hindfoot; three metatarsals (Digits 2, 3 and 4) fused to form cannon bone, undersurface of hind digits and 4 are fused (now called the ‘cannon bone’) and Digits 1 and 5 with abundant strong bristles, three cheekteeth without premolar are very short and lie close and parallel to the cannon bone (in A. in each upper jaw; Jaculus (2 spp.). tetradactyla, Digit 1 on the inner side of the hindfoot is absent); and in Jaculus and Dipus the cannon bone is extremely elongated and D. C. D. Happold Digits 1 and 5 are absent. In all species with a cannon bone, the three hind digits are well developed. In contrast to the hindlimbs, 135

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GENUS Allactaga Jerboas Allactaga F. Cuvier, 1836. Proc. Zool. Soc. Lond., 1836: 141 [1837]. Type species: Mus jaculus Pallas, 1778 (= Dipus sibericus major Kerr, 1792).

Allactaga tetradactyla.

Widespread genus with 11 species in ‘cold deserts’ from Iran and Afghanistan to Manchuria, and in ‘hot deserts’ of the Middle East and NE Africa (Holden 1993). Holden & Musser (2005) place Allactaga as the only genus in the subfamily Allactaginae. The principal characters of the genus are: fusion of the metatarsal bones of Digits 2, 3 and 4 to form the ‘cannon bone’ (as in Jaculus). In most species, Digit 1 is short, although it is absent in A. tetradactyla (the only species of this genus in Africa). Digit 5 (outer digit) is short, ending half way along the length of the cannon bone. In general shape, Allactaga is reminiscent of Jaculus, except that the ears are much longer and often held upright from the head. Hindlimbs used for synchronized bipedal hopping, and for asynchronized bipedal trotting. Very long tail acts as counterbalance when hopping. Large eyes and ears. Pads of digits large and naked. The skull is characterized by a rudimentary premolar in each upper jaw (cf. Jaculus), perforations in the small angular process of the lower jaw, large auditory bullae but less developed than in Jaculus, well developed maxillary process, large orbit and wide zygoma (Figure 20). Baculum absent. The single African species, Allactaga tetradactyla, is present only in Egypt and Libya.

Figure 20. Skull and mandible of Allactaga tetradactyla (BMNH 14.3.6.1).

D. C. D. Happold

Allactaga tetradactyla FOUR-TOED JERBOA Fr. Gerboise tétradactyle; Ger. Vierzehen-Jerboa Allactaga tetradactyla (Lichtenstein, 1823). Verz. Doublet. Zool. Mus. Univ. Berlin, p. 2. ‘Libyan Desert between Siwa and Alexandria’ (= Egypt).

Taxonomy Originally described in the genus Dipus. Synonyms: brucii. Subspecies: none. Chromosome number: not known. Description Small jerboa with large rounded head, extremely long hindlimbs, four toes on each hindfoot, and very long tufted tail. Dorsal pelage pale orange streaked with black, tending to grey on the flanks; hairs grey at base, orange on terminal half, sometimes with black tip. Ventral hairs pure white, without white colouration extending onto rump at base of tail. Head similar in colour to dorsal pelage. Eyes large. Ears long and darkly pigmented. Hindlimbs,

especially metatarsals, very long (ca. 51% of HB). Digits 2, 3 and 4 fused to form cannon bone; Digit 1 absent; Digit 5 short, ending with short claw visible about half way along outer surface of cannon bone (contra Kingdon 1997:191; inner digit [Digit 1] shown on right cannon bone is in error). Undersurface of cannon bone and base of toes with blackish hairs. Tail very long (ca. 150% of HB), with terminal tuft of black hairs, each hair black at base with white tip. Skull as in genus profile. Nipples: not known. Geographic Variation

None recorded.

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Abundance Rare and localized. Much less common than J. orientalis, which lives in similar habitats. Remarks Terrestrial and nocturnal. Many of the adaptations of these jerboas are likely to be similar to those of the two species of Jaculus, but their rarity precludes detailed information. However, unlike Jaculus spp., Four-toed Jerboas appear to have a very limited ecological tolerance. Burrows are simple, 60–150 cm deep. Burrows are occupied only for brief periods; at other times unoccupied burrows of Greater Egyptian Jerboas are utilized (Hoogstraal 1963). Occasionally infested by the flea Xenopsylla nubica, and by four other species of fleas. Another flea species, Hopkinsipsylla occulta, is speciesspecific to A. tetradactyla, and is mostly found in the nests and not on the host itself. Only 25 animals (of 200 examined) were infected by fleas (Hoogstraal & Traub 1965b). Conservation IUCN Category: Vulnerable. This species is threatened with extinction because of its rarity, small geographical range and reclamation of its habitat for agriculture and development (Hoogstraal 1963). Allactaga tetradactyla

Similar Species Jaculus jaculus: Similar in general size, shorter ears (21 mm); three hind digits, Digits 1 and 5 absent. Jaculus orientalis. Much larger (HB: 137–160 mm; HF: 71–78 mm); three hind digits, Digits 1 and 5 absent. Distribution Endemic to Africa. Eastern part of Sahara Arid BZ. Regs and hamadas along the Mediterranean coast of Libya and Egypt west of the Nile Delta. Does not extend southwards into the Sahara Desert. Habitat Salt marshes and valleys in coastal regions; further inland, recorded from clay deserts, especially near barley fields, and Anabasis shrublands (Osborn & Helmy 1980).

Measurements Allactaga tetradactyla HB: 110 (102–119) mm, n = 19 T: 169 (154–180) mm, n = 17 HF: 56 (51–59) mm, n = 19 E: 41 (37–43) mm, n = 19 WT: 52 (48–56) g, n = 3 GLS: 28.9 (27.3–30.4) mm, n = 20 GWS: 20.9 (19.1–22.6) mm, n = 16 P4–M3: 5.9 (5.2–6.2) mm, n = 20 Auditory bulla: n. d. Egypt (Osborn & Helmy 1980) Key References

Hoogstraal 1963; Osborn & Helmy 1980. D. C. D. Happold

GENUS Jaculus Jerboas Jaculus Erxleben, 1777. Syst. Regni Anim. 1: 404. Type species: Mus jaculus Linnaeus, 1758.

Widespread genus with three species in arid habitats of North Africa and the Middle East, extending eastwards to western Pakistan and northwards to Turkmenistan and Uzbekistan (Holden 1993). The genus is placed in the subfamily Dipodinae, together with the nonAfrica genera Dipus (1 sp.), Eremodipus (1 sp.), Paradipus (1 sp.) and Stylodipus (3 spp.). Two species of Jaculus occur in Africa. The principal characters of the genus are elongated hindlimbs and fusion of the metatarsals of Digits 2, 3 and 4 to form a ‘cannon bone’ (as in Allactaga); Digits 1 and 5 absent. Hindlimbs used for synchronized bipedal hopping, and for asynchronized bipedal trotting. Very long tail acts as counterbalance when hopping. Eyes very large. Ears large, rounded at tip. Pads of digits small and covered with dense bristles. Lacks small premolar in upper jaw (cf.

Allactaga). Skull broad, auditory bullae greatly inflated and elongated (ca. 40% of GLS), zygoma absent (c.f. Allactaga), large orbit, and with perforations in small angular process of lower jaw (Figure 21). Baculum present. Members of the genus are highly adapted for life in sandy arid habitats where the climate is alternately very hot during the day and cool or cold at night. They are nocturnal, and spend the day in deep burrows. The diet is seeds and dried grass. They are independent of free water, primarily because they have efficient kidneys that produce very concentrated urine. Fossils of the genus are known from the late Miocene of Kazakhstan, late Pliocene of Morocco and Ethiopia, and PlioPleistocene of Kenya (details in Holden & Musser 2005). 137

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Family DIPODIDAE

Figure 21. Skull and mandible of Jaculus jaculus (HC 504).

Two species are present in Africa: one widespread and common in North Africa; the other confined to the Mediterranean coastal zone. The species are distinguished by body size and geographical distribution. D. C. D. Happold

Jaculus jaculus LESSER EGYPTIAN JERBOA Fr. Petite Gerboise d’Egypte; Ger. Kleine Ägyptische Springmaus Jaculus jaculus (Linnaeus, 1758). Syst. Nat., 10th edn, 1: 63. Giza Pyramids, Egypt.

Taxonomy Originally described in the genus Mus. About 19 subspecies (Misonne 1974) have been described throughout the extensive African range of the species. Individual countries may have several subspecies, e.g. Algeria five subspp. (Kowalski & RzebikKowalska 1991), Egypt four subspp. (Osborn & Helmy 1980), Libya eight subspp. (Ranck 1968) and Sudan two subspp. (Setzer 1956). Some of these taxa were used as specific names in the past, but now all are regarded as synonyms (Holden & Musser 2005). Jaculus deserti, as recorded by Ranck (1968) in Libya, is a synonym of J. jaculus (Osborn & Helmy 1980; Musser & Carlton 1993). Synonyms (of J. jaculus): airensis, arenaceus, butleri, centralis, collinsi, cufrensis, favonicus, gordoni, microtis, sefrius, tripolitanicus, vulturnus, whitchurchi; (of J. deserti): favillus, fuscipes, rarus, schlüteri, vastus. Subspecies: none (but see Geographic Variation). Chromosome number: 2n = 50; FN = 90 (Senegal; Granjon et al. 1992); 2n = 48 (Niger; Dobigny et al. 2002b). Description Small pale-coloured jerboa with large rounded head, extremely long hindlimbs and very long tufted tail. Dorsal pelage long and silky, pale sandy-brown to sandy-rufous; hairs grey at base, sandy or brownish on terminal half, sometimes with black tip. Black-tipped hairs more numerous in individuals from moister habitats resulting in a streaked darker-coloured pelage. Ventral pelage pure white. White colouration extends dorsally onto rump forming white band at base of tail. Head similar in colour to back, broad and rounded, with small muzzle, many vibrissae, large eyes and large rounded ears. Hindlimbs, especially metatarsal bones, very long (ca. 56% of HB); three hind digits well covered on undersurface with whitish bristles. Forelimbs small, held close

Jaculus jaculus

under chin. Tail very long (ca. 170% of HB), basal two-thirds covered with short sandy-coloured hairs; terminal third with long hairs forming a large tuft, black on basal half, white on terminal half. Skull as in genus profile. Females tend to be heavier than "" (adult mean weights at Khartoum: !! [non-pregnant] 60 g; "" 49 g). Nipples 2 + 2 = 8.

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Jaculus jaculus

Geographic Variation Colour of dorsal pelage (the basis for most subspecies descriptions) appears to be related to the colour of the soil or sand and the degree of aridity. Individuals from C Sahara are very pale sandy- or creamy-white (e.g. airensis), those from moister environments are sandy-brown to orange-brown with greater numbers of black-tipped hairs. Mean HB length varies with locality (and ‘subspecies’): e.g. 105 mm (butleri, Khartoum); 110 mm (jaculus, Egypt); 111 mm (flavillus, Egypt). Similar Species J. orientalis. Larger (HB: 137–160 mm; HF: 71–78 mm); North African coastal regions only. Allactaga tetradactyla. Similar in general size, ears longer (40 mm), four digits including short Digit 5 on outside of cannon bone; coastal regions Libya and Egypt only; rare. Distribution Sahara Arid and Sahel Savanna BZs. Recorded from S Morocco, Mauritania and N Senegal in the west to Egypt and Sudan in the east. Not recorded in the coastal regions of Morocco north of the High Atlas and Saharan Atlas mountains of Morocco and Algeria. Isolated populations in coastal Eritrea, coastal Ethiopia and N Somalia. Perhaps occurs in Djibouti. There is some evidence that, since the 1970s, the distribution of jerboas has been moving progressively southwards within the Sahel Savanna BZ and into the Sudan Savanna BZ as a result of desertification of the savannas by humans. Also recorded from Israel, Iraq, Iran, Syria, Pakistan and Saudi Arabia. Habitat Lesser Egyptian Jerboas live in a variety of open sparsely vegetated arid and semi-arid habitats, including loose sandhills and hillocks, sandy plains and wide sandy wadis. Also occur on flat solid substrates, such as coastal regions and higher altitude plateaux (up to ca. 1500 m), provided the sand or soil is suitable for burrowing. Also recorded near fields of barley, and in Anabasis steppe country, along the Mediterranean coast. Not found in rocky habitats or on jebels.

a

Abundance In most habitats, Lesser Egyptian Jerboas are never abundant and distribution is patchy; tend to be rare over much of their geographic range especially where soil type is unsuitable, rainfall is very low and where food resources are limiting. Usually never abundant in Egypt (Osborn & Helmy 1980). Comparatively common in selected habitats near Khartoum (Happold 1967a, c, 1975a), and recorded as ‘common on the meidan’ in Somalia (specimens in BMNH, collected 1912). Where the three species of jerboas occur parapatrically, as in parts of Egypt, this species is rare (Hoogstraal & Traub 1965b). Adaptations Nocturnal and terrestrial. Lesser Egyptian Jerboas emerge from burrows after dusk. They exhibit several methods of locomotion above ground (Figure 22) including walking on all four limbs when foraging for food, hopping bipedally and slowly when ‘pottering’, and hopping very quickly when escaping from danger. Occasionally they run with a fast bipedal trot (Happold 1967a). The tail is held out horizontally as a counterbalance when moving on the hindlimbs. They gain friction on the substrate from bristles on undersurface of the hindfeet. When disturbed or pursued, they hop extremely rapidly (up to ca. 30 km/h), often changing direction quickly and erratically. Mobility is necessary to find adequate food, and speed and manoeuvrability are essential to escape from predators in open environment. Lesser Egyptian Jerboas groom by licking their fur and by lying in a little concavity in the sand with the hindlimbs stretched out and rubbing the body backwards and forwards (‘sandbathing’). Lesser Egyptian Jerboas dig burrows in the sand using their teeth and forefeet to excavate the burrow, and push the excavated sand out of the burrow with the blunt flattened nose and muzzle (nostrils can be closed), and scatter it with the hindlimbs. Burrows are simple,

b

e

Figure 22. Behavioural chararacteristics of Jaculus jaculus. (a) movement on all fours; (b) medium speed; (c) fast speed; (d) high jump; (e) sleeping; (f) sitting submissive; (g) standing alert; (h) washing rump; (i) cleaning tail; (j) sandbathing; (k) social contact between three jerboas; (l) pushing sand with nose (after Happold 1967a).

Their ability to live in many arid zone habitats and climates is one reason for their wide distribution.

c

f

h

k

d

g

i

j

l

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usually with two entrances, and vary in depth according to season; at Khartoum, winter burrows are shallow (20–25 cm deep with almost horizontal passages, and in summer, burrows are deeper (70–75 cm) with steeply sloping passages (Ghobrial & Hodeib 1973). After an individual has entered its burrow, the entrance is plugged from the inside with sand so that the entrance is practically invisible. Small sleeping chamber at base of burrow is lined with grass. Temperature and relative humidity in the burrow is relatively constant (temperature 26–28 °C in cool season when air temperature is 5–27 °C, and 37 °C in hot season when air temperature is 28–46 °C; humidity 60–100%) and does not experience the temperature or humidity fluctuations of the sand surface. Skull has enlarged zygomatic plates, which protect the orbits when anterior part of head is used for pushing sand. Auditory bulla is large (ca. 40% of GLS) and greatly inflated; it assists detection of vibrations in sand when animal is in the burrow, and airborne sounds when it is on the surface. Lesser Egyptian Jerboas can obtain all their water from their food (but will lick dew and standing water when available), and possess many special anatomical and behavioural features for conserving water. The kidney produces only small amounts of extremely concentrated urine (4320 mMol/l), about double the maximum concentration produced by a white rat. In experimental conditions, without free water and feeding on barley, an individual lost ca. 30% of body weight after three weeks (Schmidt-Nielsen 1964). In this respect, these jerboas are not as well adapted to lack of water as some other species of desert rodents (e.g. some Gerbillus and Meriones spp.). Other ways to reduce water loss and energy loss include nocturnal activity, resting in relatively cool and moist burrows, sleeping curled in a ball to reduce water loss from the lungs, feeding on foods with high water content when available (moist vegetation, bulbs) and seasonal changes in activity (Happold 1967a). In cool weather, Lesser Egyptian Jerboas rest in their burrows curled into a tight ball, tucking the head close to the abdomen and covering the long hindlimbs with the body; activity above ground is greatly reduced. In hot weather, they rest with legs and tail stretched away from the body, and sometimes lick saliva on to the fur to increase heat loss. Foraging and Food Foraging takes place only at night, when seeds and stems are collected from the substrate with short forelimbs. Lesser Egyptian Jerboas sit in a crouched position while eating, with the whole length of the cannon bone resting on the ground, and the food is held with the forelimbs. Near Khartoum (and presumably elsewhere) the diet consists of small seeds, dried desert grasses and roots, as well as bulbs and corms in the dry season. Sometimes the fleshy leaves of some succulent plants are also eaten. After rainfall, Lesser Egyptian Jerboas feed on newly germinated grass sprouts. Their weak jaws prevent them from feeding on large hard seeds. There is no evidence that these jerboas collect and store food (cf. J. orientalis and many species of Gerbillus). Social and Reproductive Behaviour In the desert, Lesser Egyptian Jerboas are seen singly (or occasionally in pairs or trios). They show many of the behavioural characters of solitary species: agonistic behaviour (fighting, chasing) when confined with an unfamiliar individual; a tendency to nest alone and to have limited

interactions with other individuals. In an established group, members show tolerance towards each other. No information available on reproductive behaviour. Reproduction and Population Structure Reproduction is seasonal. Near Khartoum, young individuals were found from Sep to Dec, and in Feb and Mar (1964–66); this suggests two periods of pregnancies, one in Jun and Jul during the annual ‘wet season’ (average 150 mm/year), and the second in Oct–Dec when monthly temperatures are declining and food is still relatively abundant (Happold 1967a). In other years (1970–72), reproduction was almost continuous with peaks in Sep–Nov and Dec–Feb (Ghobrial & Hodeib 1973). Reproduction is probably opportunistic, and dependent on availability of nutritive food (and hence rainfall). Gestation: 44–46 days. Litter-size/number of embryos (regardless of season): 3.4 (2– 5, mode 3, n = 18; Happold 1967a). Mean litter-size at Khartoum varies according to season: 4.8 in Sep (end rains, food abundant), 3.0 in Dec (cool temperature, food less abundant) and 1.83 (Jun, hot temperatures, food scarce) (Ghobrial & Hodeib 1973). Young born naked; development is slow for a rodent of this size: hair develops by Day 22; eyes open Day 38. Hindfoot and auditory bullae develop rapidly, reaching adult size quicker than other structures. At birth, cannon bone formed of three separate metatarsal bones, each 9 mm in length; at Day 40, bones fused and 50 mm in length, but unable to support weight of body. Weight when first active above ground ca. 20 g (= 35% adult weight, age 50–60 days) (Happold 1970a). The relative age of an individual can be estimated by the wear on the cheekteeth: youngest animals are Age-class 3 (least wear on molars) and oldest animals are age-class 17 (maximum wear) (Happold 1967a). At Khartoum, in Sep–Dec (1964–66), after the rains and the first breeding period, there were individuals of all age classes, suggesting three age-groups: (a) age-classes 3–8, weight 40 g, born previous Oct–Dec, age 9–12 months; and (c) age-classes 10– 17, wt >40 g, age more than 12 months. By Jan–Mar, most of the individuals in age-group (a) had moved to age-group (b), and many of the oldest individuals in age-group (c) had disappeared from the population. (No detailed information for Apr–Aug.) Individuals of age-classes 14–17 may be 3–4 years of age. Only 25% of individuals attained an age-class of 10 or above (Happold 1967a). Predators, Parasites and Diseases Remains of Lesser Egyptian Jerboas have been found in owl pellets in Algeria (Kowalski & Rzebik-Kowalska 1991) and they are likely to be preyed upon by owls in many parts of their geographic range. In Egypt, they are hosts to many species of fleas, the commonest being Xenopsylla nubica, X. cheopis and Synosternus cleaopatrae; none is specific to J. jaculus and are found also on other small rodents. Infection rate is low; infested individuals typically host 3–4 fleas (Hoogstraal & Traub 1965b). The blood parasite Hepatozoon balfouri infected 41% of jerboas examined in Egypt (n = 370) as well as many other species of Egyptian small mammals (Hoogstraal 1961). Conservation IUCN Category: Least Concern. The wide distribution of Lesser Egyptian Jerboas, and the sparseness of humans in their habitats, suggest that they are not threatened.

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Jaculus orientalis

M1–M3: 5.3 (4.8–5.6) mm, n = 77 Auditory bulla: 13.2 (12.8–13.5) mm, n = 10* Egypt (Osborn & Helmy 1980) *Sudan (J. j. butleri; BMNH)

Measurements Jaculus jaculus jaculus HB: 110 (98–118) mm, n = 78 T: 181 (160–203) mm, n = 76 HF: 62 (56–66) mm, n = 80 E: 21 (19–23) mm, n = 80 WT: 55 (45–73) g, n = 60 GLS: 31.3 (30.2–32.7) mm, n = 77 GWS: 22.6 (19.9–24.4) mm, n = 71

Key References Happold 1967a, 1970a, 1975a; Osborn & Helmy 1980; Ghobrial & Hodeib 1973. D. C. D. Happold

Jaculus orientalis GREATER EGYPTIAN JERBOA (ORIENTAL JERBOA) Fr. Gerboise Orientale; Ger. Orientalische Springmaus Jaculus orientalis Erxleben, 1777. Syst. Regn. Anim. 1: 404. ‘In the mountains separating Egypt from Arabia’ (= Egypt).

Taxonomy Several subspecies have been described (orientalis, gerboa, mauritianus) but none is recognized by Corbet (1978). This species was referred to (incorrectly) as Dipus aegyptius by Kirmiz (1962). The ‘Desert Rats’ of the North African campaign in World War II were named after this species. Synonyms: bipes, gerboa, locusta, mauritanicus. Subspecies: none. Chromosome number: not known. Description Medium-sized jerboa with large rounded head, extremely long hindlimbs and very long tufted tail. Dorsal pelage brownish-orange, becoming paler on flanks; hairs grey at base, orange-brown on terminal half, sometimes with black tip. Ventral pelage pure white. Very similar in general characters to J. jaculus. Hindlimbs, especially metatarsal bones, very long (ca. 51% of HB); three hind digits well covered on undersurface with whitish bristles. Forelimbs short. Tail very long (ca. 146% of HB), ending with large tuft, black at base and white at tip. Nipples: not known.

Geographic Variation

None recorded in Africa.

Similar Species J. jaculus. Considerably smaller (e.g. HB: 98–118 mm; HF: 56– 66 mm); widespread in desert habitats. Allactaga tetradactyla. Considerably smaller but with longer ears (37–43 mm), four hind digits including short Digit 5 on outside of cannon bone; coastal regions Libya and Egypt only; rare. Distribution Mediterranean Coastal BZ and coastal regions of Sahara Arid BZ in Libya and Egypt. Recorded from Morocco, Algeria and Tunisia, and the high plateaux of E Morocco and Algeria south to about 34° S, and in the regs and hamadas near the coast from W Libya to Egypt west of the Nile Delta. Prefers more humid environments to J. jaculus and does not extend southwards into the Sahara Desert. Also recorded from Sinai and S Israel. The geographical range overlaps with that of J. jaculus in a few areas of Algeria, Libya and Egypt. Habitat Salt marshes with Salicornia bushes; limestone slopes covered by Suada bushes above the salt marshes; coastal dunes; gardens, meadows, olive growths and old barley fields covered with annual plants (Osborn & Helmy 1980). Compared with J. jaculus, these jerboas live in habitats that are less arid and have much more vegetation. Individuals found up to 1500 m on the High Atlas and Saharan Atlas mountains. Does not live in the sclerophyllous forests of Morocco, Algeria and Tunisia. Abundance Very varied and dependent on food availability and character of the habitat. In Egypt, recorded as ‘1–50 jerboas per 0.8 km’, and ‘more or less common’ on the desert slopes near the Mediterranean Sea from Egypt west of the Nile Delta to Libya and Algeria (Hoogstraal 1963). In Algeria, common on the high plateaux (Kowalski & Rzebik-Kowalska 1991).

Jaculus orientalis

Adaptations Terrestrial and nocturnal; similar in many ways to J. jaculus but confined to the Mediterranean coastal regions and hence less well adapted for desert environments. Greater Egyptian Jerboas have two main forms of locomotion: a bipedal hop with asynchronized foot support (rather like a gallop), and a bipedal trot, which is used for moving around and through bushes and shrubs (cf. J. jaculus) (Schröpfer et al. 1985). 141

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Burrows usually 1–2 m in length, dug in hard ground, often on the slope of a hill (Kirmiz 1962). Burrow ends in a nest chamber lined with camel hair, shredded pieces of cloth or shredded vegetation. Some burrows have a food chamber (see below). Position of burrows changes seasonally: higher on hillsides during winter rains, and on lower ground close to fields in summer.When occupied, burrows are blocked with plugs of sand. Water conservation has been measured by the percentage of body water that is ‘turned over’ in 24 h, and the volume and concentration of the urine. In cold conditions (Ta = 8 °C), water turnover (regardless of food) is ca. 10%/24 h, and in hot weather (Ta = 32 °C), it is reduced to 5%/24 h (when fed only on barley) and 9%/24 h (when fed on barley and ‘salad’). Greater Egyptian Jerboas are able to produce small amounts of concentrated urine when required. For example, when fed on barley and ‘salad’, at Ta = 8 °C, urine production was 8.7 ml urine/24 h with an osmotic concentration of 714 mOsm/L, and at Ta = 32 °C they produced 0.6 ml/24 h with an osmotic concentration of 2227 mOsm/L (Baddouri et al. 1985). In these respects, this species of jerboa is not as efficient at water conservation as J. jaculus. In the winter, on parts of Mediterranean coast and high plateaux, air temperatures at night may be below freezing although the temperature in burrows is ca. 10 °C. On the high plateaux of Morocco (1500 m), Greater Egyptian Jerboas taken from burrows in winter were immobile, breathing was spasmodic and body temperature was 10–11 °C. On exposure to an air temperature of 17 °C, shivering commenced and body temperature gradually increased to normal in ca. 4 h. Thus Greater Egyptian Jerboas show true hibernation, and are able to rewarm using endogenous mechanisms (El Hilali & Veillat 1975). They appear to be more tolerant of cold than Common Jerboas (Hooper & El Hilali 1972). Foraging and Food Foraging is similar to that in Lesser Egyptian Jerboas but because of their large size, Greater Egyptian Jerboas are able to consume larger seeds. Food is mainly sprouting vegetation, plants, roots and barley grains (Kirmiz 1962). Various succulent shrubs such as Salicornia and Suada may be browsed by individuals inhabiting salt marshes (Osborn & Helmy 1980). Dates, barley and the seeds of several wild plants have been found in burrows.

Social and Reproductive Behaviour Sociable, and not usually encountered as solitary individuals (cf. J. jaculus) (Osborn & Helmy 1980). Parapatric (and perhaps syntopic) with Allactaga tetradactyla on the coasts of Libya and Egypt. Reproduction and Population Structure Limited information. In Egypt, breeding occurs during winter (Nov–Feb) and summer, and less frequently during spring (Hoogstraal 1963). Births recorded in Feb, Apr and early Jul. Litter-size: 3, 4, 7 in Algeria (Kowalski & Rzebik-Kowalska 1991); 2, 3, 4, 5 (mode 3) in Egypt (Flower 1932). Predators, Parasites and Diseases Remains found in owl pellets in Algeria (Kowalski & Rzebik-Kowalska 1991). In Egypt, a flea Mesopsylla tuschkan is found commonly in nests and on animals; nine other species of fleas have been recorded occasionally (Hoogstraal & Traub 1965b). Conservation

IUCN Category: Least Concern.

Measurements Jaculus orientalis HB: 148 (137–160) mm, n = 31 T: 224 (195–243) mm, n = 31 HF: 75 (71–78) mm, n = 31 E: 33 (28–35) mm, n = 30 WT: 134 (108–147) g, n = 17 GLS: 36.9 (36.2–38.0) mm, n = 26 GWS: 28.3 (27.1–30.0) mm, n = 24 M1–M3: 6.7 (6.2–7.4) mm, n = 25 Auditory bulla: 15.6 (15.2–16.7) mm, n = 7* Egypt (Osborn & Helmy 1980) *Egypt (BMNH) Key References

Osborn & Helmy 1980; Kirmiz 1962. D. C. D. Happold

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Family SPALACIDAE BLIND MOLE-RATS, AFRICAN ROOT-RATS, ZOKORS AND BAMBOO-RATS Spalacidae Gray, 1821. London Med. Repos. 15: 303.

Ehrenberg’s Mole-rat Root-rats

Spalax (1 species) Tachyoryctes (2 species)

p. 145 p. 148

This small family (six genera and 36 species) represents an old lineage of Muroidea whose extant members have acquired striking fossorial adaptations and evolved into predominantly subterranean niches. There are four subfamilies: Myospalacinae (Zokors), Rhizomyinae (Bamboo-rats), Spalacinae (Blind Mole-rats) and Tachyoryctinae (African Root-rats). The first two subfamilies are extralimital to Africa, myospalacines occurring in Siberian Russia and northern China, and rhizomyines in north-eastern India, southern China and the Malay Peninsula. The latter two subfamilies, the spalacines (one genus, one species) and tachyoryctines (one genus, two species), are geographically localized within Africa. The only African species of Spalax is found along the Mediterranean coastal region, and the two species of Tachyoryctes live in the highlands of Ethiopia and East Africa. Although each subfamily is readily diagnosed by unique traits (Carleton & Musser 1984), underground life in burrows has entailed the evolution of a suite of morphological, physiological, sensory and behavioural specializations common to subterranean forms in other rodent families and suborders (Nevo 1979, 1999, Nevo & Reig 1990, Lacey et al. 2000). General characters of Spalacidae include a cylindrical body shape with broad head and massive cervical musculature; very soft and fine pelage; small or vestigial pinnae and eyes; short or inconspicuous tail; short, stocky limbs and powerful appendicular musculature; enlarged claws, especially on the forelimbs; hypertrophy and procumbency of the lower incisors for chisel-tooth digging; and robust, hypsodont molars associated with their largely herbivorous diet of roots, bulbs and rhizomes (Carleton & Musser 1984, Stein 2000). Dental formula: I 1/1, C 0/0, P 0/0, M 3/3 = 16 (Table 15).

Whether such phenotypic resemblance connotes phylogenetic relationship or evolutionary convergence, it has accounted for most of the differences in the classification of these rodents. Most taxonomic arrangements, implicitly or explicitly, reflect the view that the similar fossorial adaptations have evolved in parallel and have variously grouped the six genera within three or four separate subfamilies and in one to three families (see Topachevskii 1969, Carleton & Musser 1984 for reviews). In the prevalent classificatory treatment, Spalacidae and Rhizomyidae (including Tachyoryctes) have been retained as small outlying families separate from core Cricetidae and/or Muridae (e.g. Miller & Gidley 1918, Ellerman 1940, Simpson 1945, Pavlinov et al. 1995). Some palaeontologists have echoed a similar viewpoint and have considered that Spalacidae and Rhizomyidae (a non-African taxon) originated independently from different muroid stocks (Flynn et al. 1985). As early as 1899, however, Tullberg interpreted their shared resemblance as phylogenetic relationship and placed Myospalax, Spalax, Rhizomys and Tachyoryctes in Spalacidae, separate from his Cricetidae and Muridae. Monophyletic union of these fossorial genera has earned substantial support from recent studies, including evidence from morphology of the cephalic arterial system (Bugge 1971, 1985) and repeatedly from phylogenetic reconstruction based on nuclear gene sequences (Robinson et al. 1997, Debry & Sagel 2001, Michaux et al. 2001, Jansa & Weksler 2004). These data collectively portray myospalacines, rhizomyines, spalacines and tachyoryctines as a clade (Spalacidae) that is a sister-group to representatives of all other muroid families so far investigated (Calomyscidae, Cricetidae, Muridae, Nesomyidae) and infer an early divergence (and possibly one of the earliest divergences) of Spalacidae within Muroidea from a middle or late Oligocene ancestral stock.

Table 15. Subterranean rodents in Africa. Number of upper cheekteeth

Upper cheektooth row

Upper incisor teeth

Infraorbital foramen

Eyes

Tail

Spalacidae: Spalax

3 (M1, M2, M3)

Diverge anteriorly

Orthodont, ungrooved, not extrabuccal

Large, well developed

Absent (non-functional)

Absent externally

Spalacidae: Tachyoryctes

3 (M1, M2, M3)

Converge anteriorly

Pro-odont, ungrooved, slightly extrabuccal

Large, well developed

Very small

Bathyergidae: Bathyergus

4 (P4, M1, M2, M3)

Parallel

Pro-odont, grooved, strongly extrabuccal

Very small, round

Small

Bathyergidae: Heliophobius

6 (but usually 4 or 5; left side may differ from right)

Parallel

Pro-odont, grooved, strongly extrabuccal

Very small, round

Small

3 (sometimes 2)

Parallel

Pro-odont, ungrooved, strongly extrabuccal

4 (P4, M1, M2, M3)

Parallel

Pro-odont, ungrooved, strongly extrabuccal

Very small, round Small, teardrop or ellipitcal shape

Family: genus

Bathyergidae: Heterocephalus Bathyergidae: all genera except Heterocephalus

Very small Small

Short, visible externally (20–30% of HB); well haired Short, visible externally (18–24% of HB); stiff bristles Very short, visible externally (9% of HB); stiff bristles Moderately long (ca. 50% of HB); naked Short, visible externally (7–18% of HB), stiff bristles

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Although Oligocene fossils are so far unknown, the documented fossil history of Spalacidae is suitably deep in time to be consistent with a middle to late Oligocene origin. The earliest indisputable records among the four subfamilies are early Miocene, around 19–20 mya in Asia Minor (Debruijnia, Spalacinae) and southern Asia (Prokanisamys, Rhizomyinae) (see Flynn 1990, Nevo 1999 and Cook et al. 2000 for palaeontological summaries). In Africa, the earliest occurrences of the family originate from middle to late Miocene strata of Kenya (as Pronakalimys and Nakalimys; Flynn & Sabatier 1984, Tong & Jaeger 1993) and the late Miocene of Namibia (as Nakalimys and Harasibomys; Mein et al. 2000a). Although dental contrasts between living spalacid genera are highly distinctive, it is noteworthy

that molar similarities among certain African Miocene forms render assignment to subfamily – whether rhizomyine, spalacine, or tachyoryctine – as indefinite (Mein et al. 2000a). Spalacids were more geographically widespread and taxonomically diverse during the late Miocene through Pliocene (see Flynn 1990, Ünay 1999, Cook et al. 2000), a comparison that reinforces the impression that the relatively few extant spalacids, so strongly differentiated from one another, are relicts from a much older muroid radiation. Two subfamilies of Spalacidae occur in Africa: Spalacinae (1 genus, 1 species) and Tachyoryctinae (1 genus, 2 species). Guy G. Musser & Michael D. Carleton

Subfamily SPALACINAE – Mole-rats Spalacinae Gray, 1821. London Med. Repos. 15: 303.

The Spalacinae encompasses a single living genus, Spalax, whose range stretches from SE Europe and SW Asia around the Black and Caspian Seas, through Asia Minor and the near Middle East, to North Africa (see Nevo et al. 2001). The recent distribution broadly follows the hilly, uplifted region that corresponds to the ancient basin of the Mediterranean Sea and is generally concordant with that of known spalacine fossils (Ünay 1999). Conventionally, eight living species of Spalax are recognized (e.g. Topachevskii 1969, Musser & Carleton 1993), but this number may grossly underestimate the specific diversity (see later). Only a single species, S. ehrenbergi, occurs in Africa; its distribution is limited to the Mediterranean coastal region

Figure 23. Skull and mandible of Spalax ehrenbergi (RMCA no number).

of W Egypt and Libya (Lay & Nadler 1972). As currently understood, these North African populations are separated from those of S. ehrenbergi in the southern Levant by the Nile Delta and arid Sinai Desert (rainfall 510 mm. Occurs at altitudes above 1200 m, and reaches >4000 m in Kenya (Mt Kenya, Aberdare Ranges) and Tanzania (Mt Kilimanjaro). Habitat Deep well-drained soils in savanna grasslands, open forests, afroalpine regions, agricultural fields and gardens. Abundance Patchy and disjunct, but may be very common in suitable localities. Densities can be very high in cultivated land. Densities of 80 individuals/acre (= 200/ha) in a grassed area at Nairobi, Kenya (Jarvis 1973a), and of one individual/144 m2 in E DR Congo (Rahm 1980) have been recorded. One field (100 m × 80 m) in E DR Congo contained 17 adult "", 16 adult !!, nine juvenile "" and eight juvenile !! (Rahm 1980). Adaptations Subterranean. African Root-rats live in a system of burrows 15–44 m long, comprised of foraging burrows 15–30 cm below ground, a slightly deeper nest chamber and a deeper bolt-hole (n = 20 burrow systems; Jarvis & Sale 1971, Rahm 1980). The nest chamber includes a sleeping area, a food store and a toilet area. The sleeping area, a hollow ball of grass and roots, lies close to the entrance of the nest chamber. The small food stores containing grass rhizomes, roots and geophytes are at the sides of the nest. The toilet area, behind the sleeping area, contains composting faeces and old nesting material, which generates heat. A rich invertebrate fauna, including 151

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pseudoscorpions that feed on mites, is found in the nest chamber. African Root-rats dig by biting at the soil with their incisors. While digging, an animal will periodically turn round, and use one side of its head, and the foot of the same side, to hold the soil as it is pushed up a side branch leading to the surface. Once a mound has been completed, the side branch is thoroughly sealed. African Root-rats are diurnal, and active between 10:00h and 19:00h. Less than 25% of the 24-h day is spent out of the nest (Jarvis 1973b). They feel their way along the burrow by making small lateral movements of the head so the facial vibrissae brush the sides of the burrow. Unlike species of Bathyergidae, they rarely move backwards. They can rapidly turn around by curling on the side and walking round their almost stationary hindquarters. Foraging and Food Herbivorous. The diet includes a wide variety of plants including grass rhizomes, stems and leaves, herbs, roots and storage organs of geophytes. African Root-rats may either dig up under a plant and pull the entire plant into the burrow, or forage around the periphery of an open hole. In this latter instance, they need to keep their eyes wide open and brace the hindfeet in the hole ready to effect a rapid retreat if alarmed. A foraging hole is plugged with a small and very characteristic heap of soil when foraging is complete (Jarvis & Sale 1971). Prior to eating small items of food, animals will grasp the food with the incisors and lightly brush the food with their cupped hands. Unless the item is very large, they walk forwards in the burrow while transporting food to the nest where it is eaten or stored. Stored bulbs and tubers are not disbudded when they sprout. African Root-rats do not drink water, obtaining their water requirements from their food. Social and Reproductive Behaviour African Root-rats aggressively defend their burrow systems. They communicate seismically through the soil by tapping on the burrow floor with their upper incisors (Jarvis 1969a). Tapping consists of 3–10 taps, a pause and then a repeat of the sequence. Neighbouring animals will tap in response to each other’s taps. When threatened, African Root-rats adopt an aggressive posture: the head is thrown back, jaws are agape and the feet are widely spaced and held stiff; the animal lunges forwards, snorting or squeaking and chattering its teeth. During aggressive encounters, or when alarmed, "" produce a strong musk-like odour from scent glands situated ventral to the eye and ear; !!, in contrast, have very small glands. The position of the glands is clearly indicated by hairs that are stuck together by the secretions. The defensive posture is similar to the aggressive posture, but lacks vocalizations or tooth chattering. If completely submissive, an animal will lower its head and retreat (Jarvis 1969a). Courtship is preceded by both animals tapping on the floor of their burrows. On meeting, " emits a soft high-pitched twitter, the animals repeatedly lock incisors and ! will also squeak and gnash her teeth. Eventually ! turns, adopts the lordosis position, and " mounts and bites her neck; during copulation, " kneads flanks of ! with his hindfeet.Throughout copulation, which lasts 2–3 minutes, ! squeaks. Further copulations, which may continue at intervals for two days, are initiated by !. Between copulations, " often rubs genital region on the floor, possibly marking the area with secretions from large bilobed preputial glands.The cellular structure of the two lobes is different: one produces a more oily secretion than the other and it is possible that they serve different functions. Unlike during the aggressive encounters, strong musk-odours are not produced during courtship (Jarvis 1969a).

Reproduction and Population Structure African Root-rats breed throughout the year, and are polyoestrous. Gestation: 46–49 days (Rahm 1969a). The ovary of pregnant !! contains accessory corpora lutea, and !! with full-term embryos have 2.5 corpora lutea per embryo. Foetal resorption occurs during pregnancy (Jarvis 1969b, Rahm 1980). Mating and a second pregnancy can occur during lactation. Litter-sizes are small: 1.65 (1–3) at Nairobi, 1.2 (1–3) at Nakuru, Kenya (Jarvis 1969b); 1.39 (1–4) at Kivu, E DR Congo (Rahm 1969a, 1980). At birth, young weigh 11–18 g (n = 14), are hairless and toothless. Fine hairs and small white incisors appear by Day 4. First solid foods eaten ca. Day 15–20. First leave nest Day 15– 21. Eyes open Day 21–28. Digging and carrying food Day 24–30. Weaned ca. Day 35. Inter-sibling sparring begins at Day 37–60 and establishment of own burrow by Day 80.The vaginal closure membrane of young !! breaks down when 3 months old (Jarvis 1969a). Sex ratio at birth is parity (48.6%": 51.4% !, n = 3517), but biased towards !! in adults (58%, n = 9583) in E DR Congo (Rahm 1980). A similar bias towards !! occurred also in Kenya (Jarvis 1969a). Predators, Parasites and Diseases Predators include Barn Owls (Tyto alba), diurnal birds of prey, small carnivores and snakes. These probably capture African Root-rats as they forage above ground in the vicinity of open holes. Ectoparasites include 15 species of fleas and 12 species of mites (Rahm 1980). Conservation IUCN Category: Least Concern. African Root-rats are considered agricultural pests in crops of cassava, sweet potato, peanuts, lucerne and maize. They also damage the roots of young trees and disfigure lawns and golf courses. Measurements Tachyoryctes splendens HB (""): 200 (170–215) mm, n = 50 HB (!!): 189.5 (159–205) mm, n = 55 T (""): 62.7 (53–77) mm, n = 50 T (!!): 60.4 (51–77) mm, n = 55 HF (""): 31.8 (28–35) mm, n = 50 HF (!!): 30.3 (27–34) mm, n = 55 E (""): 11.3 (9–15) mm, n = 50 E (!!): 10.5 (9–12) mm, n = 55 WT (""): 248 (180–305) g, n = 50 WT (!!): 218 (140–315) g, n = 55 GLS (""): 45.9 mm, n = 20* GLS (!!): 43.6 mm, n = 20* GWS (""): 33.2 mm, n = 20* GWS (!!): 32.1 mm, n = 20* M1–M3 (""): 8.6 mm, n = 20*† M1–M3 (!!): 8.3 mm, n = 20*† Body measurements and weights: Nairobi, Kenya (J. U. M. Jarvis unpubl.) Skull measurements: E DR Congo (Rahm 1980) *Mean only; minimum and maximum values not available †Number of molar teeth variable – see genus profile Key References Rahm 1980.

Jarvis 1969a, b, 1973a, b; Jarvis & Sale 1971; J. U. M. Jarvis

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Family NESOMYIDAE POUCHED RATS AND MICE, SWAMP MOUSE, CLIMBING MICE, LARGE-EARED MOUSE, FAT MICE, WHITE-TAILED RAT AND ROCK MICE Nesomyidae Major, 1897. Proc. Zool. Soc. Lond. 1897: 718. Cricetomyinae (3 genera, 5 species) Beamys (1 species) Cricetomys (2 species) Saccostomus (2 species) Delanymyinae (1 genus, 1 species) Delanymys (1 species) Dendromurinae (6 genera, 23 species) Dendromus (11 species) Dendroprionomys (1 species) Malacothrix (1 species) Megadendromus (1 species) Prionomys (1 species) Steatomys (8 species) Mystromyinae (1 genus, 1 species) Mystromys (1 species) Petromyscinae (1 genus, 4 species) Petromyscus (4 species)

Pouched Rats and Pouched Mice Long-tailed Pouched Rats Giant Pouched Rats Pouched Mice Swamp Mouse

p. 153 p. 154 p. 157 p. 161 p. 165

Delany’s Swamp Mouse Climbing Mice, Largeeared Mouse, Fat Mice African Climbing Mice Velvet Climbing Mice Long-eared Mouse Bale Mouse Climbing Mouse Fat Mice White-tailed Rat

p. 166 p. 168 p. 169 p. 184 p. 186 p. 188 p. 189 p. 191 p. 201

White-tailed Rat Rock Mice

p. 201 p. 203

Rock Mice

p. 204

The family Nesomyidae encompasses several small groups of archaic muroid rodents whose living members are confined to subSaharan Africa (Cricetomyinae, Delanymyinae, Dendromurinae, Mystromyinae, Petromyscinae) and to Madagascar (Nesomyinae). In Africa, the family is represented by five subfamilies, 12 genera and 34 species. Each of the subfamilies is morphologically well characterized, but the family itself lacks clear diagnostic features in view of the immense heterogeneity embraced. Collectively, the five African subfamilies are highly diverse in size and morphology, habits, trophic niche and ecology (see subfamily and genus profiles). Tullberg (1899) and later Chaline et al. (1977) recognized the Nesomyidae, but the content of the family was largely restricted to the indigenous Malagasy rodents as previously identified by Major (1897) at the rank of subfamily. Although differing in contents, the family composition observed here owes its conceptual roots to Lavocat (1973, 1978), who identified a number of small but morphologically well defined groups as relicts of a middle Tertiary (late Oligocene–Miocene) cricetodontine presence in Africa and

broadened the definition of Nesomyidae to embrace their diverse descendants (also see Carleton & Musser 1984). Prior to Lavocat’s contributions, these archaic African muroids had been variously and inconsistently divided between Cricetidae and Muridae, or all were placed in an inclusive Muridae (see Carleton & Musser 1984, for classificatory review). The family’s expanded composition was initially based on tenuous dental links to middle Tertiary fossils, but results of molecular phylogenetic studies, although not wholly concordant, supply additional empirical support for Lavocat’s view of Nesomyidae. These gene-sequence investigations associate Cricetomyinae, Dendromurinae, Mystromyinae and Nesomyinae as a monophyletic lineage (Nesomyidae) basal to other muroid taxa that represent Cricetidae and Muridae (DuBois et al. 1996, Jansa et al. 1999, Michaux & Catzeflis 2000, Michaux et al. 2001). Tong & Jaeger (1993), on the other hand, considered Lavocat’s family to be a polyphyletic wastebasket that encompasses the remnants of early evolutionary branches leading to the major radiations of Cricetidae or Muridae.The evidence needed to settle these issues of relationship and classification will require greater emphasis on molecular and genetic characters and more extensive studies on morphology (rather than on dentition). The antiquity of the five African subfamilies is substantiated by palaeontological information. Evolutionary origin of each has been linked, with varying degrees of confidence, to early Miocene to early Pliocene fossil genera, most of these known from sub-Saharan sites (see subfamily accounts). Only the Dendromurinae is firmly documented outside of sub-Saharan Africa in the middle Tertiary (e.g. Aguilar et al. 1984, De Bruijn 1999). The palaeontological argument for such phyletic connections remains sketchy and the hard evidence from critical middle Tertiary beds is scanty. Further discoveries from the Tertiary of Africa will help to clarify the validity of the family. Compared with the African radiations of Gerbillinae and Murinae (Muridae), each of the subfamilies of Nesomyidae contains few genera and species: Cricetomyinae (3 genera, 5 species); Delanymyinae (1 genus, 1 species); Dendromurinae (6 genera, 23 species); Mystromyinae (1 genus, 1 species); and Petromyscinae (1 genus, 4 species). Michael D. Carleton & Guy G. Musser

Subfamily CRICETOMYINAE – Pouched Rats and Pouched Mice Cricetomyinae Roberts, 1951. Mammals South Africa, p. 434.

The three genera and five species comprising this subfamily are endemic to Africa, occurring in sub-Saharan savannas and in lowland and montane rainforest. Only one genus is found in West Africa (Cricetomys), but all three genera are represented in eastern and southern Africa (Beamys, Cricetomys and Saccostomus). Species in the

three genera are terrestrial and nocturnal, or predominantly so, consume seeds, fruits and bulbs, build relatively complex burrow systems, and hoard foods within those burrows for immediate or later consumption. The last habit is due, in part, to the possession of internal cheek pouches (a cardinal morphological trait of the 153

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subfamily), which are used to carry food from the foraging area to the burrow. Capacious cheek pouches have also evolved within other lineages of Rodentia, namely Sciuridae (squirrels) and Geomyoidea (pocket gophers, kangaroo mice and rats), but in muroid rodents, such pouches characterize only this subfamily and the Palaearctic hamsters (Cricetidae: Cricetini). Details of buccal histology, myology and innervation indicate that pouches were independently acquired by these muroid subfamilies (Ryan 1989). The infrequency of elastin fibres and absence of distensible folds in the pouch walls of cricetomyines suggest that they do not hoard foodstuffs or accumulate the vast underground larders to the extent documented for the cricetine hamsters. Species in the subfamily are small (Saccostomus) to very large (Cricetomys) in size.They have a robust body, large head, comparatively short and stout limbs and very short (Saccostomus) or very long (Beamys, Cricetomys) tail. The cheek pouch retractor is derived from facial muscle, innervated by cranial nerve VII and originating from the anterior thoracic vertebrae (Ryan 1989). Hindfeet strongly built, broad across the metatarsum, with short toes that have inconspicuous ungual tufts; plantar surface naked with six pads, the thenar and hypothenar positioned distally and close to four interdigitals. The skull characters include strong construction, rostrum moderately long and interorbital region hourglass-shaped with edges squared to slightly beaded; zygomatic plate with slight dorsal notch, jugal forming a prominent element of the middle zygomatic arch; alisphenoid strut present; subsquamosal fenestra absent, postglenoid foramen present; tegmen tympani not overlapping squamosal (Carleton & Musser 1984). Molars cuspidate, uppers with three roots and lowers two; upper molars with accessory lingual conules positioned to form transverse laminae suggestive of a rudimentary triserial arrangement, lowers with labial conulids; longitudinal enamel connections between lamina absent (Petter 1966a, c). Upper incisors without grooves, lowers with inconspicuous parallel enamel striae (Pocock 1987). The cephalic arterial circulation lacks a supraorbital branch of stapedial artery (sphenofrontal foramen and squamosal-alisphenoid groove are absent), but the infraorbital branch is present (stapedial foramen and parapterygoid groove are present). For so distinctive and closely related a group, African Pouched Rats and Mice were not formally acknowledged taxonomically until Roberts’s (1951) classification of South African mammals. In the early systematic literature, the three genera were classified within

Murinae (e.g. Thomas 1897, Ellerman 1941, Simpson 1945), but subsequent systematic arrangements have followed Petter (1966a, c) in allying cricetomyines with cricetids (e.g. Misonne 1974, Rosevear 1969, Skinner & Smithers 1990). Molar occlusal configuration (Petter 1966a) and anatomy of internal cheek pouches (Ryan 1989) convincingly sustain the monophyly of the subfamily. Mitochondrial and nuclear DNA sequence data (Jansa et al. 1999, Michaux & Catzeflis 2000, Michaux et al. 2001) also support monophyly and indicate that Cricetomyinae is phylogenetically close to Dendromurinae and Mystromyinae, two other endemic African subfamilies. Indisputable fossil representatives are known from the late Miocene to Recent of eastern and southern Africa (Denys 1988, Senut et al. 1992, Avery 1995, 1996, Mein et al. 2004), and the autochthonous African origin of Cricetomyinae has been speculatively linked to Miocene Afrocricetodontinae, a phyletic connection that so far lacks persuasive demonstration (Chaline et al. 1977, Tong & Jaeger 1993). Roberts (1951) segregated Saccostomus, as a lone member of Saccostomurinae, from other African pouched rats (Cricetomyinae), a division not recognized by later systematists (Petter 1966a, Ryan 1989); however, morphological traits and gene-sequence data link Beamys and Cricetomys as cognate relatives separate from Saccostomus (Carleton & Musser 1984, Corti et al. 2004). Two tribes within the subfamily may be recognized: (1) Cricetomyini (Beamys, Cricetomys): tail longer than combined head and body; anterior palatal foramina short; bony palate relatively short, lacking posterolateral palatal pits; mesopterygoid fossa moderately long; alisphenoid bone possessing dorsal orbital flange; accessory foramen ovale present; ectotympanic bullae (part of auditory bullae) small; vertebral column with 13 thoracic and six lumbar vertebrae; entepicondylar foramen of humerus present; corpus of stomach densely papillated; (2) Saccostomurini (Saccostomus): tail conspicuously shorter than head and body; anterior palatal foramina long; bony palate long with prominent posterolateral palatal pits; mesopterygoid fossa short and wide; alisphenoid lacking dorsal flange; accessory foramen ovale absent; ectotympanic bullae moderately inflated; vertebral column with 12 thoracic and seven lumbar vertebrae; entepicondylar foramen absent; corpus smooth, lacking papillae. The three genera and five species are listed alphabetically below. Michael D. Carleton & Guy G. Musser

GENUS Beamys Long-tailed Pouched Rat Beamys Thomas, 1909. Ann. Mag. Nat. Hist., ser. 8, 4: 107. Type species: Beamys hindei Thomas, 1909.

The genus Beamys contains of one (or two) species confined to evergreen forests of eastern Africa. Beamys is probably close to the ancestral stock of the Cricetomyinae. Two of its features, its habitat and the presence of an ectoparasite Hemimerus, which also occurs on Cricetomys, are considered to be primitive characters. Many of the characters of Beamys such as cheek pouches, body shape, shape and colouration of tail, reduction of M3 and only two cusps on the lamina of M1 are similar to those of Cricetomys (Hanney & Morris 1962,

Petter 1966c) (Figure 25). Musser & Carleton (2005) regard the two taxa, hindei and major, as separate species. An alternative arrangement is that there is a single species that shows a geographic cline in size – the smaller hindei in the northern part of range and the larger major in the southern part of range. Here, the genus is considered to have single species Beamys hindei. See also profile Beamys hindei below. D. C. D. Happold

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Beamys hindei LONG-TAILED POUCHED RAT Fr. Petit rat à abajoues; Ger. Langschwanz-Hamsterratte Beamys hindei Thomas, 1909. Ann. Mag. Nat. Hist., ser. 8, 4: 108. Taveta, Coastal Province, Kenya.

Beamys hindei.

Taxonomy Beamys hindei was described in 1909 from an immature individual with a small hindfoot collected in S Kenya. Subsequently the species was shown to occur throughout E Tanzania. A second species (B. major), with a larger hindfoot, was described in 1914 from Malawi and is now known to occur also in E Zambia. Some authors (e.g. Misonne 1974, Musser & Carleton 1993, 2005) retain these two species, but others (e.g. Ansell & Ansell 1973, Fitzgibbon et al. 1995) consider major as a subspecies of hindei. Fitzgibbon et al. (1995) show that there is a trend for individuals from southerly latitudes to be slightly larger than those from northerly latitudes. Specimens of major from Malawi are only slightly larger than those from S Tanzania, as would be expected from the general trend of increasing size from north to south. Here, all populations from Kenya to Malawi are considered as belonging to a single species, B. hindei, which shows clinal variation in size; northern populations are referred to as B. h. hindei and southern populations as B. h. major. Synonyms: major. Subspecies: two. See also profile genus Beamys. Chromosome number: 2n = 52 (Fitzgibbon et al. 1995). Description Medium-sized rat, soft grey dorsally, white ventrally, and with thick whitish blotched tail. Dorsal pelage warm grey, sometimes with a russet tinge on rump and back; dorsal hairs medium grey with warm grey tips. Ventral pelage, chin, throat pure white. Pelage dense and soft. Face pointed, ears small and rounded; vibrissae long and black; eyes black, small. Some individuals have white blaze on forehead. Limbs short, fore- and hindfeet white. Forefeet with four digits; hindfeet five digits. Tail long (ca. 100% of HB), scaly, thick (especially at base), whitish often with irregular dark blotches. Dorsal pelage of juveniles pale grey. For most measurements, "" are larger on average than !!. Geographic Variation B. h. hindei: northern part of range. On average, smaller in size. B. h. major: southern part of range. On average, larger in size.

Figure 25. Skull and mandible of Beamys hindei (HC 2787).

Similar Species Cricetomys gambianus. Much larger and heavier (mean HB: 326 mm); relatively long tail (ca. 85–130% [mean 107%] of HB); terrestrial, wider distribution. Saccostomus campestris. Smaller (mean HB: ca. 114 mm), broader head; relatively short tail (ca. 44–50% of HB); terrestrial, wider distribution, not confined to forested habitats. Distribution Endemic to Africa. Zambezian Woodland BZ and Coastal Forest Mosaic BZ. Recorded from coastal forests within about 100 km of coast from S Kenya to S Tanzania (B. h. hindei), and extreme SW Tanzania, Malawi (and perhaps parts of N Mozambique) and NE Zambia (B. h. major). Investigations in suitable habitats between the ranges of the two subspecies have failed to find any individuals (Fitzgibbon et al. 1995). Habitat Evergreen or slightly deciduous forests preferably on sandy soils, and gallery forests associated with streams. At ArabukoSokoke, SE Kenya, these Pouched Rats also occur in Afzelia forest and, to a lesser extent, Brachystegia forest. Occasionally found on fallow land and cassava plantations (Kenya only). Abundance Sparsely distributed; recorded from only a few localities but commoner than previously realized. In most localities, only one or two individuals have been found (perhaps because of trap-shyness). In a few optimal habitats, populations may be high, e.g. at Arabuko-Sokoke, where trees were close together and there 155

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Beamys hindei

was dense vegetation below 4 m, density was 14–30 individuals/ha. Comparable abundance is known in at least one mixed dry forest in the northern East Usambara lowlands, but the species was surprisingly scarce 700 m higher in sub-montane habitats (N. Cordiero unpubl.) Adaptations Nocturnal, scansorial. Although mainly terrestrial, can clamber about on twigs and small branches. When climbing, the tip of the prehensile tail is twisted around a twig, or it is held straight backwards as a counterbalance. During the day, individuals rest in a burrow. Most burrows have a straight vertical shaft with a nest chamber and latrine chamber. The nest is lined with fresh green leaves, which are changed regularly. When foraging, seeds and other foods are collected and stored in the cheek pouches. The food is taken to the nest and disgorged. Pouched Rats store considerable quantities of food, which is eaten when conditions for foraging are poor (see below). Pouched Rats are slow-moving and lethargic, spending a lot of time curled up in their nests. During cool weather, they may become dormant. Foraging and Food Forage on the ground and in trees. Omnivorous, feeding primarily on seeds and fruits, and occasionally insects. Food is stored in the nest. One excavated nest (B. h. major) contained nearly 1400 seeds of several species (weight ca. 1200 g), which would have taken about 200 separate forays to collect (Hanney & Morris 1962). Social and Reproductive Behaviour Usually solitary. In captivity, !! not in breeding condition attack "", biting them on hips, tail and scrotum. Receptive !! have been observed to perform a ‘forwards and backwards’ dance in a circular area in front of ", as well as biting him, prior to copulation (Hubbard 1970a).

Reproduction and Population Structure Times of reproduction depend on the locality. At Arabuko-Sokoke, reproduction occurs in most months of the year (Fitzgibbon et al. 1995). Lactating !! were present for nine months of the year (but not in Nov, Mar and Aug, the months of lowest rainfall); "" had scrotal testes in all months. However, only a proportion of all individuals were reproductively active in any month: 20–60% of !! were lactating during the nine months, and 50–90% of the "" were scrotal (except Sep – 25%). The highest percentage of breeding !! was in May at the height of the wet season. Females bred at least once per year. In Malawi, reproduction is seasonally polyoestrous; pregnant !! have been recorded Nov–May during the wet season and the beginning of the dry season (Hanney & Morris 1962); during the (cold and hot) dry season (Jun–Nov), testes of "" were abdominal. Studies on captive animals (B. h. hindei) have provided detailed information on reproduction (Egoscue 1972): age at first conception: ca. 7–9 months. Gestation 22–23 days. Eyes of young open Day 21. Weaning Day 35–40. Minimum interval between births 62 days; no postpartum oestrus. Litter-size: 2.8 (1–5, mode 3, n = 39). In Malawi, litter-size: 4.6 ([n = 4] and 7 [n = 1]). Young born with pink-coloured skin and a fine down of grey hairs; wt 3.2 (2.1–4.3) g. Growth is rapid; at four weeks young are still suckling and weigh ca. 43 g (Hanney & Morris 1962). Longevity in captivity: 3–4 years. At Arabuko-Sokoke, where breeding occurred throughout most of the year, there was a low but continuous recruitment of young individuals into the population. Numbers remained fairly constant throughout the year (14–30/ha) but with a peak in May and Jun due to recruitment of many young. There appears to be no seasonal change in weight. Predators, Parasites and Diseases Remains of Pouched Rats have been found in owl pellets at Lunzu, Malawi (Hanney 1962), and one individual was found in the stomach of a Twig Snake (Thelatornis capensis) at Namakutwa, Tanzania (Fitzgibbon et al. 1995). In Malawi, nests of Pouched Rats may be infested with ectoparasitic earwigs (Araeomerus morrisi, formerly Hemimerus morrisi [Nakata & Maa 1974]) where it is assumed they feed on stored fruits, and less frequently on the skin of Pouched Rats (Popham 1984). One species of flea, Xenopsylla microphthalma, has been recorded (Beaucornu & Kock 1996). (See also Genus Cricetomys.) Conservation IUCN Category: Least Concern. Populations are small and widely dispersed, and their forest habitats are being modified or reduced in area by human activities. (IUCN previously recognized hindei and major as full species; both were considered as Near Threatened.) Populations numbers are considered to be declining. Measurements Beamys hindei HB (""): 146 (135–158) mm, n = 5 HB (!!): 140 (115–164) mm, n = 10 T (""): 135 (127–144) mm, n = 5 T (!!): 127 (112–145) mm, n = 10 HF (""): 22 (21–24) mm, n = 5 T (!!): 21 (18–24) mm, n = 10 E (""): 20 (18–22) mm, n = 5 E (!!): 20 (17–22) mm, n = 10

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M1–M3 (!!): 5.1 (4.7–5.2) mm, n = 10 Mnina, Tanzania (RMCA)

WT (""): 69 (47–96) g, n = 5 WT (!!): 67 (49–94) g, n = 10 GLS (""): 37.5 (35.7–39.1) mm, n = 5 GLS (!!): 36.0 (32.6–39.0) mm, n = 10 GWS (""): 17.4 (16.0–18.7) mm, n = 5 GWS (!!): 16.5 (14.7–18.2) mm, n = 10 M1–M3 (""): 5.3 (5.2–5.4) mm, n = 5

Key References & Morris 1962.

Egoscue 1972; Fitzgibbon et al. 1995; Hanney D. C. D. Happold

GENUS Cricetomys Giant Pouched Rats Cricetomys Waterhouse, 1840. Proc. Zool. Soc. Lond. 1840: 2. Type species: Cricetomys gambianus Waterhouse, 1840.

Cricetomys gambianus.

The genus contains two to four species, and is distributed extensively throughout sub-Saharan Africa to about 27° S, including Zanzibar and Bioko Islands. The genus is represented in nearly all habitats from dry savanna to rainforest, and is often associated with human settlements. Giant Pouched Rats are distinguished from most other African murids by their very large size, unscaled and bi-coloured tails (basal half dark, terminal half white), and ungrooved incisors (Figure 26). They have internally opening cheek pouches, a character shared with the other two genera in the subfamily Cricetomyinae (Beamys and Saccostomus). The number of species in the genus is uncertain. Allen (1939) recognized six species, and Ellerman (1941) reduced these to subspecies of a single species, C. gambianus. Genest-Villard’s (1967) revision provided evidence of two ‘species’, a predominantly savannadwelling species (C. gambianus) and a rainforest species (C. emini) (see Musser & Carleton 1993). These two species are recognized here. Musser & Carleton (2005) accept these two species for West Africa, but claim that Genest-Villard’s character states and univariate analysis do not discriminate the southern African forms of the genus. These authors name the southern African forms of the genus as C. ansorgei and C. kivuensis. Cricetomys ansorgei (given as synonym of C. gambianus by Musser & Carleton 1993) is the savanna-living form, occurring in East Africa (including Zanzibar), westwards to Angola and southwards to South Africa. Cricetomys kivuensis (given as

Figure 26. Skull and mandible of Cricetomys gambianus (HC no number).

synonym of C. emini by Musser & Carleton 1993) is a forest-living form, known from montane habitats of E DR Congo, S Uganda, Rwanda and Burundi. The two species recognized here are: Cricetomys emini, which has soft, sleek dark brown pelage, a pointed face and is restricted to the Rainforest BZ; and Cricetomys gambianus, which has rather coarse greyish-brown pelage, a wide blunt face, and is restricted to savanna habitats and ‘savanna-like’ habitats around the margins of the Rainforest BZ. Justina C. Ray & J.-M. Duplantier

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Cricetomys emini EMIN’S GIANT POUCHED RAT (FOREST GIANT POUCHED RAT) Fr. Rat Géant d’Emin; Ger. Emins Riesenhamsterratte Cricetomys emini Wroughton, 1910. Ann. Mag. Nat. Hist., ser. 8, 5: 269. Monbuttu, Gadda, DR Congo.

Taxonomy Synonyms: dissimilis, dolichops, kivuensis, liberiae, luteus, poensis, proparator, sanctus, tephrus. The form kivuensis is regarded as a valid species by Musser & Carleton (2005; see profile Genus Cricetomys). Grubb (2004) suggests that the correct specific name for this species may be C. dissimilis de Rochebrune, 1885 (see also Hatt 1940). Subspecies: none. Chromosome number: 2n = 80, aFN = 80 (Codjia et al. 1994). Description Largest of the African forest murid rodents; similar to C. gambianus but with slender body form. Pelage short, sleek and soft in texture. Dorsal pelage orange-brown to dark greyish-brown or brownish-black. Head and flanks generally paler in colour. Ventral pelage white; usually well delineated from colour of flanks. Guard hairs few in number but very long. Face narrow in appearance with large, pale, naked ears standing well above pelage. Whiskers numerous and long. Dark eye-ring absent. Limbs long and thin. Feet whitish, long and powerful, but with relatively short claws. Digit 5 of the forefoot rudimentary; Digit 5 of hindfoot reaches to about half the length of Digit 4.Tail very long (ca.115% of HB), smooth without scales, dark on basal half, white on terminal half. Nipples: 2 + 2 = 8. Geographic Variation None recorded. Similar Species C. gambianus. More thick-set in body form, pelage rough, shaggy and longer; ventral pelage off-white, not clearly delineated from flanks; face broader with dark eye-ring; ear smaller, partly submerged in pelage; occurs in grasslands, woodlands and anthropogenic habitats in savannas, but sympatric with C. emini in some localities along edge of Rainforest BZ. Distribution Endemic to Africa. Widespread in Rainforest BZ and Rainforest–Savanna Mosaics. Recorded from Sierra Leone to S Uganda, Rwanda, Burundi, DR Congo, NE Angola, Equatorial Guinea, Gabon and Congo, Bioko I. (Fa 2000). Habitat Prefers ‘high forest’ as opposed to open savanna or commensal habitats (Rosevear 1969; Genest-Villard 1967). Within rainforest, demonstrates no marked preference for habitat type (Ray 1996). Also occurs in secondary forest along logging roads (Ray 1996, Malcolm & Ray 2000). In Dzanga-Sangha, Central African Republic, Forest Giant Rats were captured most often where the understorey was relatively open (Ray 1996). Abundance Mean population density at three sites in central Africa is 134.0 ± 16.9/km2 (Fa & Purvis 1997). Probably much more abundant than assessed by trapping (Dosso 1975b). Although only ten individuals were captured during 23,291 trap-nights in Taï Forest, Côte d’Ivoire, these Giant Rats were the species most often purchased from villagers or collected during night surveys on forest trails (Dosso 1975b).

Cricetomys emini

Adaptations Generally terrestrial, but able to climb and jump (Kingdon 1974). Captured frequently at height of 2 m in DzangaSangha, Central African Republic, but significantly more abundant on the ground (Malcolm & Ray 2000). Nocturnal (Rahm 1967), but occasionally seen in daylight (Rosevear 1969). Shelters in underground burrows, either modified or self-constructed, among large tree roots, and in holes in rotten logs and fallen tree trunks (J. C. Ray unpubl., Sanderson 1940). Burrows are characterized by a ‘complex system’ of galleries with side branches and separate chambers for food storage, sleeping, or waste; there are several escape routes. Nests are lined with fresh leaves. Individuals constantly change burrows (Rosevear 1969). Well worn pathways are used regularly for travelling within the home-range (J. C. Ray unpubl., Sanderson 1940). Foraging and Food Mainly vegetarian. Consumes wild and cultivated fruits (Rahm & Christiaensen 1963, Rosevear 1969). Snails also an important food-source (Rahm & Christiaensen 1963). Termites discovered in one stomach (Hatt 1940a). Detailed analysis of diet not available. Coprophagous. Social and Reproductive Behaviour Solitary except when raising young. One individual to a burrow (Rosevear 1969). Produces call that is modulatory in nature, the function of which is unknown (Genest-Villard 1967). Reproduction and Population Structure Lives up to 4.5 years in captivity. Gestation 42 days. Litter-size: 2–4 (Rosevear 1969).

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Predators, Parasites and Diseases Humans are the most important predators. These Giant Rats are particularly common in local food markets throughout central and West Africa in those areas where ungulates and primates are rare (Fa 2000). Ranked the second commonest species at bushmeat markets on Bioko I., and eighth commonest species at markets in Rio Muni (Equatorial Guinea). Daily availability ranged from 0.43 carcasses (Rio Muni) to 5.3 (Bioko I.; Juste et al. 1995). Most common rodent in markets in Kisangani, DR Congo, representing 90% of 35,992 rodent carcasses (Colyn et al. 1987). Remains are commonly found in the scats of mammalian carnivores, such as mongooses Herpestes naso and Bdeogale nigripes, genets Genetta servalina, African Civets Civettictis civetta and Golden Cats Profelis aurata, at a frequency of occurrence ranging from 3.3 to 11.8% (Ray & Sunquist 2001). Found in 4.9% (n = 150) of scats of Leopards Panthera pardus from Dzanga-Sangha, but in only 1.9% (n = 215) of scats from Taï Forest, Côte d’Ivoire (Hoppe-Dominik 1984). Absent in leopard scats (n = 222) from Ituri Forest, DR Congo, where leopards prey on larger species (Hart et al. 1996). As for C. gambianus, ectoparasites include Hemimerus spp. The nematode Capillaria heopatica parasitizes the liver, and is known to have zoonotic potential, causing human hepatic capillariasis. A relatively high prevalence (27%) of this nematode has been recorded in wild-caught C. emini in DR Congo; because Giant Rats are frequently consumed by humans, this level of parasitism has important implications for public health (Malekani 1994).

where there has been deforestation. A well-known pest in cocoa farms where these rats climb the trunks of cocoa trees to feed on cocoa pods. In Sierra Leone, C. emini may be displaced by C. gambianus following removal of rainforest habitats (Grubb et al. 1998). Measurements Cricetomys emini HB (""): 336.4 (274–379) mm, n = 14 HB (!!): 328.2 (276–378) mm, n = 34 T (""): 392.2 (339–426) mm, n = 13 T (!!): 386.5 (332–435) mm, n = 34 HF (""): 66.8 (62–72) mm, n = 14 HF (!!): 65.8 (60–69) mm, n = 33 E (""): 44.4 (39–51) mm, n = 14 E (!!): 45.1 (40–50) mm, n = 33 WT (""): 935.5 (455–1300) g, n = 22 WT (!!): 902.7 (514–1200) g, n = 39 GLS (""): 71.2 (62.6–74.8) mm, n = 10 GLS (!!): 71.9 (67–76) mm, n = 10 GWS (""): 30.8 (28–32.7) mm, n = 9 GWS (!!): 31.4 (28.5–33) mm, n = 10 M1–M3 (""): 10.3 (9.5–10.8) mm, n = 10 M1–M3 (!!): 10.6 (9.4–11) mm, n = 9 Dzanga-Sangha, Central African Republic (J .C. Ray & J. R. Malcolm unpubl.) Key References

Genest-Villard 1967; Rosevear 1969.

Conservation IUCN Category: Least Concern. Potential threats are overhunting close to human population centres where primates and ungulates have been depleted, and

Justina C. Ray

Cricetomys gambianus GAMBIAN GIANT POUCHED RAT Fr. Rat Géant de Gambie; Ger. Gambia-Riesenhamsterratte Cricetomys gambianus Waterhouse, 1840. Proc. Zool. Soc. Lond., 1840: 2. River Gambia, Gambia.

Taxonomy Considered to include C. emini until the revision of Genest-Villard (1967) who clearly separated the two species and distinguished seven subspecies (currently synonyms) of C. gambianus. Synonyms: adventor, ansorgei, buchanani, cosensi, cunctator, dichrurus, dissimilis, elgonis, enguvi, gambiensis (spelling lapsus), goliath, grahami, haageni, langi, microtis, oliviae, osgoodi, selindensis, servorum, vaughanjonesi, viator. The form ansorgei is considered as a valid species by Musser & Carleton (2005; see profile Genus Cricetomys. Subspecies: none. Chromosome number: 2n = 78 (unknown origin, Matthey 1954), 2n = 80, aFN = 82 (Senegal; Granjon et al. 1992) and 2n = 82, aFN = 88 (Benin; Codjia et al. 1994).

body size. Forefoot with a rudimentary thumb; hindfoot strong but with rather small claws. Tail long (85–130% [mean 107%] of HB), dark blackish-brown on proximal half, white on terminal half; smooth without scales and with short and sparse hairs on proximal end. Length of white tip varies regionally (31–68% [mean 50%] in Senegal [J.-M. Duplantier unpubl.]; 37–46% [mean 41%] for "", 31–45% [mean 38%] for !! on N Transvaal, South Africa [Smithers 1983]). Nipples: 2 + 2 = 8.

Description The largest murid species in Africa (together with C. emini). Dorsal pelage coarse, rough and shaggy, ranging in colour from grey (savannas of West and central Africa to Uganda) to brown in the eastern and southern parts of the range. Flanks paler. Ventral pelage white to off-white, not clearly delineated from colour of flanks. Face broad with elongated muzzle; very long vibrissae. Eyes relatively small with dark eye-ring. Ears relatively large, lower part usually submerged in pelage. Limbs relatively short compared to

C. gambianus: West and central Africa. Dorsal pelage predominantly grey; proximal part of tail very dark. C. ansorgei: south of the Rainforest BZ. Dorsal pelage predominantly brown, more yellowish than C. g. gambianus; longer body length than C. g. gambianus. (Musser & Carleton [2005] recognize this form as a valid species, and give the five forms listed below as synonyms of C. ansorgei.) C. microtis: Virunga Mts, DR Congo. The darkest and smallest form.

Geographic Variation Genest-Villard (1967) recorded seven subspecies (not recognized here):

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C. elgonis: Mt Elgon, Uganda/Kenya. Dark thick dorsal pelage and longer body length. C. kenyensis: Mt Kenya; skull narrow. C. enguvi:Taita hills and base of Mt Kilimanjaro (Kenya). Skull narrow and longer body length. C. cosensi: Zanzibar. Very large anterior palatal foramina; narrowest skull and the longest body length. Similar Species C. emini. Slender body form; pelage shorter and softer, ventral pelage white, clearly delineated from flanks; face narrow without dark eye-ring; ears larger and standing above pelage, skull more elongated (Genest-Villard 1967); restricted to rainforest habitats. Distribution Endemic to Africa.Widespread in Sudan and Guinea Savanna BZs, Zambezian Savanna BZ, Rainforest–Savanna Mosaics and marginally in southern part of Somalia–Masai BZ. Occasionally in Sahel Savanna BZ (see below). North of the Congo Basin recorded from Senegal and Guinea to Sudan, Uganda and Kenya; south of the Congo Basin recorded from Angola, S DR Congo, Zambia, Malawi, Tanzania, Mozambique and South Africa (KwaZulu–Natal and former Northern Transvaal Provinces) (Genest-Villard 1967, Smithers 1983). The distribution of C. g. ansorgei (see above) includes SE Kenya, Tanzania and all of the area from Angola to Mozambique and N South Africa. Zanzibar I. Habitat Widespread in savanna habitats. Also recorded in humanmodified habitats on edge of Rainforest BZ (Rosevear 1969, Happold 1987). In the Sahel Savanna BZ, found only in large cities. In the southeastern part of its range, restricted to evergreen forests and moister habitats (Morris 1963, Skinner & Smithers 1990). Often commensal. Abundance Generally considered as abundant throughout the range, but few quantitative data. Forty-five burrows found on a 5 ha

farmland in Nigeria (Ajayi 1975), 42 individuals taken from 0.5 ha garden in Zimbabwe (Smithers 1983). Adaptations Typically terrestrial, and predominantly nocturnal (Morris 1963, Ewer 1967). Giant Rats walk and run on all four legs, usually with the tail raised; they are good climbers and jumpers (Happold 1987). Nocturnal activity shows two peaks of activity, one at the beginning and one at the end of the night (Knight 1984). Burrows are often located in termite mounds or within the root system of large or dead trees (Morris 1963, Ajayi 1977), and range from 0.9 m to 2.9 m in length, with 50% less than 1.8 m (Ajayi 1977). A typical burrow has a large entrance leading to a nesting chamber composed of a nest, food store and sanitary area, and smaller additional burrows leading to small exit (escape) holes (Ajayi 1977). Burrows are excavated using the incisor teeth rather than claws (Morris 1963). May also live in rainwater drains and under houses. Tooth-gnashing, inflation of cheek pouches (thus making the head appear larger than normal) and a ‘puffing sound’ are produced when threatened. Vocalizations for intra-specific communication include ‘loud squeaks’, ‘high squeaks’ and ‘piping squeaks’ (Ewer 1967). Foraging and Food Omnivorous, but mainly vegetarian. The large cheek pouches are used to transport food (and nesting materials) to food stores in the nest (Ewer 1967). Food is frequently hoarded in nests: in Transvaal, South Africa, 8.7 kg of macadamia nuts were found in a single burrow (Knight 1984) and in Nigeria, 50% of stored food was palm fruits (Ajayi 1977). Diet of "" in S Nigeria (from stomach analysis, n = 5) was palm fruits (72%), seeds (12%), insects (7%) with some quantities of other fruits. Females consumed fewer palm fruits, but larger amounts of other fruits and vegetables. Diet varies according to location and food availability, and may include small quantities of animal matter (Iwuala et al. 1980). Most diets include adequate moisture but free water is drunk when necessary. Coprophagy appears to be a common habit (Ewer 1967). Social and Reproductive Behaviour Generally considered to be solitary in the wild, but Happold (1987) mentions the presence of several individuals in the same burrow. In captivity, "" and !! can be kept together easily (Ewer 1967). Litter-mates may show aggressive behaviour toward each others (Ewer 1967). Home-ranges are 2.2–11 ha (mean ca. 5 ha; Skinner & Smithers 1990); homeranges of "" larger than those of !!. Detailed descriptions of complex courtship and mating behaviour, as well as parental–young interactions, in captivity are given by Ewer (1967). Reproduction and Population Structure Pregnant !! and adults in breeding condition found all year round in Nigeria (Anizoba 1982). Young recorded between Sep and May in Malawi, suggesting that breeding takes place during the wet season (Morris 1963). Gestation: 27–42 days (Morris 1963, Ewer 1967, Ajayi 1975). Litter-size: 3 (1–5). Average weight at birth: 16–27 g, depending on litter-size; altricial; eyes closed, hairless. Silky covering of hair Day 10. First walking Day 16. First eating of solid food Day 18. Eyes open Day 20–24. Collects food in cheek pouches ca. Day 25, coprophagy Day 25 (Ewer 1967). Sexual maturity about 20 weeks for both sexes (Ajayi 1975).

Cricetomys gambianus

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Predators, Parasites and Diseases Predators include the Steppe Eagle Aquila rapax and eagle-owls Bubo capensis and Bubo lacteus (De Graaff 1981). Highly appreciated by humans as bushmeat. In W Nigeria, ranked as the 2nd (22%) or 4th (8%) commonest species for sale as bushmeat (Martin 1983, Anadu et al. 1988) after small antelopes, Cane Rats and Brush-tailed Porcupines (Martin 1983). Ranked 20th of preferred wild animals as a source of food in N Cameroon (Njiforti 1996). Eaten frequently in East Usambara Mts as well as in other coastal and montane forests in E Tanzania (N. Cordeiro unpubl.). Several species of Hemimerus ectoparasites (Insecta: Dermaptera; earwig family) are specific to Cricetomys gambianus, including H. talpoides, H. prolixus, H. deceptor, H. bouvieri (Ashford 1970, Popham 1984). Hemimerus is the only known parasitic member of this insect family (Happold 1987) and is found only on species of Cricetomys and Beamys. Other ectoparasites include a variety of ticks, mites and fleas; major intestinal parasites are tapeworms. Among human pathogens, Babesia and Bartonella bacteria have been identified in the blood (Dipeolu & Ajayi 1976); in West Africa, several viruses (including Bandia, Uganda S, Dugbe and Gabek-Forest) have been isolated from body organs of C. gambianus (Saluzzo et al. 1986).

Conservation IUCN Category: Least Concern. Common and widespread species, and not threatened. Bred in captivity as a supply of ‘bushmeat’ (Ajayi 1975). Measurements Cricetomys gambianus HB: 326 (273–407) mm, n = 66 T: 352 (277–423) mm, n = 58 HF: 62 (52–79) mm, n = 65 E: 37 (28–46) mm, n = 64 WT: 786 (500–1550) g, n = 65 GLS: 66.4 (58.8–70.5) mm, n = 34 GWS: 32.2 (28.7–34.5) mm, n = 35 M1–M3: 10.3 (9.3–11.0) mm, n = 34 Senegal (J.-M. Duplantier unpubl.) Key References De Graaff 1981; Ewer 1967; Genest-Villard 1967; Smithers 1983. J.-M. Duplantier & L. Granjon

GENUS Saccostomus Pouched Mice Saccostomus Peters, 1846. Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin, 11: 258. Type species: Saccostomus campestris Peters, 1846.

Saccostomus campestris.

The genus comprises two species widely distributed in savanna habitats in East and South Africa. Species in the genus are small-medium in size (and smaller than other members of the family), solidly built with broad heads, short limbs and short tails. Cheek pouches are well developed.The skull is characterized by long anterior palatal foramina and a long bony palate with prominent posterolateral palatal pits; the mesopterygoid fossa is short and wide, the alisphenoid lacks a dorsal flange, the accessory foramen ovale is absent, and the ectotympanic bullae are moderately inflated (Figure 27). These skull characteristics contrast with those of the other genera in the subfamily (Cricetomys, Beamys) and hence Saccostomus may be placed in a separate tribe, the Saccostomurini (see profile Subfamily Cricetomyinae). Species in the genus are nocturnal and terrestrial, and live in burrows during the day. They are primarily granivorous; individuals collect seeds in their cheek pouches and deposit them in caches in burrows for later consumption. When food is abundant, they

Figure 27. Skull and mandible of Saccostomus campestris (HC 2435).

accumulate fat and increase in weight; when food is scarce during cool and dry weather, they become inactive and torpid and survive, partly, on accumulated fat reserves. Litter-sizes are comparatively large, with up to ten young/litter. 161

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Two species are recognized. One species, S. campestris, has a large variation in chromosome number and may represent more than one species. The significance of such variation is uncertain. The second species, S. mearnsi from East Africa, was until recently considered as a subspecies of the more widespread S. campestris.

The species are distinguished by colour of body pelage, body size, number of chromosomes and geographic distribution. D. C. D. Happold

Saccostomus campestris CAPE POUCHED MOUSE (SOUTHERN AFRICAN POUCHED MOUSE) Fr. Rat à abajoues du Cap; Ger. Kap-Hamsterratte Saccostomus campestris Peters, 1846. Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin 11: 58. Tete, Zambezi River, Mozambique.

Taxonomy The species originally included mearnsi as a subspecies. The species, as currently defined, has considerable variation in chromosome numbers and in the structure of the chromosomes (Gordon & Rautenbach 1980, Gordon 1986). The variation is primarily geographic (see below), and more than one 2n number may occur in the same locality (Gordon 1986). There may be a complex of two or more species in southern Africa (Gordon 1986). Synonyms: anderssoni, angolae, elegans, fuscus, hildae, lapidarius, limpopoensis, mashonae, pagei, streeteri. Subspecies: none. Musser & Carleton (1993, 2005) suggest that anderssoni and mashonae deserve attention in respect of their relation to campestris. Chromosome number: 2n = 28 to 2n = 50, FN = 46–62 (details in Gordon 1986). Description Small, stocky mouse with soft, thick coat and short tail. Pelage fine and dense. Dorsal pelage pale brownish-grey to grey with some black-tipped hairs along mid-dorsal line; hairs dark grey at base, medium grey or brownish-grey at tip. Colour of dorsal pelage varies geographically. Ventral pelage pure white. Colour of dorsal pelage clearly delineated from colour of ventral pelage. Chin, throat, lower cheek and base of muzzle white. Head broad, with rounded nose. Well-developed cheek pouches extend to near shoulders; very conspicuous when full of seeds. Ears short and rounded, held sideways from head. Fore- and hindlimbs white, short and stocky; four digits on forefeet, five digits on hindfeet. Tail short (ca. 44% of HB), without scales, dark above, white below, with sparse short bristles. Young animals are darker than adult animals. Body measurements of "" slightly larger on average than for !!. Nipples: 3 + 2 = 10. Geographic Variation Sixteen chromosomal forms are recorded in southern Africa (Gordon 1986). There is a general decrease in 2n number from west (e.g. 2n = 46 at the coast of KwaZulu–Natal Province, South Africa) to east (e.g. 2n = 30, 31, 32 in W Namibia), but there are exceptions to this trend (e.g. 2n = 26–28 in northern populations such as SE Angola and N Zimbabwe). In southern Africa, specimens from the drier western part of the range are paler in colour than those from the wetter eastern areas. Three colour forms – pale buffy-brown, blackish-grey and medium dark grey – recorded in Malawi; these may be partly associated with age (Hanney 1965). Similar Species S. mearnsi. HB on average larger; tail relatively longer; dorsal pelage dark grey to brownish-grey; ventral pelage dark grey, hairs sometimes with white tip (never pure white); East Africa; probably allopatric to S. campestris in C Tanzania (see also below).

Saccostomus campestris

Distribution Endemic to Africa. Zambezian Woodland and South-West Arid (Kalahari) BZs. Recorded from Angola, Zambia, Malawi, S Tanzania, Mozambique (mainly south of the Zambezi R.; records north of the river are sparse), SE DR Congo, C and N Namibia, Botswana, Zimbabwe and South Africa (SE Transvaal, C and N Free State, S KwaZulu–Natal, SW and E Cape Province). Not recorded from Lesotho (Lynch 1994). The northern boundary of the range of S. campestris in C Tanzania, where it adjoins the southern boundary of S. mearnsi, is uncertain. Habitat Occurs in many types of woodland and grassland, and close to marshes. In Botswana, found in sandy regions, kopjes, open short grasslands, rocky kopjes, dry river beds, Acacia grasslands and mopane woodlands. In Malawi, found primarily in low altitude dry woodlands, but not on the high plateaux. In southern Africa, recorded in habitats where rainfall is 250 to over 1200 mm/annum, and from sea level to 1800 m. Sandy substrates with cover provided by bushes or open woodland are preferred habitat. In several parts of their range, burrows are dug in large abandoned termite mounds. Abundance Widely distributed and locally common. Numbers fluctuate seasonally and individuals are rarely encountered during cool

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dry weather. In South Africa, variations in density (0.31–1.70/ha) are related to the habitat, when the habitat was last burnt, and frequency of burning (Korn 1981). In Acacia tortilis savanna in north KwaZulu– Natal, densities varied from 0/ha to 3.8/ha, and in Acacia nigrescens savanna, from 0/ha to 1.8/ha (Swanepoel 1976). In thicket-clump savanna in Lengwe N. P., Malawi, mostly uncommon during the dry season (0–1/ha), but more abundant during and immediately after the wet season (up to 6/ha); during a ten-month period, they contributed 11% to the small mammal community, ranging from 0 to 1% in drier months and up to 28% in wetter months (Happold & Happold 1991). Adaptations Terrestrial, scansorial and nocturnal. Excavates burrows in sandy soils or in termite mounds, or utilizes burrows made by springhares or Aardvarks. Burrows vary greatly in complexity and usually contain stores of seeds. When food is abundant, individuals store fat and put on weight. During the cool dry weather (or ‘winter’ in southern Africa), stored fat is metabolized and individuals lose weight; at this time, overall energy demands are reduced (Korn 1989), body temperature declines to 21–25 °C for 2–6 hours/day, there may be short bouts of daily torpor (Ellison & Skinner 1992), and overall activity is reduced. The stomach comprises a non-glandular forestomach and a distinctly differentiated glandular hindstomach; however, the relatively low density of bacteria in the forestomach suggests that bacteria do not contribute greatly to digestion and metabolism (Perrin & Kokkinn 1986). Foraging and Food Predominantly granivorous. The diet includes a wide diversity of seeds. Seeds are gathered in the cheek pouches and taken to food caches in the burrow. The forefeet are used to help fill the cheek pouches. Seeds found in cheek pouches and in food caches include Grewia monticola, the umbrella thorn Acacia tortilis, sweet thorn A. karroo, scented thorn A. nilotica, camel thorn Acacia erioloba, nyala tree Anthocercis zambesiaca, red thorn Acacia gerrardi, Combretum spp., mopane Colophospermum mopane, sekelbos Dicrostachys cinerea, raisin bush Grewia bicolor, G. flavescens, Burkea africana, Euclea crispa and Peltophorum africanum (Smithers 1971, Swanepoel 1976, De Graaff 1981). Termites, grasshoppers and other insects are also eaten (De Graaff 1981). Kerley (1989) classified these Pouched Mice as ‘partially insectivorous granivores’ that forage widely from the burrows. The diet varies seasonally (Watson 1987): in Kruger National Park, during the dry season, it comprised 31% insects, 12% herbage and 57% seeds (n = 14), compared with 9% insects, 12% herbage and 79% seeds during the wet season. Social and Reproductive Behaviour Solitary. Home-range in Terminalia-Dichrostachys grasslands of Kruger N. P., South Africa, is 1200 m2 in control (unburnt) habitat and 1200–2800 m2 in habitat burnt every three years (Korn 1981). Females in oestrus or prooestrus are aggressive and attack "" (Swanepoel 1976).

Reproduction and Population Structure Breeding and recruitment varies according to locality: Oct–Feb in KwaZulu– Natal (Swanepoel 1972), Oct–Apr in Transvaal (Rautenbach 1982), Jan–Apr in Botswana (Smithers 1971) and Feb–Apr in Zimbabwe (Smithers & Wilson 1979). These months are mostly during the warmer wetter months of the year. In Malawi, pregnant !! found in Apr/May (warm late wet season), Aug/Sep (cool dry season) and Dec/Jan (warm early wet season) (Hanney 1965). Litter-size also varies geographically, e.g. litter-size: 4.8 (2–8) in South Africa (Earl 1978); 7 (5–10) in Botswana (Smithers 1971); 6.7 (1–10, n = 7) in Zimbabwe (Smithers & Wilson 1979); and 5.1 (2–9, n = 10) in Malawi (Hanney 1965). Gestation: 20–21 days (Earl 1978). At birth, young fully furred, weight 2.8 g.Weaned Day 19–25 when 11 = 15 g. First litter Day 96 (Earl 1978). Females are spontaneous ovulators with a four-day cycle, without a postpartum oestrus (Westlin-Van Aarde 1988), and they exhibit a lactational anoestrus (Westlin-Van Aarde 1989). The energy demands of !! increase by 55% during pregnancy and by 110% during lactation (Perrin & Clarke 1987). Predators, Parasites and Diseases Preyed on by Barn Owls Tyto alba, Grass Owls T. capensis and Giant Eagle-owls Bubo lacteus, and probably several small mammalian carnivores. They are hosts to a wide range of ectoparasites including 11 spp. of mites, 18 spp. of fleas and 4 spp. of ticks (details in De Graaff 1981). The nematode Inermicapsifer madagascariensis is a common endoparasite (details in De Graaff 1981). Conservation IUCN Category: Least Concern. Widely distributed and locally common, and not threatened in natural habitats. Measurements Saccostomus campestris HB: 114 (83–145) mm, n = 21 T: 50 (32–83) mm, n = 21 HF: 21 (17–30) mm, n = 21 E: 14 (12–22) mm, n = 21 WT: 48.5 (33–68) g, n = 20 GLS: 30.7 (29.0–33.5) mm, n = 17 GWS: 15.0 (13.9–16.0) mm, n = 17 M1–M3: 4.6 (4.0–4.9) mm, n = 17 Body measurements and weight: Botswana (Smithers 1971; "" only) Skull measurements: Zambia, Malawi (BMNH) Key References

De Graaff 1981; Hanney 1965; Smithers 1983. Mike Perrin

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Saccostomus mearnsi MEARNS’S POUCHED MOUSE (EAST AFRICAN POUCHED MOUSE) Fr. Rat à abajoues de Mearns; Ger. Mearns Hamsterratte Saccostomus mearnsi Heller, 1910. Smithson. Misc. Coll. 54: 3. Changamwe, Coast Province, Kenya.

Taxonomy Morphological and chromosomal evidence indicate that S. mearnsi is distinct from S. campestris (Hubert 1978a, Gordon 1986). Synonyms: cricetulus, isiolae, umbriventer. Subspecies: none. Chromosome number: 2n = 40–42 (Hubert 1978a), FN = 40–42. Description Medium-sized, stocky mouse with soft, thick coat and short tail. Dorsal pelage pale to dark grey, dark brown or brownish-grey; hairs medium grey at base, grey or brownish-grey at tip. Flanks slightly paler. Ventral pelage grey (cf. S. campestris); some hairs tipped with white to give frosted appearance. Chin and base of muzzle whitish-grey. Head broad, with rounded nose and large eyes. Well-developed cheek pouches. Ears large and rounded, held sideways from head. Fore- and hindlimbs dark grey, short and stocky; four digits on forefoot, five digits on hindfoot. Tail short (ca. 50% of HB), thick at base, without scales, well covered with grey to brownish-grey hairs above, white below. Males often heavier than !!. Nipples: probably 2 + 3 = 10, as for S. campestris. Geographic Variation None recorded. Similar Species S. campestris. Dorsal pelage medium grey; ventral pelage pure white; southern Africa as far north as C Tanzania. Distribution Endemic to Africa. Somalia–Masai Bushland BZ, and marginally around the Afromontane–Afroalpine BZ of Kenya. Recorded from SW Ethiopia, S Somalia, E Uganda, Kenya and N Tanzania. The southern boundary of the range is uncertain; current records suggest that S. mearnsi and S. campestris are allopatric inTanzania. Habitat Savanna; details of habitat preferences are poorly known. In C Kenya, abundant in woodland savanna on poorly drained soils and in shrubby thickets along seasonal watercourses and on kopjes. Abundance Comprised ca. 80% of a small mammal community in the Laikipia District of C Kenya (Keesing 1998a). Population numbers fluctuated from 45/ha following heavy rains to 6/ha after a prolonged drought, with a 5-year average of 20/ha. Mearns’s Pouched Mice were twice as abundant in areas from which large mammals had been excluded (Keesing 2000), presumably because of increased availability of food. In the few other areas from where they have been reported, S. mearnsi is less abundant than at Laikipia.

Saccostomus mearnsi

with smaller amounts of seeds (9%) and arthropods (7%); in the dry season, the diet contains more seeds (33%) and arthropods (22%) and lesser amounts of forbs and browse (43%) (Neal 1984a). Diet varies at different localities (Keesing 1998a, Metz & Keesing 2001). Pouched Mice clip and consume both forbs and tree seedlings and often leave small piles of harvested vegetation. In cafeteria trials and in the field, they show strong preferences for certain forbs (e.g. Commelina spp., Monsonia angustifolia) and for the seeds and seedlings of Acacia trees. Because of their consumption of seeds and seedlings, they may influence the recruitment of Acacia trees (Keesing 2000).They appear to compete with ungulates for some food resources, since the removal of ungulates results in rapid increases in their numbers (Keesing 1998b).

Adaptations Terrestrial and nocturnal, and strikingly slow-moving. Capable of digging deep burrows in hard soil with their strong legs and toes. Adults continue to grow after attaining reproductive maturity at ca. 45 g; old "" weigh as much as 120 g. See also S. campestris.

Social and Reproductive Behaviour Mostly solitary, living alone in burrows constructed in termite mounds, under shrubs and at the bases of trees. At high population densities, however, Pouched Mice live in close proximity, occasionally sharing burrow entrances and possibly entire burrow systems with conspecifics (Keesing 1998a). In encounters between pairs of same-sex individuals, both "" and !! exhibited a number of behaviours typical of more social rodents (e.g. vocalizations, allogrooming, appeasement). Based on data from long-term trapping records, !! appear to be territorial, while the home-ranges of "" are broadly overlapping and superimposed on those of !! (Keesing & Crawford 2001). Home-range of "": 0.21 ± 0.02 ha; home-range of !!: 0.06 ± 0.01 ha.

Foraging and Food Omnivorous. Diet varies seasonally; in the wet season, forbs and browse (82%) form the majority of the diet

Reproduction and Population Structure Reproduction is seasonal in C Kenya: almost all adult !! are in breeding condition

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in Aug–Nov following the wet season (Apr–Jul), while only 10% are in breeding condition during the dry season (Jan–Mar). At least 85% of adult "" remain in breeding condition throughout the year. Litter-size: 3–5. Gestation, ontogeny and time to maturity unknown (but see S. campestris). Predators, Parasites and Diseases Predators unknown. Reported to be hosts for juvenile stages of some Rhipicephalus ticks. However, larval ticks experimentally placed on mice were groomed off and never recovered, suggesting that these mice are poor hosts for juvenile ticks (F. Keesing unpubl.). Conservation

IUCN Category: Least Concern.

T: 69 (58–79) mm, n = 29 HF: 21 (20–22) mm, n = 31 E: 20 (19–23) mm, n = 29 WT (""): 79 (48–121) g, n = 121 WT (!!): 62 (39–83) g, n = 97 GLS (""): 34.8 (33.0–37.1) mm, n = 3 GLS (!!): 33.6 (30.4–36.3) mm, n = 5 GWS (""): 17.7 (16.9–19.1) mm, n = 4 GWS (!!): 17.2 (16.0–17.8) mm, n = 5 M1–M3: 5.9 (5.5–6.1) mm, n = 8 Laikipia District, Kenya (F. Keesing unpubl.) Key References Keesing 2001.

Hubert 1978a; Keesing 1998a, b; Metz &

Measurements Saccostomus mearnsi HB: 137 (115–160) mm, n = 30

F. Keesing

Subfamily DELANYMYINAE – Delany’s Swamp Mouse Delanymyinae Musser & Carleton, 2005. In Mammal Species of the World, 3rd Ed., p. 934.

The subfamily Delanymyinae contains only a single genus and species, Delanymys brooksi, endemic to Western Rift Mts in East Africa. This very small, gracile mouse possesses an exceptionally long and semi-prehensile tail and large hindfeet with slender toes. Such traits impressed Hayman (1962a), who described the species as convergently resembling Eurasian birch mice (Sicista, Dipodoidea); similar convergence in body form is also seen in the Sundaic and Sulawesian murine Haeromys (Musser 1990). Like these mice, Delanymys is an expert climber, adept at negotiating slim stems and branches and filling a semi-arboreal, granivorous niche. Until recently, Delanymys had been grouped with Petromyscus in Petromyscinae, but the two genera differ in many essential features. In Delanymys, the tail is twice as long as the head and body and semiprehensile (less than to moderately longer in Petromyscus), and the hindfoot is long and narrow (short and broad in Petromyscus). The cranium of Delanymys possesses a very short rostrum (long and slender in Petromyscus), narrow interorbital constriction (broad in Petromyscus), narrow zygomatic plate with a shallow dorsal notch (plate broad and notch deep in Petromyscus), and large subsquamosal foramen (closed in adult Petromyscus). Examples of Delanymys have a short bony palate, in contrast to the long palate that projects as a prominent shelf behind the third molars in Petromyscus. An alisphenoid strut is absent in Delanymys (present in Petromyscus), and the carotid circulation is fully derived, lacking supraorbital and infraorbital arteries and the accompanying squamosal-alisphenoid groove and sphenofrontal and stapedial foramina (partially derived in Petromyscus, which retains the infraorbital artery and a large stapedial foramen). The molars in both genera are brachydont and cuspidate, but those of Delanymys have anterolophs(ids) and mesolophs(ids) (all absent in Petromyscus), and its M3 is relatively large with an occlusal pattern that resembles M2 (M3 very small in Petromyscus, with C-shaped occlusal pattern unlike M2) (Figure 28). The unique morphology of Delanymys, especially characters of the molar dentition, has generated various interpretations of its phylogenetic placement. Hayman (1962) regarded Delanymys

as morphologically close to Petromyscus and considered both to be dendromurines, and Petter (1967) acknowledged this relationship by classifying Delanymys and Petromyscus in Petromyscinae, separate from Dendromurinae. Others have emphasized resemblances between Delanymys and Mystromys or have viewed Delanymys as a structural link between Mystromys and typical Dendromurinae (Lavocat 1964, Verheyen 1965a). These conflicting views hinge, in part, on disputes over the origin and homology of the lingual cusp on the upper molars of Delanymys and Petromyscus, believed to represent either the

Figure 28. Skull and mandible of Delanymys brooksi (skull - RMCA 96-038-M3386 with some detail from BMNH 61.1610; mandible after Verheyen (1965a); upper cheek teeth RMCA 96.038-M-3385).

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protocone (Petter 1967) or a neomorphic acquisition comparable to those of dendromurines and certain Miocene fossils (Lavocat 1964, Jaeger 1977b). Aside from the questionable phylogenetic significance of these lingual cusps, Delanymys and Petromyscus share no other derived features that persuasively justify their joint membership in Petromyscinae as a natural group (Carleton & Musser 1984, Denys 1994a). Further, the dual retention of mesolophs(ids) and longitudinal enamel connections on its molars marks Delanymys as unique among living African Muroidea and suggests that it

represents a relictual descendant from a very early radiation of the superfamily within the continent. Musser & Carleton (2005) erected the subfamily Delanyminae to contain the genus and urged further inquiry to illuminate its phylogenetic relationships. Fossils of the Delanymyinae, represented as the extinct Stenodontomys, are known from the early Pliocene to the early Pleistocene of southern Africa (Pocock 1987, Senut et al. 1992, Denys 1994). Guy G. Musser & Michael D. Carleton

GENUS Delanymys Delany’s Swamp Mouse Delanymys Hayman, 1962. Rev. Zool. Bot. Afr. 65: 1–2. Type species: Delanymys brooksi Hayman, 1962.

Montotypic genus. In the description of the holotype, Hayman (1962) placed Delanymys in the subfamily Dendromurinae. An alternative view based on the ‘longitudinal crest’ in the molar row, is that it is closely related to Mystromys (subfamily Mystromyinae) (Lavocat 1964). Petter (1967) interpreted the molar structure of Delanymys as very similar to that of Petromyscus (both have an additional lingual cusp on M1 and M2, which is connected to other cusps by a longitudinal ridge); he placed both genera in the subfamily Petromyscinae, even though they are very different in other skull characters, external morphology, habits and distribution. The current opinion seems to be that Delanymys is far removed from the Dendromurinae, but whether the similarities to Petromyscus are phylogenetic or due to convergence is uncertain. Further details are given in the subfamily and species profiles. Fritz Dieterlen

Delanymys brooksi.

Delanymys brooksi DELANY’S SWAMP MOUSE Fr. Souris palustre de Delany; Ger. Delanys Sumpfklettermaus Delanymys brooksi Hayman, 1962. Rev. Zool. Bot. Afr. 65: 1–2. Echuya (or Muchuya) Swamp, near Kanaba, Kigezi, SW Uganda.

Taxonomy See genus profile. Synonyms: none. Chromosome number: not known. Description Very small climbing mouse with an extremely long tail and long hindfeet. Pelage long (8–10 mm). Dorsal pelage warm russet or rufous to hazel-brown; hairs dark slate-grey on basal twothirds, russet or rufous on terminal one-third. Long black guard hairs (each with subterminal buff band) project well above pelage tending to give a generally darker colouration. Ventral pelage warm buff; hairs mostly ca. 7 mm, with tuft of long white hairs surrounding the urinogenital opening in both !! and "". Throat with longitudinal patch of pure white hairs. Eyes surrounded by short black hairs; black patch on nasal region between eyes and rhinarium. Lips with very long (20–25 mm) vibrissae. Ears relatively large, round, with welldeveloped ear folds, and long (5–7 mm) hairs in places. Upperparts of fore- and hindlimbs with dark hairs; inner surfaces with whitish hairs. Small tuft of white hairs on each wrist, thought to have a tactile function (Dieterlen 1969b). Fore- and hindfeet very small with long

digits and long curved claws (except for vestigial Digit 1 on forefoot, which has small nail); Digits 3 and 4 very long, Digits 2 and 5 long but shorter than 3 and 4. Tail extremely long (ca. 180% of HB), thin, almost hairless with small scales forming ring-like patterns. Males on average slightly smaller than "". Palatal ridges: three antemolar and four intermolar. Skull: rostrum very short, palate short, mesopterygoid fossa completely open anteriorly, zygomatic plate narrow; see also subfamily and genus profiles. Nipples: 2 + 2 = 8. Geographic Variation

None recorded.

Similar Species Dendromus spp. Black mid-dorsal stripe in some species; tail long but mostly relatively shorter; more widespread distribution for most spp. Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Recorded from a small area bordering the Albertine Rift Valley in

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tightly to prevent sliding down the plant. The hindfeet are unusually long (ca. 30% of HB) and have six plantar pads; Digits 2 to 5 are long, and Digit 1 is shorter but capable of being spread sideways. Digit 5 is especially long and strong, and is opposable. The long prehensile tail is used to provide balance when climbing, and the distal end can curl around stems to provide support. In these respects, the adaptations of the feet are very similar to those of Dendromus spp. The spreading ability of digits of the fore- and hindfeet is also very important when walking on the muddy ground. Foraging and Food Vegetarian. Stomach contents (n = 2) contained only the whitish pulp of farinaceous seeds, probably from grasses, without any green material or animal remains. Captive animals preferred seeds, especially sorghum; water was permanently available and used not only for drinking but also for regular defecation (Dieterlen 1969b). Social and Reproductive Behaviour Delany’s Swamp Mice are gentle mice, and individuals of both sexes live peacefully together in captivity. In this respect, they differ from Dendromus spp., which tend to show aggressive behaviour. Delanymys brooksi

SW Uganda, W Ruanda and E DR Congo (near L. Kivu). The most northern and eastern locality is Muchuya Swamp (01°15´–01°18´ S; 29°47´–29°51´ E), near Kanaba in Uganda where the type specimen was found in 1961 (Hayman 1962a). The southernmost record is Kitabi in Rwanda (02°34´ S, 29°26´ E; 2200 m). The geographic range may also include the area from the volcano region north of L. Kivu southwards to the Itombwe Mts north-west of L. Tanganyika. Habitat Most individuals have been found in high altitude marshes, rich in plant species, at altitudes of 1700–1760 m.The typical habitat is swamp where sedge (Cladium mariscus) is standing in water, and the vegetation in the shallows is elephant grass (Pennisetum), Hyparrhenia grass and non-grassy plants such as Abutilon, Rubus, Acanthus and Impatiens. The medium height of such vegetation is 1.5–2.0 m. A few individuals have been found in non-marshy habitats such as dense grassy vegetation (Mt Gahinga [2700 m] and Mt Karisoke [3100 m]), and between ferns in a plantation of Eucalyptus at Kitabi (2200 m) (Van der Straeten & Verheyen 1983). Abundance Using an enclosure and removal method, Delany’s Swamp Mice comprised 6.2% of all small mammals (n = 355; >20 species) (Dieterlen 1967a, b). In non-marshy habitats, they are extremely rare (Dieterlen 1967a; Van der Straeten & Verheyen 1983) and it can be assumed that even in marshes they are not numerous, except in a few locations. Adaptations Nocturnal and arboreal on grass stems. The foreand hindfeet are extremely well adapted for climbing on the stems of grass and other low herbaceous plants in marshes and grasslands. The forefoot has a rudimentary Digit 1 (thumb), four relatively long fingers (Digits 2, 3, 4 and 5), five large plantar pads and numerous small but prominent tubercles on palm and fingers. When grasping, the proximal pads and the thumb form fit tightly against the flexible fingers. The fingers can be spread widely for climbing, and closed

Reproduction and Population Structure Embryo number: 3 (n = 2 !!); one ! found in a nest had four young. At birth, young are altricial. At Day 10, the eyes are closed, the incisors have not yet erupted and the short hazel-coloured pelage is brighter than in adults, WT = 2.5–2.7 g. Individuals in captivity did not reproduce. Predators, Parasites and Diseases A dead individual found near the nest of an African Grass-owl Tyto capensis in E DR Congo suggests these owls may be potential predators (James Chapin, in Hayman 1963). Conservation IUCN Category: Vulnerable. Schlitter (1989) classified the species as Rare, and hence of some conservation concern. The small geographical range has experienced considerable habitat destruction in recent years and has a very high human population density. Measurements Delanymys brooksi HB: 56.8 (50–63) mm, n = 24 T: 104.1 (90–111) mm, n = 22 HF: 18.5 (17.0–20.5) mm, n = 25 E: 11.3 (10–13) mm, n = 17 WT: 5.8 (5.2–6.5) g, n = 8 GLS: 18.3 (17.3–19.0) mm, n = 12 GWS: 9.8 (9.5–10.1) mm, n = 12 M1–M3: 2.5 (2.3–2.8) mm, n = 15 Throughout geographic range (Dieterlen 1969b; Hayman 1962a, b; Van der Straeten & Verheyen 1983; Verheyen 1965a) Key References Verheyen 1965a.

Dieterlen 1969b; Hayman 1962a, 1963a; Fritz Dieterlen 167

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Subfamily DENDROMURINAE – African Climbing Mice Dendromurinae G. M. Allen, 1939. Bull. Mus. Comp. Zool., Harv. Coll. 83: 349.

The Dendromurinae encompasses a small muroid radiation (six genera and 24 species sensu Musser & Carleton 2005) whose living representatives are indigenous to sub-Saharan Africa. Although few in number, members of the six genera are highly diversified in their morphology, behaviour and ecology. Dendromus and Megadendromus, though occasionally active at ground level, are primarily adept climbers of slender grasses and shrubs where they forage and construct nests; accordingly, they are found in habitats where grasses and shrubby vegetation predominate (marshes, savannas, forest edges, alpine bamboo and heath zones). They are largely omnivorous, feeding upon seeds, berries and insects. In contrast, Steatomys is terrestrial, dwells primarily in savanna habitats, accumulates copious fat reserves during periods of abundant food and can aestivate in response to unfavourable environmental conditions. Malacothrix, endemic to the South-West Arid BZ, is terrestrial, granivorous and gerbil-like in certain aspects of its morphology and habits. Dendroprionomys and Prionomys are arboreal and insectivorous inhabitants of lowland evergreen rainforest. All are nocturnal. Diagnostic features that unite so heterogeneous a group are few: first and second upper molars consist of bicuspid laminae, with a lingual accessory cusp adjacent to the middle lamina of M1 and front lamina of M2. M3 and M3 are tiny and single-rooted. Each upper incisor has a single deep groove (except Prionomys). Other characters include: infraorbital foramen wide and ovoid; zygomatic plate narrow; dorsal notch indistinct or shallow; masseteric tubercle prominent; carotid circulatory pattern partially derived (sphenofrontal foramen and squamosal-alisphenoid groove absent, stapedial foramen spacious); postglenoid foramen large and subsquamosal fenestra present, middle lacerate foramen small; strut of alisphenoid bone present,

delineating an accessory foramen ovale; and tegmen tympani reduced, not contacting the squamosal (Petter 1966c, Dieterlen 1971, Carleton & Musser 1984) (Figure 29). Dendromurines are small to very small in size, and the genera vary substantially in external characters.The pelage is short, soft and slightly woolly (Dendromus, Megadendromus), short and velvety (Dendroprionomys, Prionomys), or dense and silky (Malacothrix); a black, mid-dorsal stripe is well developed in Megadendromus and some Dendromus. Compared with body size, the ears are small (Prionomys), moderately large (Dendromus, Megadendromus) or exceptionally large (Malacothrix). Relative to the head and body, the tail may be longer (Dendromus, Prionomys), about equal to (Megadendromus) or conspicuously shorter (Malacothrix, Steatomys); and its surface naked to thinly haired and visibly scaly (Dendromus, Megadendromus, Prionomys) or moderately to thickly haired with caudal scales inconspicuous (Malacothrix, Steatomys). The forefoot may possess only three functional digits (Digits 2, 3 and 4; Digits 1 and 5 present but small and non-functional; Dendromus, Megadendromus), four digits (Digits 2, 3, 4 and 5; Malacothrix, Steatomys), or four plus a stubby but definitive Digit 1 (Dendroprionomys, Prionomys). Malacothrix has four digits on the hindfoot, the other genera have five digits (Figure 30). In certain dendromurines, the claw is replaced by a nail on Digit 1 of the hindfoot (Dendromus, Dendroprionomys, Prionomys) and on Digit 5 (some Dendromus, Megadendromus). Palmar and plantar surfaces are naked (Dendromus, Megadendromus, Dendroprionomys, Prionomys), hairy over the proximate third (Steatomys), or densely furred so that the plantar pads are obscured (Malacothrix). The skull typically possesses a long and slender rostrum (most genera) or only moderately long and wide (Steatomys).The interorbital region is strongly constricted with smooth edges (Dendromus, Malacothrix, Megadendromus, Steatomys) or moderately wide with a weak postorbital ledge (Dendroprionomys, Prionomys). Anterior palatal foramina are slender and long, posterior margins reaching middle of first upper molars (Dendromus, Malacothrix, Megadendromus, Steatomys), or short and wide, the posterior margins set notably anterior to molar rows (Dendroprionomys, Prionomys). The auditory bulla is inflated in Malacothrix and Steatomys relative to other genera. Most genera have orthodont or opisthodont upper incisors, but in Prionomys the incisors

a

Figure 29. Skull and mandible of Dendromus melanotis (RMCA 38416).

b

c

Figure 30. Right hindfoot of (a) Dendromus sp., (b) Dendroprionomys rousseloti, (c) Prionomys batesi to show relative sizes of Digit 1 and Digit 5 (after Petter 1966b). Hindfeet drawn to same length for each species.

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are slightly pro-odont. The upper molar rows are parallel in most genera but divergent anteriorly in Steatomys; occlusal surfaces are cuspidate, or somewhat laminate in Steatomys; cusp rows of M1 and M1 lack longitudinal enamel crests in most genera, interconnecting longitudinal crests present in Malacothrix; posterior cingulum of M1 and M2 is small (Dendroprionomys, Malacothrix, Prionomys), laminar (Steatomys), large and oblong (Dendromus), or exceedingly large and forming an oblong cusp (Megadendromus). In view of the variability and diversity of dendromurines, the employment of the taxon Dendromurinae in muroid classifications has suggested a ‘waste-basket’ for African taxa of highly specialized morphology and hence obscure relationships. The subfamily has been thought to include Lophuromys (Alston 1876), Leimacomys (Thomas 1897, and others), Beamys and Saccostomus (Allen 1939), Petromyscus (Ellerman 1941, Simpson 1945) and Deomys (Simpson 1945), genera that are now affiliated with four other muroid subfamilies (Cricetomyinae, Deomyinae, Leimacomyinae, Petromyscinae). While others have noted that Dendromurinae as arranged in former classifications is polyphyletic (e.g. Rosevear 1969, Carleton & Musser 1984), the cladistic analysis by Denys et al. (1995), based largely on dental traits, has convincingly demonstrated that the subfamily has a polyphyletic origin. However, strict interpretation of their study would exclude Prionomys and Dendroprionomys from the core dendromurines (Dendromus, Malacothrix, Megadendromus, Steatomys), a hypothesis that warrants continued testing in a broader systematic context. As with the uncertainty surrounding its generic contents, Dendromurinae has been alternatively associated with murids (Miller & Gidley 1918, Simpson 1945) or cricetids (Allen 1939, Misonne 1974), and even ranked as a separate family (Chaline et al. 1977). In part, this vacillation hinges on the homology of the lingual accessory cusp (enterostyle) on the lamina of the upper molars,

whether a presumptive t4 corresponding to that in the murid triserial arrangement or a neomorphic acquisition evolved in parallel from a cricetid stock. Phylogenetic implications issuing from recent studies of morphology (Breed 1995, Denys et al. 1995) and DNA sequences (Verheyen et al. 1995, Michaux et al. 2001, Jansa & Weksler 2004) provide stronger vindication for the latter interpretation and support closer affinity of dendromurines to other archaic African muroids, here arranged in Nesomyidae, than to murids proper. Although living species are today restricted to the sub-Saharan region, the subfamily was more widespread in the Tertiary. Fossils are known from the late Miocene of Africa’s Mediterranean rim (Ameur 1984), the Iberian Peninsula (Aguilar et al. 1984) and the Arabian Peninsula (De Bruijn & Whybrow 1994, De Bruijn 1999). Certain extinct genera from the middle Miocene of North-West Africa, Pakistan and Thailand have been referred to Dendromurinae (e.g. Lindsay 1988, Mein & Ginsburg 1997), but their identification as such is disputed (e.g. Tong & Jaeger 1993,Wessels 1996). The earliest indisputable dendromurine thus far documented (Ternania) appears in the middle Miocene of Kenya, about 14 mya (Tong & Jaeger 1993). Representatives of Dendromus appear in the late Miocene of Ethiopia (Geraads 2001), and Steatomys is known from the late Miocene of Namibia (Senut et al. 1992). These genera, as well as Malacothrix, are commonly and widely recorded from the late Pliocene to the Quaternary in southern and eastern Africa (e.g. Jaeger 1979, Denys 1987a, b, 1994, Pocock 1987, Avery 1996, 1998). Four of the six dendromurine genera are monospecific (Dendroprionomys, Malacothrix, Megadendromus, Prionomys), but Dendromus (ten species) and Steatomys (eight species) each contain several species and may prove to be even more diverse after further systematic study. Guy G. Musser & Michael D. Carleton

GENUS Dendromus African Climbing Mice Dendromus Smith, 1829. Zool. J. London, 4: 38. Type secies: Dendromus typus Smith, 1829 (= Mus mesomelas Brants, 1827)

The genus Dendromus contains 11 or 12 species of small climbing mice distributed throughout sub-Saharan Africa. Most species occur in southern, central and East Africa; only one, with a very disjunct distribution, is recorded from West Africa. Three species have wide distributions; most have a small distribution, and others are restricted to one or more widely scattered locations. Typical habitats are long grasslands, bracken, dense scrub, grassy wetlands, and subalpine or alpine vegetation. In some regions, several species are sympatric (e.g. Kivu, E DR Congo; Dieterlen 1971). Species in the genus are characterized by delicate build, small size, soft brown or reddish-brown pelage, long thin tail (100–160% of HB, depending on the species) and specialized feet. Several species have a black mid-dorsal stripe and one species (D. lovati) has three black stripes. The forefoot has only three well-developed digits (Digits 2, 3 and 4); Digit 1 is extremely reduced and Digit 5 is short.The hindfoot has five elongated digits; Digits 2, 3 and 4 are long and slender, Digit 1 is reduced but functional, and Digit 5 is long and semi-opposable and widely separated from other digits (Figure 30). Some species have a nail on Digit 5 of the hindfoot, rather than a claw. Skull characters

Skeleton of Dendromus.

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include: small in size, delicate build, rostrum narrow, zygomatic plate narrow with the masseteric knob at lower corner of the plate, anterior palatal foramina extending to middle row of M1, supraorbital ridges absent and auditory bullae comparatively well developed. Upper incisors small, slightly opisthodont, each with single longitudinal groove. Cheekteeth small; M2 about half size of M1; M3 very small; cusps biseral (two cusps in each row), with additional small lingual cusp on middle lamella of M1 and (to a lesser extent) on M2. Most species of Dendromus climb using the long digits of the forefeet and hindfeet. The opposable Digit 5 of the hindfeet (which can be

opposed to contact Digit 1) and the semiprehensile tail are especially important for climbing and balancing on twigs and grass stems. Some species tend to be more terrestrial than others. They feed primarily on seeds and are mostly nocturnal. Some species, kept in captivity, tend to rather aggressive to conspecifics and to other co-habiting species. Systematically, Dendromus is one of the most difficult genera of African rodents. Rosevear (1969) records that over 50 forms have been given names, and that eight species have been recognized. Bohmann (1942) disposed of these forms in three ‘Rassenkreisen’, roughly equivalent to what are now considered as D. mesomelas, D. melanotis

Table 16. Species in the genus Dendromus, arranged alphabetically. (n. d. = no data.) HB mean (range) (mm)

T mean (range) mm [% HB]

Colour of dorsal pelage

Colour of ventral pelage

Mid-dorsal stripe [width of stripe]

D. insignis

80.1 (68–96)

97 (84–113) [120%]

Rich brown, reddish or buff

Buffy-grey; grey at base

One. Back of head to rump [4 mm]

D. kahuziensis

(77, 82)

(120, 132) [157%]

Dark brown, dusky

White; dark grey at base

74 (57–95)

73 (57–87) [100%]

Warm greyishbrown

Greyish; grey at base

(56–81)a

90 (76–108)a [120–130%]

Medium-brown to rufousbrown; flecked with grey

Greyish; grey at base

D. mesomelas

76 (69–80)a

99 (91–105)a [130–140%]

Rufous-brown; woolly texture

Off-white; hairs dark at base with whitish tip

D. messorius

63.5 (60–68)

88.6 (72–95) [140%]

Gingery-brown

Pure white

D. mystacalis

60.2 (55–65)a

84.8 (75–95) [140–160%]

Bright rufousbrown

Pure white

D. nyasae/ kivu

74 (64–80)

92 (84–105) [124%]

Rich brown

Silvery-white; grey at base

86.4 (80–93) [130–160%]

Pale to dark cinnamon

Species

D. lovati

D. melanotis

a

D. nyikae

66.8 (50–78)

D. oreas

69.3 (60–74)

95.3 (89–104) [135%]

Medium brown

63.6 (60–66)

82.0 (80–84) [130%]

Ochre- to cinnamonbrown; long and silky

D. vernayi

a

68

Creamy-white; white at base (sometimes grey) Dark rufous to greyish-yellow; dark at base Pinkish-buff to grey; grey at base

One. Forehead to base of tail [8 mm] Three. Middle stripe 40–50 mm long, rump to neck; outer stripes 25–30 mm long, rump to shoulders [n.d.]

Claw/nail on Digit 5 of hindfoot Claw (short)

Claw

Notes

Montane areas of eastern Africa. Chin/throat white. Pelage long and dense Mt Kahuzi, E Zaire only. Chin, throat and chest white

Nail

Afroalpine, Ethiopia. Black stripe on head from between eyes to back of head

Nail

Widespread in southern and eastern Africa. Common

Claw

Widespread in southern and eastern Africa. Common

Claw

Mostly West Africa; often found in banana plants

Claw

Widespread in southern and eastern Africa

Claw (short)

Albertine Rift Valley 1300–4200 m

One. Shoulder to base of tail [variable]

Nail

Mostly eastern Africa; higher altitudes. White subauricular patch

One. Indistinct; mid-back to base of tail [3–4 mm]

Claw

Cameroon Mts only. White patches on chin, throat and chest

One [4.5 mm]

n. d.

Chitau, Angola. Rare. White patch on throat and axillary region

One. Shoulder to base of tail [3–4 mm] One. Shoulder to base of tail (absent in some) [variable] Absent (Occasionally faint brown in West Africa) One. Shoulder to base of tail [variable] One. Neck to base of tail [2–3 mm]

Males only.

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and D. mystacalis; but this reduction created additional problems. The genus as a whole possesses a very homogeneous morphology with respect to skull and dentition, but a great variability of pelage colour and pattern, so that classifying the species often depends on rather few details (Heim de Balsac & Lamotte 1958, Dieterlen 1971). Musser & Carleton (2005) recognize 12 species and list 44 synonyms for the whole genus.The distributional limits of many species are unresolved, and karyological information is unavailable for most species. The

genus is in need of further study and revision. Here, 11 species are recognized; the twelfth species of Musser & Carleton (2005), D. leucostomus, is placed as a synonym of D. melanotis. The species are distinguished by body size, pelage colour, presence or absence and form of the mid-dorsal stripe, presence of nail or claw on Digit 5 of hindfoot, and distribution (Table 16). Fritz Dieterlen

Dendromus insignis MONTANE AFRICAN CLIMBING MOUSE Fr. Souris arboricole des montagnes; Ger. Gebirges-Klettermaus Dendromus insignis (Thomas, 1903). Ann. Mag. Nat. Hist., ser. 7, 12: 341. Nandi, Kenya.

Taxonomy Originally described in the genus Dendromys. This species belongs to the mesomelas species-complex, although there is some doubt about which forms may be included within insignis (Thomas 1916b, Bohmann 1942, Ellerman et al. 1953).The holotype may be unrepresentative of the species as a whole, and its relationship to mesomelas is uncertain. According to Musser & Carleton (2005), most literature references to D. mesomelas in montane habitats north of the southern African sub-region prior to 1991 actually represent either D. insignis or D. nyasae (kivu), both of which may co-occur in the Albertine Rift Valley mountains. Synonyms: abyssinicus, kilimandjari, percivali. Subspecies: none. Chromosome number: not known (identity of Matthey’s [1967, 1970] specimens is questionable). Description Very small climbing mouse with broad mid-dorsal stripe; the largest species of Dendromus. Pelage long (8–9 mm) and dense; shows considerable individual variation (see below). Dorsal pelage rich brown tending to reddish or buff; hairs grey at base, brown at tip. Underfur dark grey, which may provide greyish tinge to pelage. Broad (ca. 4 mm) black mid-dorsal stripe from back of head to base of tail. Ventral pelage and flanks mainly buff or buffygrey; hairs grey at base, buff at tip. Small area of chin and throat pure white. Indistinct black longitudinal stripe (ca. 10–20 mm in length) on head. Ears with short brown or black hairs. Hindfeet silver-grey. Hindfoot with five digits; Digit 1 short with nail, Digit 5 long with claw. Tail very long (ca. 120% of HB, but relatively short compared with other Dendromus spp.), dark brown above, much paler below with long silvery-grey hairs. Nipples: 2 + 2 = 8. Geographic Variation Pelage variable: longer on individuals living above 2000 m; darker in colour in young and subadults than in adults; ventral colouration tends to be more variable than dorsal colour. Similar Species D. nyasae (kivu). HB on average slightly smaller (mean 73.4 mm); tail on average slightly shorter (mean 91.8 mm) but of similar relative length (ca. 124% of HB); mid-dorsal stripe 2–3 mm wide, from neck to base of tail; pelage not as dense. D. mesomelas. HB on average smaller (mean 74–76 mm); tail on average longer (mean 99–103 mm) and relatively longer (130– 140% of HB); narrower mid-dorsal stripe, sometimes absent; pelage woolly; larger geographic range.

Dendromus insignis

Distribution Endemic to Africa. Afromontane–Afroalpine BZ of eastern Africa. Recorded from the Rwenzori Mts (up to 4500 m; Misonne 1963, Verschuren et al. 1983); Virunga Range and Mitumba Mts west of L. Kivu (1700–3300 m; Dieterlen 1976a); Mt Kenya (2300–4300 m; Hollister 1919), Matthews Range, Mau Escarpment, Aberdare Range, Mt Kilimanjaro (3500– 4700 m) and Uluguru Mts (Bohmann 1942, F. Dieterlen unpubl.). The total range of the species is uncertain because of the uncertain taxonomic status of some related forms and lack of material from large areas. Maybe sympatric with D. nyasae (kivu) at some localities in the mountains of the Albertine Rift Valley (Dieterlen 1971, 1976a, as D. mesomelas kivu). Habitat In the cultivated regions near L. Kivu, DR Congo (1500– 2000 m), lives with other species of Dendromus in open comparatively dry dense grassy areas (Dieterlen 1971) and, less frequently, in marshes and moist herbaceous vegetation. Also occurs in sparse bamboo and secondary forest above 2000 m, and in wet grass and 171

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herbaceous vegetation in sub-alpine and alpine zones above 3000 m (Kingdon 1974, Dieterlen 1976a). Abundance May be abundant at lower altitudes, but becomes progressively less common with increasing altitude.

Predators, Parasites and Diseases Small carnivores, birds and snakes are potential predators. Predatory ants are certainly a great danger for young in the nest during the first three weeks of life, especially when nests are on the ground or in burrows. Conservation

Adaptations Probably nocturnal and diurnal. Although comparatively large and heavy (15 g) and often found on the ground, these mice are agile climbers. The opposable Digit 5 of hindfoot and the semi-prehensile tail are used to provide support when climbing. More terrestrial than many species of Dendromus. Because of its terrestrial habits, this species is trapped more often than other Dendromus spp. Most nests seem to be on the ground or in burrows. Foraging and Food Omnivorous and/or insectivorous. Two stomachs contained remains of well-chewed seeds (including sorghum), reddish berries and arthropods (Dieterlen 1971, 1976a). Social and Reproductive Behaviour No information, but it is presumed that individuals are mostly or entirely solitary. Reproduction and Population Structure Near L. Kivu, most young born in the wet season between Sep and May. Gestation: not known (although probably similar to D. nyasae [kivu]). Embryo number: 4 (n = 3). Young extremely altricial at birth; postnatal development is slow. Sex ratio: 69 : 31% (ca. 7 : 3; n = 48). In most respects, reproduction and population structure appear to be similar to D. nyasae (kivu) (Dieterlen 1971).

IUCN Category: Least Concern.

Measurements Dendromus insignis HB: 80.1 (68–96) mm, n = 34 T: 97.0 (84–113) mm, n = 34 HF: 22.3 (20.5–24) mm, n = 34 E: 15.7 (13–18) mm, n = 34 WT: 15.1 (9–27) g, n = 34 GLS: 24.1 (21.4–25.7) mm, n = 19 GWS: 12.1 (11.3–12.9) mm, n = 14 M1–M3: 3.8 (3.7–4.1) mm, n = 22 West of L. Kivu, DR Congo (Dieterlen 1971, 1976a; F. Dieterlen unpubl.; SMNS, ZFMK) Individuals from higher altitudes of Mt Kenya are slightly larger (e.g. mean HB: 85.4; mean T: 102.1) Key References & Carleton 1993.

Bohmann 1942; Dieterlen 1971, 1976a; Musser Fritz Dieterlen

Dendromus kahuziensis KAHUZI AFRICAN CLIMBING MOUSE Fr. Souris arboricole du Mont Kahuzi; Ger. Kahusi-Klettermaus Dendromus kahuziensis Dieterlen, 1969. Z. Säugetierk. 34: 348–349. SSW slopes of Mount Kahuzi, Kivu, DR Congo. 2100 m.

Taxonomy Very distinct species of Dendromus with an extremely long tail, broad mid-dorsal stripe and long gracile rostrum. Systematic relationships unknown. Only two specimens known. Synonyms: none. Chromosome number: not known. Description Small climbing mouse with a wide mid-dorsal stripe and very long tail; a comparatively large species of Dendromus. Dorsal pelage dusky dark-brown; hairs greyish-black with medium brown tip. Mid-dorsal blackish stripe from head (at level with the eyes) to base of tail; stripe broad (ca. 8 mm), broader than in other species of Dendromus. Flanks paler; hairs with pale brownish tip. Chin, throat and chest white; hairs dark grey at base. Head with pale brown on cheeks and behind ears. Eyes with black rings, which extend anteriorly on to nasal region. Ears with sparse short black and reddish hairs. Forefoot with three long digits; Digit 1 rudimentary, Digit 5 minute. Hindfoot with five digits; Digit 1 short without nail or claw, Digit 5 long and opposable with long claw. Tail very long (ca. 157% of HB), comparatively longer than in other Dendromus spp., with short dark bristles, darker above, paler below. Skull with long nasals and rostrum. Nipples: not known. Geographic Variation None recorded.

Dendromus kahuziensis

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Dendromus lovati

Similar Species D. insignis. HB similar; tail shorter (ca. 80 mm and ca. 120% of HB); mid-dorsal stripe broad (ca. 4 mm) from back of head to base of tail. D. nyasae (kivu). HB slightly smaller; tail shorter (84–105 mm ca. 124% of HB); long mid-dorsal stripe (2–3 mm wide) from behind head to base of tail. D. mystacalis. HB smaller; tail shorter (75–97 mm), but relatively long (140–160% of HB); mid-dorsal stripe from shoulders to base of tail. Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Recorded only from the montane forest on the SSW slopes of Mt Kahuzi, east of L. Kivu, DR Congo. Habitat The only known habitat is a small deep valley, with a small stream in the bottom, covered with dense montane forest and bamboo Arundinaria alpina (Dieterlen 1969a). Other common plants include the trees Polyscias fulva, Symphonia globulifera and Neobutonia macrocalyx, and the ferns Cyathea manniana and Marattia fraxinea. The only species of Dendromus living in montane forest.

Foraging and Food One stomach contained homogeneous well-chewed vegetable material, probably composed of seeds and fruits; green vegetation and animal remains were not present. Conservation IUCN Category: Critically Endangered. The very small known geographic range, and presumably low population numbers, are cause for concern. Schlitter (1989) classified the species as ‘Rare’. Measurements Dendromus kahuziensis HB: 82, 77 mm, n = 2 T: 132, 120 mm, n = 2 HF: 22, 21 mm, n = 2 E: 15, 14 mm, n = 2 WT: 12, 10 g, n = 2 GLS: 23.7, 23.4 mm, n = 2 GWS: 10.8, 11.0 mm, n = 2 M1–M3: 3.5, 3.4 mm, n = 2 E DR Congo (Dieterlen 1969, 1976a) Key References

Dieterlen 1969a, 1976a.

Adaptations The structure of the forefoot and hindfoot, and the long tail, suggest that these mice are good climbers, and probably spend most their time above the ground (as do species of Dendromus that live in grasslands). The two known specimens were caught in traps on the ground, so it seems that they sometimes descend to the ground, perhaps when foraging.

Fritz Dieterlen

Dendromus lovati LOVAT’S AFRICAN CLIMBING MOUSE Fr. Souris arboricole de Lovat (Souris arboricole des plateaux éthiopien); Ger. Lovats Klettermaus Dendromus lovati (de Winton, 1900). Proc. Zool. Soc. Lond. 1899: 986. (publ. 1900). Menagesha, Ethiopia. 2800 m.

Taxonomy Originally described in the genus Dendromys. A very distinctive species, sometimes placed in a subgenus Chortomys by itself. Synonyms: none. Chromosome number: 2n = 44 (Lavrenchenko et al. 1997). Description Distinctive very small dendromurine, with three very obvious stripes over the rump (quite unlike any other species of Dendromus). Pelage dense and soft. Dorsal pelage warm greyishbrown, sometimes with a slight russet tinge; hairs dark grey at base, brown or buff at tip; some longer hairs with black tips. Three black longitudinal stripes on back, each widening anteriorly and highlighted by paler sandy edges; middle stripe (ca. 40–50 mm long, 8 mm wide on mid-back, tapering to 0–2 mm at base of tail) extends from neck to rump; two outer stripes (ca. 25–30 mm long, 5–6 mm on mid-back tapering to 0–2 mm at base of tail) extend from chest to rump. Middle stripe may be partially split lengthwise into a double median stripe. Ventral pelage greyish; hairs grey at base, offwhite at tip. Head similar in colour to back with small longitudinal black stripe from back of head to between eyes. Ears sparsely furred, sandy, with patch of black hairs on lower part of inner surface and on anterior part of outer surface. Whitish subauricular spot. Fore- and hindfeet sandy-brown. Hindfoot with five digits; Digit 1 short with

Dendromus lovati

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nail, Digit 5 with nail-like claw.Tail long (ca. 100% of HB) with short hairs, dark above, paler below; comparatively short for a species of Dendromus. Nipples: 2 + 1 = 6.

Social and Reproductive Behaviour Little known. Appears solitary, one individual was ‘dug out from nest in tuft of grass’ (collector’s notes: R. E. Drake-Brockman, BMNH)) and another was found in a nest beneath a boulder.

Geographic Variation None recorded. Similar Species No other species of Dendromus has three longitudinal stripes on the back. Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Confined to the high plateaux of Ethiopia, from 2500 to 3900 m. Known from only about nine locations (Yalden et al. 1976,Yalden & Largen 1992, Sillero-Zubiri et al. 1995b). Habitat Typically found in highland grasslands, but extending into the dry Heliochrysum heathland at 3900 m on Tullu Deemtu, Bale (Sillero-Zubiri et al. 1995a). Abundance Uncommon, known only from about 30 specimens (Yalden et al. 1976, Yalden & Largen 1992). Recent collecting in Ethiopia (1968–98, n = ca. 6300 rodents) yielded only 23 individuals of this species (Müller 1977, Rupp 1980, Yalden 1988, Sillero-Zubiri et al. 1995a, Afework Bekele 1996a, Lavrenchenko et al. 1997, Nievergelt et al. 1998). Adaptations Nocturnal and terrestrial. Compared with other Dendromus, this short-tailed species appears to have no climbing ability. Several specimens have been caught at night in open grassland, where they moved ‘in short hops’ (collector’s notes: R. E. Cheesman, BMNH). Individuals show no obvious adaptations to living in harsh high-altitude environments, leading Sillero-Zubiri et al. (1995a) to speculate that they might hibernate during the dry season (when, under clear skies, it is extremely cold at night). However, captures have been made throughout the year (author’s records, collectors’ notes).

Reproduction and Population Structure A juvenile " trapped in Dec and a parous ! in Jan in Bale, and a juvenile caught in Mar (label, BMNH), suggest that reproduction occurs in the dry season. Predators, Parasites and Diseases

No information.

Conservation IUCN Category: Least Concern. The highland grassland (woina dega) habitat is threatened by continuing modification and destruction by humans and their livestock. Measurements Dendromus lovati HB: 74 (57–95) mm, n = 21 T: 73 (57–87) mm, n = 21 HF: 18 (17–20) mm, n = 17 E: 16 (15–18) mm, n = 16 WT: 16.4 (11–23) g, n = 11 GLS: 19.4 (18.0–20.2) mm, n = 6 GWS: 10.5 (9.8–11.1) mm, n = 4 M1–M3: 3.3 (3.1–3.7) mm, n = 9 Ethiopia Body measurements and weight: D. W.Yalden unpubl., Sillero-Zubiri et al. 1995 Skull measurements: D. W.Yalden unpubl. Key References Müller 1977; Nievergelt et al. 1998; SilleroZubiri et al. 1995a;Yalden et al. 1976;Yalden & Largen 1992. D. W. Yalden

Foraging and Food Unknown, but probably granivorous.

Dendromus melanotis GREY AFRICAN CLIMBING MOUSE Fr. Souris arboricole gris; Ger. Graue Klettermaus Dendromus melanotis, Smith 1834. S. Afr. Quart. J. 2: 158. Near ‘Port Natal’ (= Durban), South Africa.

Taxonomy The taxonomic status of this species is uncertain, as manifested by karyotypic variability within the species and by the numerous synonyms currently recognized. The form leucostomus (known only from the type locality in Angola) is maintained as a synonym as in Musser & Carleton (1993) but raised to species status by Musser & Carleton (2005), following Hill & Carter (1941). CrawfordCabral (1998) recorded that the dorsal pelage of leucostomus does not have a mid-dorsal stripe (cf. typical D. melanotis). Here, pending full taxonomic revision, leucostomus is retained as a synonym of D. melanotis. Synonyms: arenarius, basuticus, capensis, chiversi, concinnus, exoneratus, insignis (Shortridge & Carter, 1938, not of Thomas, 1903), leucostomus, nigrifrons, pallidus, pecilei, pretoriae, shortridgei, spectabilis, subtilis, thorntoni, vulturnus. Subspecies: none. Chromosome number: 2n = 36 (Matthey in Robbins & Baker 1978); 2n = 52 (Dippenaar et al. 1983).

Description Very small slender mouse with soft woolly pelage and long slender tail. Dorsal pelage medium brown to rufousbrown, sometimes flecked with ashy-grey; hairs grey on basal twothirds, pale brown on terminal third, some hairs with black tip. Dark stripe (usually 3–4 mm in width) extends along mid-dorsal line from between shoulders to base of tail. Ventral pelage white to greyishwhite; hairs grey at base, off-white to pale grey at tip. Head with pointed muzzle and long vibrissae. Ears small and rounded, dark. Small white patch at base of ears. Limbs short. Forefeet with three long functional digits, Digits 1 and 5 greatly reduced but not absent. Hindfoot with five digits; Digit 1 short with nail, Digit 5 with nail (see also D. nyikae). Tail long (ca. 120–130% of HB), brownish above, paler below, sparsely haired. Nipples: 2 + 2 = 8.

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Dendromus melanotis

Geographic Variation Pelage colour varies slightly throughout range. In the form nigrifrons (from e.g. Namibia, Kenya, Zambia), the mid-dorsal stripe extends to the forehead although usually indistinct between the ears and on the neck. Similar Species D. mystacalis. Similar in size and proportions; pelage rufous-brown; tail 140–160% of HB. D. nyikae. Larger, head and body usually well over 70 mm; tail relatively longer (130–160% of HB). D. mesomelas. On average larger, head and body generally well over 70 mm; tail relatively longer (130–140% of HB). Distribution Endemic to Africa. Recorded in many biotic zones: Zambezian Woodland BZ, extending marginally into parts of Highveld and South-West Cape BZs of South Africa; Afromontane– Afroalpine BZ of Ethiopia; and (?) Guinea Savanna BZ of West Africa. Ranges widely from S and E South Africa through Botswana, E Namibia (Matson & Blood 1994), Zimbabwe, S Mozambique, Zambia, Malawi and S Angola (Crawford-Cabral 1998). Isolated populations in W Uganda (Wilson 1995), Tanzania and Kenya. Status in Ethiopia uncertain; distribution poorly known, and records may represent D. mystacalis (Yalden et al. 1976). In West Africa recorded at scattered localities in Nigeria, Benin, Liberia and Guinea (Happold 1987). Habitat Inhabits a wide range of habitats. In southern Africa inhabits stands of tall Hyparrhenia grassland (Lynch 1994), short montane grassland (Rowe-Rowe & Meester 1982a,Taylor 1998), dry Kalahari scrub, fringes of rivers, dry Baikiaea woodland (Smithers 1971) and flood-plains (Sheppe & Haas 1981). May recolonize burnt grasslands within one month of fire (Rowe-Rowe & Meester 1981). Abundance Relatively common. Third commonest species of small mammal in several different high-altitude grassland habitats in KwaZulu–Natal Province, South Africa (Rowe-Rowe & Meester 1982a). Detailed density estimates not available. Adaptations Nocturnal. Limbs and tail modified for a semiarboreal existence. Long, slender digits used to grip and climb thin stalks while long, prehensile tail provides balance. Weaves a grass nest with a single entrance, which is usually attached to grass stems or shrubs up to 1 m above the ground. Nest used only during the breeding season (Smithers 1971). May also use burrows up to 50 cm deep leading to a nest chamber with an emergency exit on the opposite side (De Graaff 1981). Foraging and Food Predominantly granivorous. Frequency of occurrence of food types (n = 14 stomach contents) in subalpine grasslands of Drakensburg Mts, South Africa: seeds 100%, arthropods 24% and vegetable material 0% (Rowe-Rowe 1986). Social and Reproductive Behaviour Poorly known. Thought to be predominantly solitary and territorial (Smithers 1971, Kingdon 1974). Known to fight ferociously with D. mystacalis in captivity. Parents reported to remain with offspring for some time after weaning.

Dendromus melanotis

Reproduction and Population Structure Little known. Breeding possibly confined to the wet season in southern Africa. Gravid !! with 2–8 embryos collected between Nov and Apr (Smithers 1971, Lynch 1994). In South Africa, juveniles captured at the end of the wet season during Apr and May (Rowe-Rowe & Meester 1982b). Predators, Parasites and Diseases Dendromus spp. are difficult to distinguish in raptor pellets (Coetzee 1972), but remains have been found in the pellets of Barn Owls Tyto alba (Vernon 1972) and Black-shouldered Kites Elanus caeruleus (Mendelsohn 1982). Conservation

IUCN Category: Least Concern.

Measurements Dendromus melanotis HB (""): 68 (56–81) mm, n = 32 HB (!!): 70 (60–86) mm, n = 17 T (""): 90 (76–108) mm, n = 26 T (!!): 83 (71–113) mm, n = 21 HF (""): 18 (16–21) mm, n = 33 HF (!!): 17 (16–20) mm, n = 18 E (""): 14 (12–18) mm, n = 30 E (!!): 16 (13–19) mm, n = 20 WT (""): 7.4 (6–10) g, n = 11 WT (!!): 7.0 (4–12) g, n = 19 GLS: 20.1 (19.8–21.0) mm, n = 17 GWS: 10.4 (10.0–11.2) mm, n = 15 M1–M3: 3.0 (2.8–3.3) mm, n = 22 Body measurements and weights: throughout range (De Graaff 1981) Skull measurements: Botswana, South Africa (Roberts 1951) Key References

Kingdon 1974; Rowe-Rowe & Meester 1982a. A. Monadjem 175

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Dendromus mesomelas BRANTS’S AFRICAN CLIMBING MOUSE Fr. Souris arboricole noisette; Ger. Brants Klettermaus Dendromus mesomelas (Brants, 1827). Het. Geslacht der Muizen, p. 122. Sunday’s River, South Africa.

Taxonomy Originally described in the genus Dendromys. The name mesomelas refers to the large Dendromus of southern and central Africa (Musser & Carleton 1993) in which Digit 5 of the forefoot is absent. Some names formally included in D. mesomelas (see Misonne 1974) are now regarded as valid species (i.e. D. insignis, D. nyasae, D. oreas and D. vernayi). Synonyms: ayres, major, pumilio, typicus, typus. Subspecies: none. Chromosome number: not known. Description Very small slender mouse with soft woolly pelage and very long slender tail. Dorsal pelage rufous-brown; hairs dark grey, rufous-brown to gingery-brown terminally. Dark mid-dorsal stripe of variable intensity from shoulders to base of tail; absent in some individuals. Ventral pelage off-white; hairs dark at base with whitish tip. Head with pointed nose and long vibrissae. Ears small and rounded. Limbs short. Forefeet with three long functional digits, Digit 1 greatly reduced and Digit 5 absent. Hindfoot with five digits; Digit 1 short with nail, Digit 5 long and opposable with claw. Tail very long (ca. 130–140% of HB), brownish above, paler below and sparsely haired. Nipples: 2 + 2 = 8. Geographic Variation None recorded. Dendromus mesomelas

Similar Species D. mystacalis. Smaller, HB less than 65 mm; ventral pelage white, hairs with white base. Tail 140–160% of HB. D. nyikae. Dorsal pelage pale to dark cinnamon suffused with grey; ventral pelage white, hairs with white base. Tail long (130–160% of HB). D. melanotis. Smaller, HB usually less than 70 mm; dorsal pelage ashygrey; ventral pelage greyish-white. Tail 120–130% of HB. Distribution Endemic to Africa. Zambezian Woodland BZ, extending to southern part of Somalia–Masai Bushland BZ and to Coastal Forest Mosaic BZ (in South Africa). Distribution disjunct. Recorded in S and E South Africa, W Swaziland, N Botswana, Namibia (Caprivi Strip), Mozambique (Gorongosa Mt) (De Graaff 1981), NE and NW Zambia (Ansell 1978), S and N Malawi (Ansell & Dowsett 1988), SE DR Congo (Musser & Carleton 1993) and C Tanzania (Swynnerton & Hayman 1950).

Adaptations Mostly nocturnal, but may also be active during the day. Limbs and tail modified for a semi-arboreal existence. Long, slender digits used to grip and climb thin stalks while long, prehensile tail provides balance. Weaves a grass nest, which may be placed either above or below ground (Kingdon 1974). Also occupies nests of birds, e.g. Ploceus spp. (De Graaff 1981). Believed to be more terrestrial than other members of the genus (see also D. lovati). Foraging and Food Predominantly granivorous. Proportional contribution of food types in stomach contents: 12% vegetable material, 87% seeds, 1% arthropods (n = 5, Swaziland; Monadjem 1997b). Elsewhere, in Botswana and South Africa, insects contribute a greater proportion of the diet (Smithers 1971, Rowe-Rowe 1986). Seasonal availability may limit the consumption of insects to the wet season. Social and Reproductive Behaviour

No information.

Habitat Inhabits a wide range of grassland habitats mostly in temperate environments. In southern Africa also inhabits swamps and damp grasslands (Hanney 1965, Taylor 1998) as well as afromontane forest (Rowe-Rowe & Meester 1982, Monadjem 1998a). Prefers wet moist habitats and is absent from hot, low-lying river basins, arid savannas and miombo woodland.

Reproduction and Population Structure Reproduction possibly confined to the wet season in southern Africa, but gravid !! have been collected in the dry season in Malawi (Hanney 1965). Embryo number: 2–6 (Hanney 1965, Taylor 1998).

Abundance Relatively common. No density estimates available, but relatively numerous in leaf-litter of afromontane forest in Swaziland (A. Monadjem unpubl.).

Predators, Parasites and Diseases Dendromus spp. are difficult to distinguish in raptor pellets (Coetzee 1972), but remains have been found in the pellets of Barn Owls Tyto alba (Vernon 1972) and Black-shouldered Kites Elanus caeruleus (Mendelsohn 1982). Ectoparasites include the fleas Ctenophthalamus verutus, C. cophurus,

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Dendromus messorius

Dinopsyllus grypurus and Nosopsyllus incisus. May be susceptible to plague (De Graaff 1981). Conservation

IUCN Category: Least Concern.

Measurements Dendromus mesomelas HB (""): 76 (69–80) mm, n = 5 HB (!!): 74 (67–85) mm, n = 7 T (""): 99 (91–105) mm, n = 4 T (!!): 103 (94–109) mm, n = 7 HF (""): 20 (19–21) mm, n = 4 HF (!!): 20 (18–22) mm, n = 7 E (""): 18 (15–21) mm, n = 3

E (!!): 14 (12–17) mm, n = 7 WT (""): 12.0 (11–13) g, n = 4 WT (!!): 10.6 (9–15) g, n = 5 GLS: 22.4 (20.8–24.7) mm, n = 9 GWS: 11.5 (10.7–12.5) mm, n = 9 M1–M3: 3.3 (3.1–3.5) mm, n = 10 South Africa Body measurements and weights: De Graaff 1981 Skull measurements: Roberts 1951 Key References

De Graaff 1981; Hanney 1965. A. Monadjem

Dendromus messorius BANANA AFRICAN CLIMBING MOUSE Fr. Souris arboricole de bananier; Ger. Bananen-Klettermaus Dendromus messorius (Thomas, 1903). Ann. Mag. Nat. Hist., ser. 7, 12: 340. Efulen, Cameroon.

Taxonomy Originally described in the genus Dendromys. Considered to be a synonym or subspecies of D. mystacalis (Rosevear 1969, Misonne 1974, Delany 1975 [referring to ruddi]) or as a valid species (Hatt 1940a, Verheyen & Verschuren 1966, Dieterlen 1971, Musser & Carleton 1993, 2005). Synonyms: haymani, kumasi, ruddi. Subspecies: none. Chromosome number: not known. Description Very small slender climbing mouse with long tail and without mid-dorsal stripe. Dorsal pelage gingery-brown; hairs dark grey on basal three-quarters, gingery-brown on terminal one-quarter. Usually without mid-dorsal stripe (cf. some other Dendromus spp.); occasionally very faint brownish stripe is visible. Flanks paler than back.Ventral pelage pure white; clearly delineated from colour of flanks. Head similar colour to dorsal pelage. Ears comparatively large covered with short ginger hairs. Upper lips, lower lips, cheeks, throat and chest whitish. Long stiff black vibrissae on muzzle. Fore- and hindlimbs whitish. Forefoot with three functional digits (Digits 2, 3 and 4). Hindfoot with five digits; Digit 1 short with nail (or absent), Digit 5 long and opposable with claw. Tail very long (ca. 140% of HB), dark, with short blackish bristles. Skull: upper incisors opisthodont. Nipples: 2 + 2 = 8. Geographic Variation Some individuals have a faint brownish stripe (e.g. kumasi from Ghana [Rosevear 1969]). Similar Species D. mystacalis. Similar size; well-developed dark mid-dorsal stripe; ventral pelage whitish; partially parapatric. D. mesomelas. Larger; well-developed dark mid-dorsal stripe; ventral pelage off-white, hairs dark at base with whitish tip; partially parapatric. Distribution Endemic to Africa. Rainforest BZ and Northern and Eastern Rainforest–Savanna Mosaics. Also Afromontane– Afroalpine BZ in Uganda and E DR Congo. Recorded from Ghana, Togo, Benin, E Nigeria (Umuahia only), Cameroon, DR Congo and Uganda. Distribution disjunct. Limits uncertain; may also occur in S

Dendromus messorius

Sudan and W Kenya (Musser & Carleton 2005). Recorded at 2000– 3000 m on Mt Elgon (Clausnitzer & Kityo 2001, as D. mystacalis ruddi; see Musser & Carleton 2005). Habitat Preferred habitat is banana plants in farmlands. Also found among sweet potatoes (Hatt 1940a), in holes in trees, in fissures, cracks and trunks of trees in gallery forest, and orange trees (Verheyen & Verschuren 1966). In Cameroon, recorded in long grass (Rosevear 1969). Abundance May be common in some localities; in Garamba N. P., NE DR Congo, commoner than sympatric Dendromus mesomelas and D. mystacalis (Verheyen & Verschuren 1966). 177

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Family NESOMYIDAE

Adaptations Nocturnal. At Medje, NE DR Congo, the nest of a ! with three young was ‘inserted between the bases of banana leaves, close to the stem, about eight feet from the ground. The nest was hardly visible being well concealed between the large bases of the closely growing banana leaves. Sliced up banana leaves were the only material used outside. On the underside larger pieces were utilised, but inside the leaves were so finely split that the material looked like fine grass’ (H. Lang in Hatt 1940a). This nest was about the size of a man’s fist, roundish in shape, with a single entrance. Two other nests, each occupied by a single ", were similar in construction. Nests also built in long grass, and a bush (type specimen) (Cameroon; Rosevear 1969). Foraging and Food Vegetable material present in two stomachs (Hatt 1940a). Also feeds on fruits of Sarcocephalus esculentus (Verheyen & Verschuren 1966). Social and Reproductive Behaviour Assumed to be solitary. The nests referred to above each occupied by a single adult animal.

Predators, Parasites and Diseases No information. Conservation IUCN Category: Least Concern. In Uganda, conserved in four National Parks (Mgahinga N. P., Bwindi Inpenetrable N. P., Rwenzori N. P. and Semliki N. P. [Wilson 1995]). Measurements Dendromus messorius HB: 63.5 (60–68) mm, n = 13 T: 88.6 (72–95) mm, n = 13 HF: 16.7 (15–19) mm, n = 13 E: 13.2 (11–14) mm, n = 13 WT: 8.8 (7.6–10.5) g, n = 10 GLS: 21.0 (20.2–21.6) mm, n = 7 GWS: 11.1 (10.5–11.6) mm, n = 7 M1–M3: 3.2 (2.9–3.5) mm, n = 7 Uganda (BMNH) Key References Verschuren 1966.

Hatt 1940a; Rosevear 1969; Verheyen &

Reproduction and Population Structure No information. D. C. D. Happold

Dendromus mystacalis CHESTNUT AFRICAN CLIMBING MOUSE Fr. Souris arboricole de Heuglin; Ger. Kastanienbraune Klettermaus Dendromus mystacalis Heuglin, 1863. Nova Acta Acad. Caes. Leop.-Carol., Halle 30: 2, suppl. 5. Baeschlo region, Ethiopia.

Taxonomy A smaller version of D. mesomelas of southern Africa. Probably closely related to D. mesomelas, the two species forming a species-pair (Avery 1998; see also Musser & Carleton 2005). Synonyms: acraeus, ansorgei, capitis, jamesoni, nairobae, ochropus, pallescens, pongolensis, uthmoelleri, whytei. Subspecies: none. Chromosome number: 2n = 38 (Matthey in Robbins & Baker 1978). Description Very small slender mouse with soft woolly pelage and very long slender tail. Dorsal pelage bright rufous-brown. Dark stripe on mid-dorsal line from shoulders to base of tail.Ventral pelage whitish, ventral hairs with white base. Head with pointed nose and long vibrissae. Ears small and rounded. Limbs short. Forefeet with three long functional digits, Digit 1 greatly reduced, Digit 5 absent. Hindfoot with five digits; Digit 1 short with nail, Digit 5 long and opposable with claw. Tail very long (ca. 140–160% of HB), brownish above, paler below, sparsely haired. Nipples: 2 + 2 = 8. Geographic Variation None recorded. Similar Species D. melanotis. On average larger; dorsal pelage ashy-grey; tail 140– 160% of HB. D. nyikae. On average larger; dorsal pelage pale to dark cinnamon; tail 130–160% of HB. D. mesomelas. Larger; black mid-dorsal stripe sometimes absent; ventral hairs with dark base; tail 130–140% of HB. D. messorius. Dorsal stripe absent. West Africa and central Africa only.

Dendromus mystacalis

Distribution Endemic to Africa. Zambezian Woodland BZ and parts of Eastern Rainforest–Savanna Mosaic and Afromontane– Afroalpine BZ in eastern Africa. Occurs widely from E South Africa northwards through Zimbabwe, Mozambique and Zambia. Ranges westwards in N Botswana and Angola. Isolated populations in E DR

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Dendromus nyasae (kivu)

Congo, Uganda, S Sudan, N Tanzania and S Kenya, and Ethiopia (Yalden et al. 1976). Habitat Occupies a wide range of savanna habitats; presence of dense grass cover is essential. In Swaziland, high vegetation density in first 10 cm above ground appears to be a critical habitat feature (Monadjem 1997a). Tall stands of Hyperthelia-Hyparrhenia grassland are favoured, but also occurs in rank vegetation fringing wetlands, drainage lines and afromontane forests. Predominantly a low-altitude species occurring below 2000 m (1200 m in southern Africa) generally preferring drier conditions than D. nyasae (kivu)(Dieterlen 1971). In Swaziland, associated with hilly landscapes (Monadjem 1999a). Abundance Relatively common to abundant. Population densities not easy to determine because individuals do not regularly enter rodent traps. Densities of up to 19.8/ha reported from DR Congo (Misonne 1963) and 2.5/ha from Swaziland (A. Monadjem unpubl.).

Reproduction and Population Structure Breeding confined to the wet season with juvenile recruitment mostly Jan–Mar (Zimbabwe; Smithers & Wilson 1979). Embryo number: usually 3–4, but up to eight (Rautenbach 1982). In Swaziland, number of individuals trapped fluctuated seasonally; highest numbers in dry season between Jun–Oct (Monadjem & Perrin 2003).‘Disappearance’ of these mice in the wet season was attributed to a change in foraging tactics rather than population fluctuations. In southern Africa, most grasses set seed in the late wet season, at which time individuals of this species probably take seeds from the grass stalks, rather than from the ground. Predators, Parasites and Diseases Dendromus spp. are difficult to distinguish in raptor pellets (Coetzee 1972), but remains have been found in the pellets of Barn Owls Tyto alba (Vernon 1972) and Black-shouldered Kites Elanus caeruleus (Mendelsohn 1982). Conservation

IUCN Category: Least Concern.

Adaptations Nocturnal. Limbs and tail modified for a semiarboreal existence. Long, slender digits used to grip and climb thin stalks while long, prehensile tail provides balance. Weaves a grass nest with multiple entrances, in which offspring are raised. Adults also use nest for resting. Nest usually located in tall grass about 1 m above ground (Monadjem 1998a) but may also be located higher up in gardens (Kingdon 1974). Known to utilize burrows and disused nests of weaver birds.

Measurements Dendromus mystacalis HB (""): 60.2 (55–65) mm, n = 6 HB (!!): 54.7 (47–62) mm, n = 3 T (""): 84.8 (75–95) mm, n = 6 T (!!): 90.3 (82–97) mm, n = 3 HF (""): 16.8 (15.9–17.6) mm, n = 6 HF (!!): 17.4 (16.4–19.5) mm, n = 3 Foraging and Food Omnivorous. Diet (assessed by stomach E: 9.3 (7–12) mm, n = 3 contents): 44% vegetable material, 40% seeds, 16% arthropods WT (""): 7.2 (5–10) g, n = 6 (n = 6, Swaziland; Monadjem 1997b). Forages both on the ground WT (!!): 8.7 (6–11) g, n = 3 and on tall, dense grass. GLS: 20.2 (18.7–21.6) mm, n = 16 GWS: 10.3 (9.4–10.8) mm, n = 16 Social and Reproductive Behaviour In Swaziland, pairs were M1–M3: 3.1 (2.8–3.4) mm, n = 16 regularly trapped together in same rodent live-trap, suggesting that Body measurements and weights: Swaziland (Monadjem 1998a) individuals often forage in pairs. Known to fight ferociously with D. Skull measurements: South Africa (Roberts 1951) melanotis in captivity. Key References Dieterlen 1971; Monadjem & Perrin 2003. A. Monadjem

Dendromus nyasae (kivu) KIVU AFRICAN CLIMBING MOUSE Fr. Souris arboricole du Kivu.; Ger. Kivu-Klettermaus Dendromus nyasae Thomas, 1916. Ann. Mag. Nat. Hist., ser. 8, 18: 241. Nyika Plateau, Malawi. Dendromus kivu Thomas, 1916. Ann. Mag. Nat. Hist., ser. 8, 18: 242. Buhamba, Kivu region, DR Congo.

Taxonomy Thomas (1916b) described kivu as a subspecies of D. insignis and hence a member of the mesomelas species group, where it was also placed by Bohmann (1942). Musser & Carleton (1993) considered kivu to be a distinct species, morphologically similar to D. mesomelas. Referring to Osgood’s (1936) two samples of Dendromus from Rwenzori, Musser & Carleton (1993) identified them as D. lunaris and D. insignis. Dendromus lunaris was found to be identical with (the pre-dated) D. kivu.The same sympatric combination – D. kivu and D. insignis – were found in two other series from the Kivu region. Dieterlen (1971, 1976a) recorded three species living sympatrically

in the region west of L. Kivu, and named them as D. mesomelas kivu (now D. insignis), D. melanotis (now D. kivu) and D. mystacalis. Musser & Carleton (2005) included kivu as a synonym of nyasae. This profile refers to what the present author considers as D. kivu from the Kivu region. The relationship between kivu and nyasae requires further investigation. Morphologically, D. nyasae seems to be most closely related to D. vernayi from Angola, D. oreas from the mountains of E Cameroon, and D. insignis from the mountains of East Africa (Musser & Carleton 2005). Synonyms: hintoni, kivu, lunaris. Subspecies: none. Chromosome number: not known. 179

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Family NESOMYIDAE

Description Small climbing mouse (although one of the largest species of Dendromus) with a long mid-dorsal stripe and a long tail. Pelage colouration varies with age (see below). Adult dorsal pelage typically rich brown, tending to be darker on anterior part of body; hairs ash-grey at base, brown at tips. Pelage may appear greyish-brown when basal hair colour visible on surface. Dorsal stripe black, 2–3 mm wide from ca. 2 cm behind head to base of tail. Flanks yellowishbrown. Ventral pelage silvery-white; hairs grey at base, silvery-white at tip. Chin and throat with pure white patches. Eyes relatively large. Hindfoot with short hair, silver-grey, rarely with reddish tinge. Hindfoot with five digits; Digit 1 short, Digit 5 long and opposable with short claw.Tail very long (ca. 124% of HB), colour and hairiness very variable. Ventral pelage in young and subadults (when several weeks or months in age) may be whitish or greyish. Nipples: 2 +2 = 8. Geographic Variation None recorded. Similar Species D. insignis. HB on average slightly larger; tail slightly longer (mean 97 mm) but of similar relative length; mid-dorsal stripe broader (ca. 4 mm) from back of head to base of tail. D. mystacalis. HB smaller; tail on average shorter (mean 84.8, 90.4 mm) but relatively longer (ca. 140–160% of HB); middorsal stripe from shoulders to base of tail. Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Recorded from the western and eastern slopes of the Rwenzori Mts and the Kivu region of E DR Congo, from 1300–4200 m (see below). May be considered as an ‘Albertine Rift Valley’ endemic. Other East African montane forms, similar or identical to D. kivu, are known from Tanzania, Rwanda, Kenya, but their taxonomic status is uncertain. Musser & Carleton (2005) give the distribution (as D. nyasae) as follows: Western Rift Mts and Rwenzori Mts (E DR Congo,W Uganda), south

through E DR Congo to the Marungu Mts in SE DR Congo, Mbizi Mts in W Tanzania, Rungwe and Ukinga in SW Tanzania, Nyika Plateau in NE Zambia and N Malawi, and highlands in S Malawi; also through the Eastern Arc Mts in E Tanzania (Uzungwa Mts, Uluguru highlands, Ukaguru Mts, Nguru Mts and South Pare Mts). Habitat Occurs over a wide range of habitats along the Albertine Rift Valley, and at altitudes of 1300–4200 m on the Rwenzori Mts (Verschuren et al. 1983) and 1600–2400 m to the west of L. Kivu (Dieterlen 1971, 1976a). All habitats are characterized by moist dense vegetation suitable for climbing. Favoured moist habitats at lower altitudes include grassy vegetation between cultivated fields, banana plantations and the edges of swamps. Above 2000 m occurs in sparse tree and/or bamboo forests and on the edges of swamps. At 4200 m (L. Marion, Rwenzori Mts), found in the afroalpine vegetation between immortelles (strawflowers) and alchemillas (Dieterlen 1976a, Verschuren et al. 1983). Abundance In favoured habitats, e.g. at the edge of grassy thickets, populations may be abundant and comprise up to 10% of individuals in a community of 10–15 small mammal species (Dieterlen 1967a, b, 1971). Adaptations Mostly nocturnal and arboreal. Agile climbers well adapted for life on stems and twigs. The large opposable Digit 5 of the hindfeet and the long semi-prehensile tail are used for holding onto small narrow stems. Ball-shaped nests, made of grass and leaves, are fastened between stalks and twigs some 30–80 cm above the ground (Dieterlen 1971). Foraging and Food Omnivorous or insectivorous. Five stomachs contained a mixture of seeds and the remains of insects, and two other stomachs contained only insects (Dieterlen 1971, 1976a). Social and Reproductive Behaviour No information, but it is presumed that individuals are mostly or entirely solitary. Reproduction and Population Structure Near L. Kivu, reproduction occurs throughout most of the year. Most births have been recorded during the wet season (Sep–May) with a peak of reproductive activity in Jan–Mar (73% of the !! active, n = 15). In the dry season, only one (11%) out of nine !! was reproductively active (Dieterlen 1971). Gestation: probably 23–27 days. Mean littersize: 4.Young are altricial at birth, unable to crawl, and the eyes and ears are closed. First hairs visible and dorsal stripe showing as pigmented skin Day 11. Incisors erupt Day 11. Molar teeth erupt Day 15–20. Eyes open, walking and climbing begins Day 22–24. Dark juvenile pelage complete (young much darker than adults) Day 24. Weaning begins Day 24.Young totally weaned and appearance like small adults (although HB not yet adult size) Day 35. Sex ratio: 69% : 31% (ca. 7 : 3; n = 83; Dieterlen 1967a, b, 1971; see also D. insignis). Predators, Parasites and Diseases Small carnivores, birds and snakes are potential predators. The small intestine of one adult contained many parasitic worms, some 50 mm long.

Dendromus nyasae

Conservation

IUCN Category: Least Concern.

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Dendromus nyikae

M1–M3: 3.4 (3.1–3.6) mm, n = 28 Lwiro and Kahuzi region, DR Congo (SMNS, Dieterlen 1971, 1976, F. Dieterlen unpubl.)

Measurements Dendromus nyasae (kivu) HB: 73.8 (64–80) mm, n = 40 T: 91.8 (84–105) mm, n = 38 HF: 18.9 (17.0–21.5) mm, n = 40 E: 13.6 (11.0–15.5) mm, n = 40 WT: 10.5 (6.0–20.0) g, n = 40 GLS: 22.2 (20.7–23.2) mm, n = 32 GWS: 10.9 (10.1–11.5) mm, n = 16

Key References Bohmann 1942; Dieterlen 1971, 1976a; Musser & Carleton 1993; Osgood 1936. Fritz Dieterlen

Dendromus nyikae NYIKA AFRICAN CLIMBING MOUSE Fr. Souris arboricole de Nyika; Ger. Nyika-Klettermaus Dendromus nyikae Wroughton, 1909. Ann. Mag. Nat. Hist., ser. 8, 3: 248. Nyika Plateau, Malawi.

Taxonomy Closely related to D. melanotis. Sometimes placed (with D. melanotis) in the subgenus Poemys, which is characterized by the flattened nail on the Digit 5 of hindfoot (Crawford-Cabral 1998). Dendromus nyikae angolensis is considered to be a synonym of D. pecilei in Angola by Crawford-Cabral (1998) but pecilei is treated as a synonym of D. melanotis by Musser & Carleton (2005). Synonyms: angolensis, bernardi, longicaudatus. Subspecies: none. Chromosome number: not known. Description A very small climbing mouse with a mid-dorsal thin black stripe, and very long tail. Dorsal pelage pale to dark cinnamon suffused with grey. Dorsal hairs short (6–7 mm), soft and dense; grey at base, cinnamon at tip. Black mid-dorsal stripe from shoulders to base of tail, sometimes rather inconspicuous and irregular; hairs of stripe grey with dark cinnamon or black tip. Ventral pelage usually pure creamy-white; hairs in some individuals medium grey at base, with small white tip. Throat and chest yellow or rufous in some individuals (?""; ?glandular secretions). Head cinnamon, without black patch on crown. Small white subauricular patch. Ears rounded, conspicuous (standing out from side of head), brown with short brown hairs. Hindfoot with five digits; Digit 1 short with nail, Digit 5 very long and opposable with nail (see also D. melanotis). Tail long (ca. 130–160% HB), scaly with small pale bristles; longer hairs at tip. Nipples: not known. Geographic Variation None recorded. Similar Species D. melanotis. Similar in size; well-defined wide dorsal stripe; flat nail on Digit 5 of hindfoot. D. mystacalis. On average smaller; black dorsal stripe; ventral pelage pure white; small claw on Digit 5 of hindfoot. D. mesomelas. On average larger HB, T and HF; dorsal stripe dark but variable; small claw on Digit 5 of hindfoot. Distribution Endemic to Africa. Zambezian Woodland and Afromontane–Afroalpine BZs. Recorded from Angola, S DR Congo, Zambia, Zimbabwe, Malawi, SW Tanzania, Mozambique and N South Africa. A specimen of D. melanotis nyikae from Ukerewe I. in L. Victoria is, in fact, D. mystacalis (Musser & Carleton 2005).

Dendromus nyikae

Habitat Long grass in grassland savannas and plateaux, mostly at higher altitudes. May occur in grassland habitats within pine plantations. On Nyika Plateau, Malawi, some individuals found in unoccupied standard beehives (specimens; HZM). Does not enter forested habitats except very rarely. Abundance Rather uncommon, although may be locally abundant. In Malawi, ‘widespread but probably limited to upland areas’ (Ansell & Dowsett 1988). Comprised 9.2% of grassland small rodents during a mark-capture study (n = 215) on Zomba Plateau, Malawi (1800 m) (Happold & Happold 1989c), but not present in tall Hyparrhenia and Panicum grasslands in Liwonde N. P., Malawi (500 m) (Happold & Happold 1990). In Malawi, numbers were highest where the montane grasslands were tall (i.e. 40– 80 cm, depending on time of year and location), and lowest (or absent) where grasslands were short and after burning (Happold & Happold 1989c). 181

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Family NESOMYIDAE

Adaptations Nyika Climbing Mice are adapted for climbing on thin tall grass stems. Their small size and grasping feet allow them to climb up and down grass stems from the flowerheads to ground level. When climbing, the hindfeet are held almost at right angles to the body. A grass stem is grasped by the forefeet as would a human hand, and the flexible Digits 1 and 5 of the hindfeet encircle the stem. The tail is stretched out from the rump at various angles to provide balance, or the tip is wound several times around a stem to provide an extra holding point. Locomotion is by swiftly climbing up and down stems, and reaching across gaps (equivalent to at least the length of head and body) to adjacent stems (D. C. D. Happold unpubl.). The burrow is a simple tunnel in the soil with a nest chamber lined with dead leaves (Hanney 1965). Foraging and Food Seed-heads of grasses are obtained by climbing to the top of a stem, biting through part of the stem below the seedhead so that the head falls over but is still attached, and then nibbling at the seeds (D. C. D. Happold pers. obs.). Analysis of stomach contents (n = 6) revealed five with white vegetable material, one with grain husks and three with insects, including beetles (Hanney 1965). Social and Reproductive Behaviour No information. Nyika Climbing Mice make a high-pitched squeal when disturbed.

Predators, Parasites and Diseases Comprised 6% (n = 100 prey) and 30% (n = 46 prey) of prey numbers in the pellets of African Grass-owls Tyto capensis on Zomba Plateau, Malawi. When foraging on the tops of grass stems, they are probably easy prey for owls. Because of their small size, climbing mice comprised only 1% and 6% of prey biomass (Happold & Happold 1986). Conservation

IUCN Category: Least Concern.

Measurements Dendromus nyikae HB: 66.8 (50–78) mm, n = 24 T: 86.4 (80–93) mm, n = 21 HF: 16.7 (11–19) mm, n = 24 E: 14 (11–18) mm, n = 21 WT: 10.5 (11–18) g, n = 15 GLS: 21.8 (20.3–24. 4) mm, n = 14 GWS: 11.1 (10.0–11. 7) mm, n = 8 M1–M3: 3.5 (3.2–3.6) mm, n = 8 Malawi (BMNH; D. C. D. Happold unpubl.) Key References

Hanney 1965; Happold & Happold 1986. D. C. D. Happold

Reproduction and Population Structure Pregnant ! in Nov (Malawi; Hanney 1965); lactating ! in Aug (S Tanzania; Kingdon 1974). Embryos: 4 (n = 1, Hanney 1965).

Dendromus oreas CAMEROON AFRICAN CLIMBING MOUSE Fr. Souris arboricole du Cameroon; Ger. Kamerun-Klettermaus Dendromus oreas Osgood, 1936. Field Mus. Nat. Hist., Zool. Ser. 20: 236. South-west side of Mount Cameroon, Nigeria (now in Cameroon). 9000 ft (2740 m).

Taxonomy Described as a distinct species, but subsequently considered to be a synonym of D. mesomelas (Bohmann 1942, Rosevear 1969, Misonne 1974). Referred to as D. mesomelas oreas by Rosevear (1969). Musser & Carleton (1993, 2005) reinstated oreas as a valid species, and considered it to be related to Dendromus lunaris (now D. nyasae [kivu]), and not to be a geographic outlier of D. insignis. Synonyms: none. Chromosome number: not known. Description Very small climbing mouse with an indistinct middorsal stripe and long tail. Dorsal pelage medium brown. Black middorsal stripe from mid-back to base of tail, often rather indistinct. Ventral pelage varied, from dark rufous to pale greyish-yellow; hairs dark at base. White or cream patches on chin and throat, and around anus. Ears blackish-brown, with fine covering of blackish-tawny hairs; pale spot at base of outer margin of each ear, sometimes indistinct. Hindfoot with five digits; Digit 5 long and opposable with claw. Tail very long (ca. 135% of HB), dusky above, paler below. Nipples: not known.

Similar Species D. messorius. On average smaller; mid-dorsal stripe absent; usually occurs at lower altitudes; recorded from Cameroon (and many other countries) but not from mountainous habitats (see below). Distribution Endemic to Africa. Afromontane–Afroalpine BZ; known only from Mt Cameroon (1700–4000 m), Mt Manenguba (1800–1900 m, ca. 120 km NE of Mt Cameroon) and Mt Kupé (ca. 80 km NE of Mt Cameroon, 850 m) in Cameroon (Eisentraut 1963, Rosevear 1969). Probably endemic to the mountains of W Cameroon. This is the only known species of Dendromus on the mountains of Cameroon. Habitat Montane savannas above the montane forest zone, from ca. 1675 m to ca. 2900 m, on the ground amongst boulders of lava (Mt Cameroon). Also recorded on dry grassy and scrubby slopes (Mt Manenguba), and in plantations and farmlands (Mt Kupé). Has not been recorded in montane forest habitats at lower altitudes on these mountains.

Geographic Variation None recorded.

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Dendromus vernayi

Adaptations Mostly terrestrial, diurnal and nocturnal. Although well adapted for climbing, these mice spend most of their life on the ground and in their subterranean burrows. Reproduction One ! with five embryos in late Nov (Eisentraut 1963). Conservation IUCN Category: Vulnerable. Geographic range is very limited and populations on each of the mountains are isolated. Measurements Dendromus oreas HB: 69.3 (60–74) mm, n = 6 T: 95.3 (89–104) mm, n = 7 HF: 19.0 (18–20.5) mm, n = 7 E: 15.3 mm (13–19) mm, n = 6 WT: 11.1 (9–13) g, n = 5 GLS: 22.0 (21.6–22.5) mm, n = 4 GWS: 11.0 (10.9–11.3) mm, n = 4 M1–M3: 3.5 (3.3–3.7) mm, n = 7 Cameroon (Osgood 1936, Eisentraut 1963, Rosevear 1969, F. Dieterlen unpubl.) Key References

Dendromus oreas

Eisentraut 1963; Rosevear 1969. Fritz Dieterlen

Dendromus vernayi VERNAY’S AFRICAN CLIMBING MOUSE Fr. Souris arboricole de Vernay; Ger. Vernays Klettermaus Dendromus vernayi Hill and Carter, 1937. Amer. Mus. Novit. 913: 4. Chitau, Angola. 4930 ft (1500 m).

Taxonomy Described as a subspecies of Dendromus mesomelas (see also Misonne 1974). Dendromus vernayi is morphologically distinct from D. nyasae and D. oreas, and is probably phylogenetically more closely related to these species than to any other species of Dendromus (Musser & Carleton 2005). Synonyms: none. Chromosome number: not known. Description Very small climbing mouse with dark mid-dorsal stripe. Pelage relatively long and silky. Dorsal pelage ochraceous and cinnamon-brown; hairs dark grey at base. Mid-dorsal stripe wide (ca. 4.5 mm). Ventral pelage varied, bright pinkish-buff or buffy-grey; hairs medium grey at base. Pure white patch on throat and in axillary region. Ears blackish on outer surface, short orange hairs on inner surface. Upper surface of fore- and hindfeet pinkish-buff. Hindfoot relatively long (>19 mm). (No information on digits of fore- and hindfoot.) Tail very long (ca. 130% of HB), dark above, paler below. Skull: rostrum heavier and shorter than in other species of Dendromus. Nipples: not known. Geographic Variation None recorded.

Similar Species D. melanotis. Body size slightly larger (mean HB: 68 mm); skull (GLS, GWS) slightly smaller; tail usually longer; hindfoot shorter (usually 8 (Coetzee 1977). Geographic Variation None recorded. Similar Species S. cuppedius. Smaller size, pelage pale, hairs soft; tail relatively longer (51–57% of HB). S. jacksoni. Same size as the largest S. caurinus, pelage dark brown, hairs soft; tail usually relatively longer (ca. 50% of HB); length of interparietal bone ca. 4.5 mm. Distribution Endemic to Africa. Sudan and Guinea Savanna BZs. Recorded from Senegal, Mali, Burkina, N Côte d’Ivoire, N Ghana, Togo, Benin, NW Nigeria; eastern limits uncertain (Swanepoel & Schlitter 1978; B. Sicard & L. Granjon unpubl.).

Steatomys caurinus

Habitat Drier savannas on sandy soils, where vegetation cover is poor. Often found in traditionally cultivated fields and fallow lands. Abundance Apparently uncommon, perhaps due to low trappability; specimens obtainable only through excavations of burrows. Infrequently found in pellets of Barn Owls Tyto alba in Senegal (L. Granjon, K. Ba & J.-M. Duplantier unpubl.). 193

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Family NESOMYIDAE

Adaptations Nocturnal and terrestrial. Individuals accumulate fat during the wet season, which allows them to go into torpor and to be inactive during the dry season. In laboratory conditions, appears better adapted to dehydration than gerbils (Lacas et al. 2000). Foraging and Food No information. Social and Reproductive Behaviour adult individual per burrow.

Generally only one

Reproduction and Population Structure Females with their young recorded in 11 burrows in Sep and Oct in S Mali; in each burrow, mean number of young 8.5 (range 5–12, n = 11) (B. Sicard & L. Granjon unpubl.). Predators. Parasites and Diseases Found in pellets of Barn Owls Tyto alba in Senegal (Heim de Balsac 1965; Granjon, Bâ & Duplantier unpubl.).

Conservation

IUCN Category: Least Concern.

Measurements Steatomys caurinus HB: 104.7 (79–122) mm, n = 9 T: 39.3 (18–51) mm, n = 9 HF: 16.5 (14.5–18) mm, n = 9 E: 15.9 (14–18) mm, n = 9 WT: 44.6 (16.5–101) g, n = 9 GLS: 24.3 (19.5–27.3) mm, n = 8 GWS: 12.9 (11.9–13.8) mm, n = 9 M1–M3: 3.9 (3.5–4.6) mm, n = 9 Senegal and Mali (B. Sicard & L. Granjon unpubl.) Key References Schlitter 1978.

Coetzee 1977; Rosevear 1969; Swanepoel & B. Sicard & J.-M. Duplantier

Steatomys cuppedius DAINTY FAT MOUSE Fr. Souris adipeux gracile; Ger. Zierliche Fettmaus Steatomys cuppedius Thomas and Hinton, 1920. Novit. Zool. 27: 318. Farniso near Kano, Nigeria. 1700 ft (518 m).

Taxonomy Although listed as a subspecies of S. parvus by Coetzee (1977), most authors (e.g. Rosevear 1969, Swanepoel & Schlitter 1978, Musser & Carleton 1993, 2005) consider cuppedius to be a valid species. Synonyms: none. Chromosome number: not known.

pelage pure white. Nose pointed. Ears large and rounded. Nasal region, cheeks, lips and throat white. Forefeet and hindfeet white. Tail short (57% of HB, range: 51–67%), white, scantily covered with white hairs above and below. Nipples: 2 + 2 = 8.

Description Small, delicate mouse with soft silky pelage and short white tail; the smallest of the West African species. Dorsal pelage pale, sandy to grey; hairs medium grey at base, sandy or grey at tip; darker on mid-dorsal line due to some black-tipped hairs. Ventral

Geographic Variation

None recorded.

Similar Species S. caurinus. Larger size, pelage darker, hairs coarse; tail relatively short (mean 38% of HB). Distribution Endemic to Africa. Sudan Savanna BZ. Recorded from several localities in Senegal, and also from N Nigeria and S Niger. Swanepoel & Schlitter (1978) expressed doubts about data concerning Senegal (Heim de Balsac 1965) but captures (in Mbour) and in owl pellets from various localities (L. Granjon, K. Ba & J.-M. Duplantier unpubl.) confirm the presence of the species in Senegal. May occur also in other parts of the Sudan Savanna BZ between these known localities. Records of this species from SE Ghana are now known to be Uranomys ruddi (see Grubb et al. 1998 for details). Habitat In Senegal, trapped in sandy areas near gardens and near the coast. Seems to prefer drier habitats than S. caurinus. Abundance Generally uncommon, but may be locally abundant: e.g. in Nigeria, 55 individuals caught within a week in a single locality (as S. parvus; Happold 1987), and in Senegal 12 individuals caught within two weeks on the same trap line. Comprised up to 30% of prey in pellets of Barn Owls Tyto alba in some localities in Senegal (L. Granjon, K. Ba & J.-M. Duplantier unpubl.).

Steatomys cuppedius

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Steatomys jacksoni

Adaptations Nocturnal and terrestrial. Individuals accumulate fat during the wet season, which allows them to go into torpor or to be inactive during the dry season. Foraging and Food No information. Social and Reproductive Behaviour Several individuals caught at the same place suggests the existence of colonies. Reproduction and Population Structure No detailed information. Juveniles (HB: 58–62 mm) in Dec in N Nigeria (labels, BMNH). In Mbour (Senegal), sex-ratio in Aug was 10 "" : 2 !! for individuals caught at the same place within a few days. Predators, Parasites and Diseases Preyed on by Barn Owls Tyto alba in Senegal (Heim de Balsac 1965; L. Granjon, K. Ba & J.-M. Duplantier unpubl.).

Conservation

IUCN Category: Least Concern.

Measurements Steatomys cuppedius TL: 132.7 (120–143) mm, n = 6 T: 48.7 (46–54) mm, n = 6 HF: 16.0 (15–17) mm, n = 6 E: 13.7 (13–14) mm, n = 6 WT: 16.2 (13–22) g, n = 6 GLS: 22.4 (21.3–23.9) mm, n = 5 GWS: 11.8 (11.2–12.4) mm, n = 6 M1–M3: 3.8 (3.6–4.0) mm, n = 10 N Nigeria (Swanepoel & Schlitter 1978) Key References

Rosevear 1969; Swanepoel & Schlitter 1978. J.-M. Duplantier & B. Sicard

Steatomys jacksoni JACKSON’S FAT MOUSE Fr. Souris adipeux de Jackson; Ger. Jacksons Fettmaus Steatomys jacksoni Hayman, 1936. Proc. Zool. Soc. Lond. 1935: 930 (publ. 1936). Wenchi, Ghana.

Taxonomy Included in S. caurinus by Coetzee (1977) but retained as a distinct species by Swanepoel & Schlitter (1978) and Musser & Carleton (1993).The wide interparietal bone on the skull (compared with other species in the genus) is the diagnostic character for the species, but the significance of this character is uncertain.The validity of the species can not be assessed until additional specimens are available (Musser & Carleton 1993). Synonyms: none. Chromosome number: not known. Description Small dark brown mouse, with a short tail and tendency to be fat (or very fat) at some seasons of the year. Pelage soft. Dorsal pelage dark brown; dorsal hairs dark slate-grey at base, medium-brown at tip. Ventral pelage pure white. Chin, throat, chest white. White postauricular spot, sometimes obscure. Fore- and hindlimbs and feet white. Forefoot with four well-developed digits. Tail short (ca. 50% of HB), bicoloured, dark above, whitish below. Skull with small cheekteeth (and M3 very small), palate not much prolonged posteriorly to cheekteeth, each upper incisor opisthodont with single groove and angled slightly outwards, and moderately well-developed auditory bullae; interparietal bone ca. 4.5 mm (cf. S. caurinus). Nipples: 2 + 4 = 12. Steatomys jacksoni

Geographic Variation None recorded. Similar Species S. caurinus. Dorsal pelage rufous-brown and paler; usually smaller, HB: 79–122 mm, HF: 14–18 mm, GLS: 19–27 mm; interparietal bone 3.0–3.5 mm; Guinea and Sudan Savanna BZs; may be parapatric with this species. S. cuppedius. Dorsal pelage sandy-grey; smaller, HB: 3800 m, near summit), diet is snails, worms andTipulidae larvae (V. Clausnitzer unpubl.). It appears that this species takes less animal food than most other species of the genus (Dieterlen 1976b) and that the respective portions of animal and plant foods depend on environment and season. Social and Reproductive Behaviour Little information available. Many individuals in the wild have torn ears and mutilated tails suggesting agonistic behaviour. In captivity, individuals (especially "") fight and injure each other and deaths (with or without physical damage) are not uncommon. In captivity, individuals are very shy, hiding themselves in vegetation and nesting materials. They are able to utter a feeble cry (Hanney 1964; Delany 1972; Dieterlen 1976b; Muhmenthaler 1999). Reproduction and Population Structure In most studies, reproduction occurs during the wet season. In Malawi, pregnancies were recorded during the late dry season, throughout the wet season, and at the beginning of the dry season (Oct–May), with the highest pregnancy rates in Nov–Mar (Hanney 1964; Happold & Happold 1989c). In Uganda, two peaks of pregnancy (Sep–Nov and Mar–Jun) coincided with the double peak of annual rainfall (Delany 1971, 1972). In Kivu, DR Congo, three populations living in different habitats and different rainfall regimes showed different patterns of reproductive activity (Table 21). Reproduction shows considerable flexibility; shorter periods of reproduction are characterized by a lower pregnancy rate and larger litters, and longer periods of reproduction by a higher pregnancy rate and smaller litters. In Uganda, most pregnant !! are primiparous although a small number of older !! may be pregnant with their second to fifth litters (Delany 1971). In Uganda, some "" are fecund at all times of the year, with a higher fecundity rate when monthly rainfall is at its highest (Delany 1971). In Malawi, testis size in adult "" fluctuates during the year, being smallest in the dry season (when perhaps "" are not fecund) and largest in the wet season when the fecundity rate is also at its highest (Hanney 1964). Litter-size varies depending on the environment (see above) and also on the size of the !. In Uganda, mean litter-size is 1.9 for !! of less than 40 g, 2.2 for !! of 40–49 g, and 2.3 for !! over 50 g (Delany 1971). In Malawi, litter-size is 2.4 (2–5); 51% of !! had 3 embryos and 25% had 2 embryos (total n = 43, Hanney 1964). Gestation: 30–31 days. At birth young are naked, with eyes and ears closed, but are rather large (6.5–9.5 g). Growth is rapid: thin

Table 21. Patterns of reproductive activity in populations of Lophuromys flavopunctatus living in different habitats with different rainfall regimes.

a

Habitat and altitude

Rainfall (months)

Number of months of pregnancies

Pregnancy ratea

Litter-size mean (range)

Cultivations, 1650–1850 m Montane forest, 2000– 2300 m Lowland rainforest, 800–900 m

9

9

42%

2.18 (1–4)

10

10

57%

1.99 (1–4)

12

87%

1.83 (1–2)

9 (but mostly all year)

Pregnancy rate = % of adult !! (40 g or more) which were pregnant, only during months when pregnancies recorded.

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Family MURIDAE

covering of dorsal hair by Day 3, and well covered with hairs by Day 3–6. Eyes open by Day 4–7. Walking by Day 5, and looking like a small adult by Day 9. Adult pelage by Day 30–35. Sexually mature by Day 50–70 when HB: 110–120 mm and weight ca. 45 g (Hanney 1964, Delany 1971, 1975, Dieterlen 1976b). Sex ratio is fairly constant but varies according to time of year and locality. In Kivu, DR Congo, "" formed 53.5% of the population at the beginning of the breeding season. In different localities, "" comprised 51.0% to 55.9% of the population (Dieterlen 1976b, 1985a). Young are born during many months of the year, and hence populations include young, subadults and adults (of various ages) in most months of the year. In Uganda (two wet seasons and two main periods of pregnancy), each of six age-classes are represented in each month with a tendency for larger numbers of younger animals in Oct–Jan and Apr–Jun (Delany 1971). In Malawi (one wet season and one period of pregnancy), the largest number of young (WT less than 40 g) occurs during and immediately after the wet season (Jan–Jun). These young are responsible for population numbers being at their highest at the time. Mortality of older individuals cause population numbers to be at their lowest in the late dry season (Sep– Dec) (Happold & Happold 1989c). The large number of !! that have only one litter (compared with the small number that have two or more litters) suggests that there is high mortality and rapid turnover of the population (Delany 1971). Predators, Parasites and Diseases The strong odour of this species makes them unattractive and unpalatable for some carnivores (see above). However, they are preyed upon by several diurnal and nocturnal birds such as eagles, herons (Misonne 1963) and Barn Owls Tyto alba (Rahm 1960b). Remains of L. flavopunctatus were not found in the pellets of African Grass-owls Tyto capensis in montane grasslands in Malawi, even though the species was common (Happold & Happold

1986). They are eaten by snakes of the genera Naja, Dendroaspis and Bitis (Allen & Loveridge 1942). Hanney (1964) reported that about 9% of individuals were diseased in Malawi; the percentage fluctuated throughout the year, being lowest during the cold dry season (3–7%) and highest (up to 15%) during the wet season. The most prevalent endoparasites were diphylobothriid larvae (Cestoda), which infested about 6% of individuals. Ectoparasites include five species of fleas (Ctenophalmus eximius, C. nyikensis, C. calceatus, Dinopsyllus sp. and Xiphiopsylla hyparetes) (Hanney 1964). Conservation IUCN Category: Least Concern. This common and widely distributed species is not threatened at present, although clearance of suitable habitats will reduce numbers and limit the geographic range. Measurements Lophuromys flavopunctatus HB: 119 (98–144) mm, n = 113 T: 63 (46–88) mm, n = 113 HF: 21.3 (20–25) mm, n = 113 E: 17.4 (13–20) mm, n = 113 WT: 52 (36–73) g, n = 102 GLS: 30.2 (26.8–31.3) mm, n = 56 GWS: 14.9 (12.9–15.9) mm, n = 56 M1–M3: 4.8 (3.9–5.4) mm, n = 56 Body measurements and weight: throughout most of geographic range Skull measurements: E DR Congo (Dieterlen 1976b, SMNS) Key References Delany 1971; Dieterlen 1976b; Hanney 1964; Happold & Happold 1989c. Fritz Dieterlen

Lophuromys huttereri HUTTERER’S BRUSH-FURRED RAT Fr. Souris hérissé de Hutterer; Ger. Hutterers Bürstenhaarmaus Lophuromys huttereri Verheyen, Colyn & Hulselmans 1996. Bull. Inst. Roy. Sci. Nat. Belgique, Biol. 66: 255. Yaenero, DR Congo (00° 12´ N. 24° 47´ E).

Taxonomy Subgenus Lophuromys. Species-group: sikapusi. An ‘unspeckled’ short-tailed Lophuromys, more closely related to L. nudicaudus than to the other representatives of the sikapusi speciesgroup (W. Verheyen et al. 1996). The characteristics that distinguish this species from other Lophuromys are based mainly on the skull and teeth. Synonyms: none. Chromosome numbers: not known. Description Similar in external characters to L. nudicaudus. Compared to L. nudicaudus, the skull has a wider and higher rostrum, the zygomatic plate is situated more anteriorly, and t3 on M2 is nearly always totally absent. Can only be distinguished unambiguously from L. nudicaudus by discriminate analysis (see W. Verheyen et al. 1996).

Similar Species L. nudicaudus. Probably similar in size; rostrum narrower and lower; zygomatic plate narrow. L. sikapusi. Zygomatic plate comparatively broad (mean 2.82 mm). Distribution Endemic to Africa. Rainforest BZ (South Central Region). Recorded from between the Lualaba and Lomani rivers close to junction with the Congo R. (the type locality), and west of the Lomani R. to Ndele (00° 51´ N, 21° 05´ E); all localities on the left side of the Congo R. (cf. L. nudicaudus).The single specimen from Ndele seems to be closer to typical L. nudicaudus than to L. huttereri. Habitat Rainforest.

Geographic Variation None recorded. Abundance

Rare; known only from about three localities.

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Lophuromys luteogaster

Remarks Apparently no other information available. Conservation IUCN Category: Least Concern. A rare and little-known species. Measurements Lophuromys huttereri HB: 93–114 mm T: 59–61 mm HF: 18–20 mm E: n. d. WT: n. d GLS: 28.8 (28.0–29.9) mm GWS: 14.4 (13.7–15.2) mm M1–M3: 4.4 (4.1–4.8) mm Skull measurements: n = 7–10 DR Congo (W. Verheyen et al. 1996, BMNH, RUCA) Key Reference

W. Verheyen et al. 1996. Fritz Dieterlen Lophuromys huttereri

Lophuromys luteogaster BUFF-BELLIED BRUSH-FURRED RAT (HATT’S BRUSH-FURRED RAT) Fr. Souris hérisse à ventre fauve; Ger. Gelbbauch Bürstenhaarmaus Lophuromys luteogaster Hatt, 1934. Amer. Mus. Novitat. 708: 4. Medje, Ituri district, DR Congo.

Taxonomy Subgenus Kivumys. Species-group: woosnami. Synonyms: none. Chromosome number: not known. Description Small unspeckled olive-brown rat with long tail, similar to L. woosnami. Pelage rather harsh. Dorsal pelage uniformly olive-brown and unspeckled, hairs reddish or yellowish at base. Ventral pelage pinkish-cinnamon to buff, hairs unicoloured. Dorsal colour merges to ventral colour on flanks. Head similar colour to dorsal pelage. Ears darkly pigmented with short greyish bristles. Forelimbs yellowish to reddish, hindlimbs pinkish-cinnamon, claws short, relatively weak, pale. Tail relatively long (ca. 100% of HB), dark above, pale below, except darkish tip. (The colour of dry skins in museums fades and looses its brightness when exposed to light). Skull: relatively long; M1 relatively broad. Nipples: 2 + 1 = 6. Geographic Variation None recorded. Similar Species L. woosnami. Similar colour; mostly larger in all body measurements; montane habitats of E DR Congo. L. flavopunctatus. Dorsal pelage reddish-brown; on average larger HB and skull; tail shorter and relatively shorter; sympatric. Distribution Endemic to Africa. Rainforest BZ (East Central Region). Recorded from only four localities in rainforest in NE and E DR Congo.

Lophuromys luteogaster

Habitat Lowland rainforests from 700 to 1100 m where dominant trees are either Gilbertiodendron dewevrei or Julbernardia and Cynometra (Hatt 1934, Dieterlen 1975, 1976b, Schlitter & Robbins 1977, Verschuren et al. 1983). Also occasionally in sub-montane forests of Julbernardia at about 1400 m (F. Dieterlen unpubl.). Of 26 records from 249

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Family MURIDAE

lowland rainforest, 11 were from primary forest, 14 from secondary forest and one from a cultivated area (Dieterlen 1975, 1976b).

Although a rare species, it is unlikely to be endangered because it lives in several habitats within the extensive rainforest.

Abundance Rare, comprising ca. 0.85% of terrestrial small mammals captured in the region of Irangi (DR Congo). Despite regular collecting during several years, almost 90% of captures (n = 34) were during the last three months of dry season and first three months of the wet season (Jun–Nov).

Measurements Lophuromys luteogaster HB: 102 (90–113) mm, n = 25 T: 106 (90–117) mm, n = 25 HF: 21 (19–22.5) mm, n = 25 E: 17.5 (16.5–19) mm, n = 25 WT: 34 (28–41) g, n = 25 GLS: 28.8 (27.2–29.6) mm, n = 12 GWS: 13.2 (12.9–13.9) mm, n = 12 M1–M3: 4.5 (4.2–4.9) mm, n = 12 E DR Congo (Dieterlen 1976b, SMNS)

Remarks Terrestrial. Daily activity pattern not well known; mostly captured at night. Mostly insectivorous. Stomach analysis (n = 9) shows that the diet is mainly (90–100%) small grubs, caterpillars, small snails, small beetles and termites. Pregnancies recorded in Jun–Nov (no data at other times of year). During these months, pregnancy rate was high (11 out of 14). Embryo numbers: 2 (n = 11). Sex ratio 12 "" : 21 !!. Sexually active "" had testes length of at least 6 mm. Conservation

Key References

Dieterlen 1975, 1976b, 1987; Hatt 1934. Fritz Dieterlen

IUCN Category: Least Concern.

Lophuromys medicaudatus MEDIUM-TAILED BRUSH-FURRED RAT (WESTERN RIFT BRUSH-FURRED RAT) Fr. Souris hérissé a queue moyenne; Ger. Mittelschwänzige Bürstenhaarmaus Lophuromys medicaudatus Dieterlen, 1975. Bonn. Zool. Beitr. 26: 295. Nyabutera near Lemera, Kivu, DR Congo.

Taxonomy Subgenus Kivumys. Species-group: woosnami. The only species of Lophuromys with a tail of 85% of HB. Closely related to L. woosnami and L. luteogaster. Synonyms: none. Chromosome number: not known. (Specimens named as L. luteogaster collected in the mountains west of L. Kivu by U. Rahm [Verheyen (1964c], and one specimen from Nyungwe Forest, Rwanda [Elbl et al. 1966] are, in fact, L. medicaudatus [see Dieterlen 1975]). Description Small beautiful rat with brightly coloured unspeckled pelage and medium tail length. Dorsal pelage and head uniform dark olive-brown, rather harsh; dorsal hairs paler at base, olive-brown at tip. Ventral pelage dark orange tinged with olive, colouration most intense on the chest; ventral hairs paler at base, olive at tip. Ears darkly pigmented, sparsely haired. Fore- and hindlimbs olive-brown. Tail long (ca. 85% of HB), darkish above, paler below. (The colour of dry skins in museums fades and loses its brightness when exposed to light.) Nipples: 2 + 1 = 6. Geographic Variation None recorded. Similar Species L. woosnami. Larger, ventral pelage pale brown; tail longer (mean 123 mm, 105% of HB); sympatric and syntopic. Distribution Endemic to Africa. Afromontane–Afroalpine BZ of the Albertine Rift Valley of E DR Congo and Rwanda around L. Kivu, at 1850–2500 m (Verheyen 1964c, Dieterlen 1975, 1976b, 1987, Verschuren et al. 1983). Specimens named as L. luteogaster collected in the mountains west of L. Kivu by U. Rahm (Verheyen 1964c), and one specimen from Nyungwe Forest, Rwanda (Elbl et al. 1966) are, in fact, L. medicaudatus (see Dieterlen 1975).

Lophuromys medicaudatus

Habitat Montane swamps of Cyperus latifolius and montane forests; syntopic with L. flavopunctatus and L. woosnami. Abundance Rare; comprised 2.3% (9 specimens) of small mammals in montane swamps and 0.6% (17 specimens) in montane forests (Dieterlen 1975, 1976b).

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Lophuromys melanonyx

Adaptations Terrestrial.There appear to be no special adaptations for locomotion in flooded swamp vegetation, but it is assumed that the ability to swim and climb must be essential for survival.

This species is threatened because of its rarity and very restricted geographic range; in this respect it is similar to the other Albertine Rift endemics, e.g. L. rahmi, L. woosnami and L. cinereus.

Foraging and Food Omnivorous. Stomach contents (n = 11) contained remains of many arthropods and some molluscs (70%, range 30–100%) as well as seeds and fruits.

Measurements Lophuromys medicaudatus HB: 103.2 (92–112) mm, n = 27 T: 87.2 (73–95) mm, n = 27 HF: 20.8 (18–23) mm, n = 27 E: 16.5 (15–19) mm, n = 27 WT: 35.4 (29–43) g, n = 27 GLS: 29.1 (27.7–30.2) mm, n = 19 GWS: 14.7 (14.1–15.6) mm, n = 19 M1–M3: 4.1 (3.9–4.3) mm, n = 19 DR Congo (Dieterlen 1975, SMNS)

Social and Reproductive Behaviour

No information.

Reproduction and Population Structure Mean embryo numbers: 1.6 (range 1–2, mode 2, n = 5 !!). Pregnant !! recorded in Feb, Apr and Jul. Testes length in mature "": 6–8 mm. Sex ratio 2 : 1 (n = 30). Predators, Parasites and Diseases No information.

Key References Conservation

Dieterlen 1975, 1976b, 1987.

IUCN Category: Vulnerable. Fritz Dieterlen

Lophuromys melanonyx BLACK-CLAWED BRUSH-FURRED RAT Fr. Souris hérissé d’Ethiopie; Ger. Schwarzkrallen-Bürstenhaarmaus Lophuromys melanonyx Petter, 1972. Mammalia 36: 177. Dinshu, Bale, Ethiopia.

Taxonomy Subgenus Lophuromys. Species-group: flavopunctatus. One of three Ethiopian endemic species of Lophuromys. A distinctive species, larger than any other species of Lophuromys. Synonyms: none. Chromosome number: 2n = 60 (Corti et al. 1995, Lavrenchenko et al. 1997). Description Medium-sized to large speckled grey-brown rat with large ears; the largest species of Lophuromys. Dorsal pelage greybrown, finely speckled with cream spots. Ventral pelage creamywhite. (Lacks the overall reddish-brown colouration of most of its congeners.) Ears prominent, grey, lightly furred with distinctive tuft of orange or cream at the base of ears. Feet whitish, with a grey wash dorsally, claws distinctively long and black. Tail short (ca. 43% of HB), dark above, whitish on the sides and below. Skull slightly larger than all congeners, with proportionately narrower interorbital region and strong zygomatic plate. Incisors strongly pro-odont (in contrast to moderately orthodont in all other Lophuromys). Skin tough (cf. fragile skin of other Lophuromys and Acomys spp.). Nipples: probably 0 + 2 = 4. Geographic Variation None recorded. Similar Species L. brevicaudus. On average much smaller size, pale claws; chromosome number 2n = 68. L. chrysopus and other species of the flavopunctatus species-group. Smaller, distinctive reddish colouration; chromosome number 2n = 54 (chrysopus) or 68 (Lavrenchenko et al. 1997). Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Recorded only from Ethiopia at 3200–4300 m, commonly east of the Rift Valley in Bale, but rarely west of the Rift Valley (only from near

Lophuromys melanonyx

Debra Sina and Addis Ababa). Range perhaps more extensive, but not reported from Simien despite extensive surveys (e.g. by Müller 1977). Habitat Afroalpine moorland, above the treeline (3500 m), ranging down to lower grasslands where open river valleys penetrate the woodland zones. Shares habitat with Arvicanthis blicki, Tachyoryctes macrocephalus and Stenocephalemys albocaudata and Otomys typus (Yalden 1988, Sillero-Zubiri et al. 1995a). 251

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Abundance The most numerous of the small mammals trapped in the afroalpine habitat of Ethiopia. Comprised 33% of 3083 small mammals in Bale Mts. Population estimates, from live-trapping, suggested densities in different months of 102–658/ha in the grasslands of the Web Valley at 3450 m, and 118–601/ha at 3800–4050 m on the Sanetti Plateau (Sillero-Zubiri et al. 1995a). Biomass estimated, on the basis of actual captures, to be 4.9 kg/ha (annual range 3.9–6.6 kg/ha). Adaptations Active only during daytime. Individuals emerge from their burrows around 08:00h, and the largest numbers of animals above ground occurs at 10:00–12:00h. Activity declines slowly towards 15:00h and sharply thereafter, and no animals are visible above ground after 18:00h (sunrise 06:00h; sunset 18:30h) (see also below). Trapping results reflect this activity: 07:00–12:30h (76%), 12:30–18:30h (23%), overnight (0.5%) (Sillero-Zubiri et al. 1995a). The long claws on the forefeet and pro-odont incisors are presumed to be adaptations for burrowing. Large size, relatively short tail and daytime activity might be adaptations to the cold at high altitudes. Foraging and Food Limited data from stomach analysis suggest that the diet is primarily leaves of dicotyledonous herbs (93% occurrence), with smaller amounts of monocotyledonous leaf (2%), seeds (3%) and insects (2%) (Yalden & Largen 1992).This diet differs from that of most other Lophuromys, which are largely insectivorous (Dieterlen 1976b), and from sympatric Arvicanthis blicki, which eats much monocotyledonous leaf (30%) as well as herbs. Social and Reproductive Behaviour Lives in mixed colonies with Arvicanthis blicki; both species are diurnal, live in burrows and utter high-pitched alarm squeaks that warn of intruding predators (including humans). Because of the difficulty of distinguishing this species from Arvicanthis blicki in the field, observations (given above) refer to the mixed colonies. Reproduction and Population Structure Pregnant and lactating !! found throughout the year. The occurrence of pregnancy is 80–100% at beginning of short wet season in Apr, falls to 40% in May (rainfall lower and variable), rises to 80–100% at the

beginning of the main wet season (Jun–Jul), declines to 40% in the wet season (Aug–Oct), and reaches its lowest level of ca. 20% during dry season (Nov–Mar). This pattern suggests at least two litters per year. Young mainly enter the trappable population in Oct–Feb, contributing ca. 15% of captures; apparent slowness of recruitment is puzzling; possibly due to slow growth or effects of high population density. Mean embryo number: 1.88 ± 0.38 (n = 52 pregnancies; Sillero-Zubiri et al. 1995a). Sex ratio close to unity. Predators, Parasites and Diseases Important prey of Ethiopian Wolves in Bale, comprising 40% of 1307 prey occurrences in 689 scats and 15% by volume of prey (Sillero-Zubiri & Gottelli 1995); the third commonest prey after Tachyoryctes macrocephalus and Arvicanthis blicki. Probably also prey of diurnal raptors, but no detailed information available. Conservation IUCN Category: Vulnerable. Although abundant in suitable habitats, geographic range is very limited. Schlitter (1989) suggested it should be listed as rare. Measurements Lophuromys melanonyx HB: 145.7 (120–180) mm, n = 700 T: 63.7 (30–99) mm, n = 671 HF: 22.8 (21–25) mm, n = 234 E: 22.1 (19–26) mm, n = 235 WT: 94.6 (60–142) g, n = 717 GLS: 33.1 (31.6–34.3) mm, n = 9 GWS: 16.7 (16.0–17.3) mm, n = 10 M1–M3: 5.7 (5.4–6.0) mm, n = 14 Ethiopia Body measurements and weight: Sillero-Zubiri et al. 1995b Skull measurements: Petter 1972a Key References Largen 1992.

Petter 1972a; Sillero-Zubiri 1995a; Yalden & D. W. Yalden

Lophuromys nudicaudus FIRE-BELLIED BRUSH-FURRED RAT Fr: Souris hérissé à ventre feu; Ger. Rotbauchige Bürstenhaarmaus Lophuromys nudicaudus Heller, 1911. Smithson. Misc. Coll. 56 (17): 11. Efulen, Bula country, Cameroon.

Taxonomy Subgenus Lophuromys. Species-group: sikapusi. Closely related to L. sikapusi and L. huttereri (see W. Verheyen et al. 1996 for a review). Previously considered to be a subspecies of L. sikapusi, but now considered to be a valid species on the basis of skull and tooth characters (Rosevear 1969; Dieterlen 1976b, 1979a). The specific name (nudicaudus = naked tail) is misleading (Rosevear 1969). Synonyms: afer, naso, nudicaudatus, parvulus, tullbergi. Subspecies: two. Chromosome number: 2n = 56, with considerable polymorphism (Verheyen & Van der Straeten 1980). Description Small unspeckled species with harsh pelage, relatively short tail and relatively short hindfeet. Dorsal pelage dark

brown with reddish tinge, stiff and harsh; hairs paler at base. Ventral pelage bright rufous (especially in young animals) or yellowish-red. Overall colouration resembles that of young L. sikapusi. Head similar to dorsal pelage, throat and chest similar to ventral pelage. Foreand hindlimbs brownish; claws long, usually pale or brown. Tail short (ca. 58% of HB) with black bristles, dark above, paler below. Skull: rostrum narrow; zygomatic plate narrow. Nipples: 2 + 1 = 6 (W. Verheyen et al. 1996). Geographic Variation The two subspecies are distinguished only by craniometric analysis (W. Verheyen et al. 1996).

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Lophuromys rahmi

L. n. nudicaudus: north and west of Congo R. to Sanaga R. L. n. tullbergi: between Cross and Sanaga rivers, and on Bioko I. Similar Species L. sikapusi. Larger (HB: 136 (122–153) mm); sympatric. L. huttereri. Probably similar in size and characters; rostrum wider and higher; zygomatic plate wide. Distribution Endemic to Africa. Rainforest BZ (West Central Region), north and west of Congo and Ubangi rivers and westwards almost to Cross R. in Nigeria. Recorded from Congo, SW Central African Republic, Gabon, Equatorial Guinea (Rio Muni and Bioko I.), Cameroon and E Nigeria. Recorded also at one locality on the right bank of Aruwimi R. (DR Congo), and may extend westwards along Congo R. to its junction with Ubangi R. Habitat Gaps and clearings within the rainforest and along logging roads (Rosevear 1969; Ray 1996; Malcolm & Ray 2000). Recorded from grassy and open habitats on the forest edge on Mt Cameroon (100–600 m) and Bioko I. (450–1200 m) (Eisentraut 1973). Does not occur in closed canopy rainforests. Lophuromys nudicaudus

Abundance Relatively rare, especially in disturbed forests. In SW Central African Republic, comprised 2.4% of small mammals (n = 704) along secondary logging roads (Ray 1996). Remarks Terrestrial and diurnal (Rosevear 1969, Malcolm & Ray 2000). Stomach contents of two individuals consisted mainly of insects (80–100%) (Dieterlen 1976b). Solitary (Rosevear 1969). Number of embryos: 2, 5 (n = 2). Conservation IUCN Category: Least Concern. Probably not threatened because of its large geographic range.

T: 61.8 (47–74) mm, n = 33 HF: 18.6 (16.3–21.0) mm, n = 44 E: 14.6 (10–18) mm, n = 40 WT: 39.5 (29–52) g, n = 23 GLS: 27.9 (26.5–29.3) mm, n = 34 GWS: 13.4 (12.2–14.2) mm, n = 43 M1–M3: 4.4 (3.9–5.0) mm, n = 47 Throughout geographic range (W. Verheyen et al. 1996) Key References 1996.

Dieterlen 1976b, 1979a; W. Verheyen et al.

Measurements Lophuromys nudicaudus HB: 106.2 (89–119) mm, n = 41

Fritz Dieterlen

Lophuromys rahmi RAHM’S BRUSH-FURRED RAT Fr. Souris hérissé de Rahm; Ger. Rahms Bürstenhaarmaus Lophuromys rahmi Verheyen, 1964. Rev. Zool. Bot. Afr. 69: 206. Bogamanda near Lemera, Kivu, DR Congo.

Taxonomy Subgenus Lophuromys. Species-group: flavopunctatus. Synonyms: none. Chromosome number: not known. Description Small beautiful dark reddish unspeckled rat with short-tail, very short hindfeet and short rounded ears. Pelage rather harsh. Dorsal pelage dark reddish-brown, dorsal hairs pale reddish usually with dark tip. Some individuals may have pale tips to dorsal hairs to give slight speckling to pelage. Flanks paler than dorsal pelage. Ventral pelage bright reddish-orange in both young and adults; hairs paler at base. Head similar to dorsal pelage but duller. Ears short and rounded. Hindfeet very short. Tail short (ca. 51% of HB), darkish-brown above, paler below. Skull: short with rather broad interorbital constriction. Nipples: not known.

Geographic Variation

None recorded.

Similar Species L. cinereus. On average larger; dorsal pelage greyish-brown; very rare. L. flavopunctatus. On average larger; dorsal pelage reddish-brown; widespread and common. L. medicaudatus. Similar in size; tail relatively longer; pelage olivebrown; limited distribution, rare. L. woosnami. On average larger; tail longer and relatively longer; dorsal pelage brown tinged with olive-grey; limited distribution, rare. Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Restricted to montane forests bordering Albertine Rift Valley of E 253

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Abundance Extremely rare, comprising less than 0.05% of terrestrial small mammals captured in montane forests and grasslands. Remarks Terrestrial. Probably burrows amongst roots of trees. Mostly active during the night (Rahm 1967). Omnivorous, preferring insects (mostly larvae), small grubs, caterpillars, beetles, ants, etc. May also eat seeds. Forages on ground in leaf litter (Dieterlen 1976b). Embryo numbers: 2 (n = 2). Pregnant !! recorded in Feb and Jul. Adult size attained when HB >95 mm and weight >30 g. Males fecund when testes 5–10 mm. Sex ratio 16"" : 5!! (Dieterlen 1976b). Conservation IUCN Category: Endangered. This species is threatened because of its rarity and very restricted geographic range; in this respect it is similar to the other Albertine Rift endemics – L. medicaudatus, L. woosnami and L. cinereus. Schlitter (1989) classified the species as rare.

Lophuromys rahmi

DR Congo and Rwanda around L. Kivu at 1900–2500 m (Dieterlen 1976b, 1987, Verschuren et al. 1983). Also Bwindi Forest, SW Uganda (Kasangaki et al. 2003). Habitat Dense primary montane forest especially in Albizia gummifera–Carapa grandiflora–Parinari excelsa forest. Also recorded in secondary forest with Hagenia and Macaranga trees, and in sparse bamboo stands (Hagenia abyssinica–Sinarundinaria alpina) with ground cover of grass (Dieterlen 1976b). May show a preference for habitats with small streams (Rahm 1967) although this preference is uncertain (Dieterlen 1976b, Verschuren et al. 1983).

Measurements Lophuromys rahmi HB: 102 (95–116) mm, n = 21 T: 52.6 (48–56) mm, n = 21 HF: 16.3 (13–18) mm, n = 21 E: 12.5 (10–15) mm, n = 21 WT: 32.5 (30–45) g, n = 21 GLS: 25.3 (24.7–25.9) mm, n = 14 GWS: 14.1 (13.3–14.8) mm, n = 11 M1–M3: 4.1 (3.8–4.4) mm, n = 16 DR Congo (Dieterlen 1976b; SMNS) Key References Dieterlen 1976b, 1987; Verheyen 1964b; Verschuren et al. 1983. Fritz Dieterlen

Lophuromys roseveari ROSEVEAR’S BRUSH-FURRED RAT (MOUNT CAMEROON BRUSH-FURRED RAT) Fr. Souris hérissé de Rosevear; Ger. Rosevears Bürstenhaarmaus Lophuromys roseveari Verheyen, Hulselmans, Colyn and Hutterer, 1997. Bull. Inst. Roy. Sci. Nat. Belgique, Biol. 67: 167. Musake (slopes of Mount Cameroon), Cameroon. 1850–2200 m.

Taxonomy Subgenus Lophuromys. Species-group: sikapusi. Previously considered to be a montane race of L. sikapusi (Eisentraut 1963, 1973, Rosevear 1969, Dieterlen 1976b, 1979a) and now considered as a valid species (Verheyen et al. 1997). Morphologically and morphometrically similar to other species in the sikapusi speciesgroup, but its status and systematic relationships need investigation (Verheyen et al. 1997, Musser & Carleton 2005). Synonyms: none. Chromosome number: not known.

Fore- and hindlimbs brown; claws very long, variable and brownish. Tail short (ca. 50% of HB), dark brown. Skull (compared to L. sikapusi): slender and fragile, with narrow choanae, more inclined borders of the zygomatic plates, supraorbital ridges and notch weakly developed, and zygomatic arches more slender; mandible relatively longer and with a more slender angular process (Verheyen et al. 1977). Nipples: 2 + 1 = 6, 1 + 1 = 4. Geographic Variation

Description Medium-sized reddish-brown unspeckled rat, with long dorsal hairs on rump and a short tail. Dorsal pelage reddishbrown; hairs reddish at base with reddish-brown tip. Hairs on rump long and dense (13–14 mm). Ventral pelage pale reddish-brown or dark cinnamon-brown. Head similar to dorsal pelage. Ears rather large.

None recorded.

Similar Species L. sikapusi. Similar in size; ears on average shorter; dorsal pelage reddish-brown (probably paler); different skull characters (details in profile); widespread and common.

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Lophuromys sikapusi

Habitat Montane forest, grasslands, forest fringes, small patches of woodland, gardens and plantations. Remarks

Apparently no other information available.

Conservation IUCN Category: Least Concern. Distribution is restricted; changes in land use on the mountain may be cause of concern. Measurements Lophuromys roseveari HB: 127.5 (104–141) mm, n = 32 T: 66.0 (50–78) mm, n = 29 HF: 22.5 (20–25) mm, n = 32 E: 18.5 (16–21) mm, n = 28 WT: 63.5 (49–88) g, n = 24 GLS: 31.3 (30.3–32.6) mm, n = 15 GWS: 14.7 (13.9–15.6) mm, n = 21 M1–M3: 4.8 (4.5–5.1) mm, n = 36 Mt Cameroon (Verheyen et al. 1997; BMNH, MNHN SMNS, ZFMK) Key References Eisentraut 1963, 1973; Rosevear 1969; Verheyen et al. 1997.

Lophuromys roseveari

Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Mt Cameroon only. All specimens (n = 44) were collected from several localities on the slopes of Mt Cameroon at 1200–3000 m (Eisentraut 1963, 1973).

Fritz Dieterlen

Lophuromys sikapusi RUSTY-BELLIED BRUSH-FURRED RAT Fr. Souris hérissé de l’Ouest; Ger. Braunbauchige Bürstenhaarmaus Lophuromys sikapusi (Temminck, 1853). Esquisses Zoologiques sur la côte de Guiné, p. 160. Dabocrom, Ghana.

Taxonomy Originally described in the genus Mus. Subgenus Lophuromys. Species-group: sikapusi. One form (eisentrauti) previously considered to be a synonym by Musser & Carleton (1993) now considered to be a valid species. Musser & Carleton (2005), following Verheyen et al. (2000), regard ansorgei (SW DR Congo near the mouth of the Congo R., Uganda, W Kenya, N Tanzania) and angolensis (SW DR Congo near the mouth of the Congo R. and NW Angola) as valid species; here they are retained within L. sikapusi pending further revision. Synonyms: afer, ansorgei, angolensis, mantufeli, pyrrhus, tullbergi. Subspecies: none. Chromosome number: 2n = 60, FN = 66–70 (Côte d’Ivoire; Matthey 1958); 2n = 64, FN = 76 (Mt Nimba, Guinea; Gautun et al. 1986). Description Small unspeckled short-tailed rat. Pelage comparatively soft and less stiff than in other Lophuromys. Dorsal pelage rufous to rusty; blackish-brown in some populations. Ventral pelage pale rufous (most) or bright red (Côte d’Ivoire). Head pointed; eyes small, ears short. Limbs short. Fore- and hindfeet short, reddish-brown, with long dark curved claws. Digit 1 of forefoot much reduced. Tail short (ca. 64% of HB), dark with scaly rings and short dark bristles. Skull: zygomatic plate comparatively broad (2.82 [2.4–3.1 mm]). Nipples: 1 + 2 = 6.

Geographic Variation Dorsal pelage is darkish-brown, and ventral pelage is also unusually dark, in ansorgei from W Kenya. Ventral pelage strikingly bright red in pyrrhus from N Uganda and S Sudan (Dieterlen 1987). Similar Species L. roseveari. Ear on average longer; dorsal pelage reddish-brown to blackish-brown; different skull characters (details in species profile); Mt Cameroon only, where may be sympatric. L. flavopunctatus. Pelage harsh and brush-like, speckled; dorsal pelage blackish-brown; ventral pelage pale brown; tail on average shorter and relatively shorter; nipples 2 + 1 = 6 or 1 + 1 = 4; 2n = 68 or 70. Distribution Endemic to Africa. Rainforest BZ (Western, West Central and East Central Regions) and adjacent Rainforest–Savanna Mosaic. Recorded from Sierra Leone to Cameroon, southwards to Gabon, Equatorial Guinea, Congo, SW DR Congo and N Angola, and east to Central African Republic, N DR Congo (north of Congo R.), S Sudan, Uganda and W Kenya. In Tanzania, patchy distribution in some montane habitats (Dieterlen 1976b).

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Republic, the diet (as assessed by volume) varied according to season: in the dry season 60% insects (especially ants), 18% vegetable pulp and 22% miscellaneous (hairs, soil and myriopods etc.); and in the wet season, 46% insects (mainly ants and termites), 38% vegetable pulp, 5% seeds and 10% miscellaneous (hair, soil, etc.) (GenestVillard 1980). In Uganda, ants and other insects comprised the major proportion of the food (n = 37 stomachs; Delany 1964). In S Nigeria, insects were found in 78% of stomachs, earthworms in 85% and plant material in 20% (n = 61 stomachs; Funmilayo & Akande 1979b). Similar diets are reported from DR Congo (Verheyen & Verschuren 1966), Rwanda (Misonne 1965a) and Côte d’Ivoire (Heim de Balsac & Aellen 1965). Social and Reproductive Behaviour Probably solitary. Torn ears and mutilated tails are not uncommon, suggesting intra-specific aggression.

Lophuromys sikapusi

Habitat Dense moist grasslands, secondary growth, agricultural fields, abandoned farmlands, swamps and grassy plantations where there is abundant low cover. In primary and secondary rainforests, occurs only in grass and herbaceous patches (e.g. after a tree fall) and where grass and bushes grow in open areas. These habitats are preferred because they provide moist soil for digging, and abundant insects, throughout the year (see below). At Mt Nimba, Guinea, occurs up to 1600 m (Gautun et al. 1986). In Uganda, very common in heavily grassed bush country (Delany & Neal 1966). Abundance Common and often numerous in preferred moist habitats, e.g. in S Nigeria comprised 46% of small rodents in swampy stream banks (n = 43, 7 spp.) and 13% in arable fields (n = 138, 8 spp.) (Funmilayo & Akande 1979a). Uncommon in primary rainforest, e.g. 2.9% of seven species of small rodents (n = 482) in a Nigerian rainforest, all individuals occurring in open herbaceous patches (Happold 1977). In Uganda, in grassland, comprised 14.5% of rodents (n = 931, 10 spp.) (Cheeseman & Delany 1979). Adaptations Terrestrial, nocturnal and crepuscular (Cheeseman 1977). The long claws are used for scratching and digging soil while making tunnels through litter and long grass (Happold 1987). Nests of dry grass are constructed on or just under the surface. As in other species of the genus, the strong odour may be unattractive, and may reduce predation by some terrestrial carnivores but not by predatory birds (Dieterlen 1976b). Normally lives in moist rank dense habitats that are not burned; however, in grassland in Uganda, numbers declined after burning (Cheeseman & Delany 1979). Foraging and Food Insectivorous and omnivorous. Forages by searching and digging in dead leaves or litter, where ants, termites, other small or large insects, millipedes, earthworms, molluscs and even carrion are devoured opportunistically. May also eat soft fallen fruits and small seeds of certain tree species. In Central African

Reproduction and Population Structure Young !! become pregnant when 40 g (Happold 1987). In Uganda, a relatively high proportion of pregnant/lactating !! occurred in the wet seasons (Mar–Jun and Sep–Dec; n = 183; Delany & Neal 1969, Cheeseman & Delany 1979). In S Nigeria, pregnancies also recorded mainly in the wet season (Mar–Jun), with reduced pregnancy rates in the ‘little dry season’ (Jul–Aug) and in the ‘long dry season’ (Nov– Feb) (Happold 1974). Gestation: about 30 days (Genest-Villard 1968). Litter-size usually 2–3. Mean embryo numbers: 2.6 (range 1–3, n = 8) in Virunga Mts, DR Congo (Verschuren et al. 1983); 3.0 (range 2–5, n = 13) in S Nigeria (Happold 1974). At birth, young are precocial, weight ca. 8 g. Animals born in captivity weighed 35 g at three weeks of age (= 50% adult weight), and adult weight when 5–8 weeks (Genest-Villard 1968). Predators, Parasites and Diseases In grasslands of Rwenzori N. P., Uganda, various species of predatory birds, carnivores (mongooses, genets, servals) and snakes were potential predators of small rodents (Cheeseman & Delany 1979). Conservation IUCN Category: Least Concern. A widely distributed and common species. Measurements Lophuromys sikapusi HB: 118.5 (110–130) mm, n = 10 T: 74.2 (65–82) mm, n = 10 HF: 22.2 (21–23) mm, n = 10 E: 16.2 (15–17) mm, n = 10 WT: 62.7 (51–79) g, n = 10 GLS: 31.2 (29.8–32.6) mm, n = 10 GWS: 15.0 (14.4–15.9) mm, n = 10 M1–M3: 5.0 (4.8–5.3) mm, n = 10 SW Nigeria (Happold 1974) Key References Cheeseman & Delany 1979; Dieterlen 1976b; Genest-Villard 1968, 1980; Happold 1974, 1987. Fritz Dieterlen

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Lophuromys woosnami

Lophuromys woosnami WOOSNAM’S BRUSH-FURRED RAT Fr. Souris hérissé de Woosnam; Ger. Woosnams Bürstenhaarmaus Lophuromys woosnami Thomas, 1906. Ann. Mag. Nat. Hist., ser. 7, 18: 146. Mubuku Valley Rwenzori East, Uganda; 6000 ft (1820 m).

Taxonomy Subgenus Kivumys. Species-group: woosnami. Synonyms: prittiei, undescribed (see below). Subspecies: two or three (Dieterlen 1976b). Chromosome number: 2n = 42, FN = 72 (Dieterlen 1976b, Maddalena et al. 1989). Description Medium-sized and slender-bodied rat with a long tail, long ears, long hindfeet and comparatively short claws. Pelage soft and conspicuously glossy; unspeckled. Dorsal pelage brown tinged with olive-grey; hairs reddish-brown at base. Ventral pelage pale brown tinged with reddish. In subadults, reddish colouration of dorsal and ventral pelage is brighter and more intense than in adults. Ears long, naked, rounded at tip. Fore- and hindfeet whitish with comparatively short claws. Hindfoot long (cf. other species in genus). Tail long (ca. 105% of HB), mostly naked with dark bristles, dark above, pale flesh-colour below. Nipples: 1 + 1 = 4. Geographic Variation L. w. woosnami: Rwenzori Mts. HB: ca. 110 mm. L. w. prittiei: highlands of Kigesi, Uganda; Virunga Mts in Uganda, Rwanda, DR Congo; Nyungwe Forest in Rwanda and Kibira Forest in Burundi. HB: ca. 115 mm. L. w. undescr.: west of L. Kivu, DR Congo. HB: ca. 121 mm. Similar Species L. flavopunctatus. Dorsal pelage reddish-brown; E shorter; HF shorter; T much shorter. Distribution Endemic to Africa. Afromontane–Afroalpine BZ in Uganda, Burundi, Rwanda and DR Congo. Recorded from the Albertine Rift Valley and bordering mountains from Rwenzori Mts in the north to the Itombwe massif (E DR Congo) and mountains of Burundi in the south. Distribution discontinuous, confined to forested mountains on both sides of the Rift Valley; 1800–3880 m.

Lophuromys woosnami

long legs, long tail and long ears suggests that these mice are very mobile and probably have a large home-range. Activity is mostly nocturnal (Rahm 1967, Delany 1972). Captive !! constructed simple nests of dry grass and leaves. Foraging and Food Omnivorous. Stomach contents contained 40–50% arthropods (and some molluscs) and 50–60% vegetable material, seeds and bulbs (but not green matter) (n = 15; Dieterlen 1976b, Verschuren et al. 1983). Captive animals preferred meal worms, grasshoppers and dry insect larvae etc., but also ate sweet apples, peanuts and grains of sunflowers. Drinking water is essential.

Habitat Undergrowth in montane forests, cleared areas in forests, old and new bamboo forests, and amongst rocks in afroalpine vegetation (Senecio, Lobelia).

Social and Reproductive Behaviour Captive animals are tame from the first day of captivity, showing no fear or panic even when handled for the first time. When in captivity for several Abundance Common in suitable habitats and often the most months, adults and their young live together peacefully and without numerous species of small rodents. Commonly syntopic with L. aggression towards one another. Individuals exhibit mutual grooming. flavopunctatus. In many habitats in Kahuzi-Biega N. P., E DR Congo, Olfactory communication seems to be important. The typical one species of Lophuromys was considerably commoner than the other odour, caused by sebaceous glands in the glossy pelage, probably suggesting inter-specific competition. Percentage occurrence of L. has a socially stimulating effect (Dieterlen 1976b). Females exhibit woosnami and L. flavopunctatus in small mammal communities was, ‘midwifery’ behaviour when a mother is giving birth; they try to get respectively, 25% and 19% in montane secondary forests, 42% and hold of the umbilical cord and, some moments later, to ‘steal’ and eat 6% in cleared montane primary forest, 13% and 4% in undisturbed the placenta.Young are raised communally, several !! participating montane primary forest, 56% and 8% in old bamboo, and 20% and in the care of the young (Dieterlen 1976b); this behaviour is similar 48% in young bamboo with grassy cover (Dieterlen 1976b). to that of Acomys spp. (Dieterlen 1962). Adaptations Terrestrial long-footed rat; runs with a ‘jumping gallop’ and good at climbing. This kind of locomotion, as well as the

Reproduction and Population Structure In Kivu Province, DR Congo, reproduction is seasonal, occurring during the wet 257

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season (Sep–Apr), with peaks in pregnancy rate (80–100%) in Oct– Dec and in Mar–Apr (Dieterlen 1976b). After a transition period in May, pregnancies not recorded during the dry season (Jun–Aug). Gestation: at least 32 days. At birth, young are precocial; postnatal development is rapid. Mean embryo numbers: 1.94 (1–3; mode 2 [84% of total]; n = 96). Females and "" became sexually active when 7–8 weeks old; at this age "" weigh 36–40 g (testes 10– 12 mm). Adult "" weighing 50–60 g have permanently large testes of 20–25 mm. Sex ratio: 58% "": 42% !! (n = 454). Predators, Parasites and Diseases No information. Conservation IUCN Category: Least Concern. The forests on the mountains of the Albertine Rift Valley, the only habitat of this species, are fragmented and declining in area. The future of the species depends on adequate protection of these forests.

Measurements Lophuromys woosnami HB: 121 (111–135) mm, n = 20 T: 123 (114–133) mm, n = 20 HF (c.u.): 26.5 (26–28) mm, n = 20 E: 22.9 (20–25) mm, n = 20 WT: 47 (38–64) g, n = 20 GLS: 31.5 (30.3–33.2) mm, n = 20 GWS: 14.4 (13.8–15.3) mm, n = 20 M1–M3: 4.7 (4.4–4.9) mm, n = 20 DR Congo (Dieterlen 1976b, SMNS) Key References Dieterlen 1976b, 1987; Kingdon 1974; Verschuren et al. 1983. Fritz Dieterlen

GENUS Uranomys Rudd’s Brush-furred Mouse Uranomys Dollman, 1909. Ann. Mag. Nat. Hist., ser. 8, 4: 551. Type species: Uranomys ruddi Dollman, 1909.

Uranomys ruddi.

A monotypic genus widespread in savannas of West and East Africa. The genus is characterized by small size, dorsal pelage composed of coarse brownish brush-like hairs, white ventral pelage and short tail. The skull has pro-odont incisors (except in one form), the palatine bones extend posteriorly to M3 and partly cover the mesopterygoid fossa, the small cheekteeth are similar to those of Acomys spp., and there is an enlarged process on the external aspect of the mandibular ramus (Figure 43). Further details are given in the species profile. The genus originally contained seven species (see synonyms below), but is now considered to contain only one species, Uranomys ruddi. The genus is closely related to Lophuromys and Acomys, and (like these genera) is placed in the subfamily Deomyinae of the Muridae (Musser & Carleton 2005) rather than in the subfamily Murinae as in previous classifications (e.g. Musser & Carleton 1993). Other investigations (biochemical, molecular and karyological) confirm the monophyletic origin of Uranomys, Lophuromys and Acomys. Full details of these relationships are given in Musser & Carleton (2005, and references therein). The single species is Uranomys ruddi.

Figure 43. Skull and mandible of Uranomys ruddi (HC 1748).

D. C. D. Happold

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Uranomys ruddi RUDD’S BRUSH-FURRED MOUSE Fr. Souris de Rudd; Ger. Rudds Bürstenfellmaus Uranomys ruddi Dollman, 1909. Ann. Mag. Nat. Hist., ser. 8, 4: 52. Kirui, Mount Elgon, Kenya. 6000 ft (1820 m).

Taxonomy Now considered as a single species with a very broad distribution (Verheyen 1964a; see also profile Genus Uranomys); however, variations in chromosome number perhaps indicate that more than one species is present. Synonyms: acomyoides, foxi, oweni, shortridgei, tenebrosus, ugandae, woodi. Subspecies: none. Chromosome number varies geographically: 2n = 50 (Senegal), 2n = 58 (Côte d’Ivoire), 2n = 52 (Central African Republic). Description Small mouse with small limbs, short tail and stiffened hairs on back and rump. Pelage short and stiff. Dorsal pelage grey to grey-brown, speckled with pale brown and black; hairs dark grey with pale brown tip, or with pale brown terminal band and black tip. Dorsal hairs stiffened as in an artist’s brush, and not easily rubbed the wrong way. Dorsal hairs not spiny as in Spiny Mice (Acomys spp.). Ventral pelage dirty-white, sometimes tinged with pale cinnamon; colour of flanks merges gradually to colour of ventral pelage. Head rather slim and pointed, with small eyes and small ears. Chin, throat, chest and limbs white. Limbs short. Tail short (ca. 60% of HB), brownish, with scales and numerous very small black bristles. Skin thin and fragile; many individuals have damaged ears, and tail is frequently shortened or completely absent. Incisor teeth pro-odont (although occasionally opisthodont in some juveniles). See also genus profile. Nipples: 3 + 3 = 12. Geographic Variation The form acomyoides from Ghana has orthodont incisors (Ingoldby 1929 in Musser & Carleton 2005).

Similar Species Acomys spp. Dorsal hairs spiny and thicker. Mus minutoides/musculoides. Smaller, without stiffened hairs. Distribution Endemic to Africa. Guinea Savanna BZ, Northern Rainforest–Savanna Mosaic, Eastern Rainforest–Savanna Mosaic, and parts of Zambezian Woodland BZ. In West Africa, recorded from Sierra Leone, Gambia, Guinea, Liberia, Côte d’Ivoire, Ghana, Togo, Benin and Nigeria. In central and eastern Africa, recorded in Uganda, Zimbabwe, Mozambique and Malawi. Isolated records in N Cameroon, Central African Republic, Chad, NE DR Congo, C Tanzania and W Ethiopia. May be more widespread than records indicate (see Abundance). Distribution disjunct. Habitat Grasslands. The rarity of the species in most places, and its abundance in a few localities (see below), suggest that the preferred habitat has abundant grasses, few trees and moist soil, which provides moist or semi-swampy conditions. Also occurs in farmland (where soil is moist and friable) and oil palm plantations (where there are moist grasses and grass litter). One individual in Malawi was found ‘in hole of Mole-rat on wooded hills’, and others were found close to a river swamp where there were many ant and termite mounds (Hanney 1965). Abundance Generally a rare species and seldom encountered. Not recorded from many savanna habitats and rare in others. However, is quite common at a few study sites, e.g. comprised 29% of small rodents in grass savanna at Lamto, Côte d’Ivoire (n = 745; Bellier 1968), 31% in farmland at Ibadan, Nigeria (n = 710; Happold 1974), 44% in grass swamp at Ibadan (n = 86; D. C. D. Happold unpubl.) and 55% in grass savanna at Dabou, Côte d’Ivoire (n = 838; Bellier 1968). Comprised 8% (n = 1448) in oil palm plantation in Côte d’Ivoire (Bellier 1968). Population numbers not adversely affected by burning of grasses at Ibadan, Nigeria, probably because of fossorial habits (D. C. D. Happold unpubl.). Adaptations Nocturnal and crepuscular; terrestrial. Brush-furred Mice dig burrows, using their short strong feet. An excavated nest (in Côte d’Ivoire) about 15 cm below ground had a nest chamber 6–8 cm in diameter lined with fresh cut grass; another tunnel descending to about 30 cm below ground (Bellier 1968). Entrance holes were plugged. Brush-furred Mice seem to be partly fossorial; specimens in captivity burrowed under litter and grass when disturbed, and they blocked the entrances to their burrows and nestbox with soil which was sometimes glued together with fluid (? saliva) (D. C. D. Happold unpubl.). The pro-odont incisor teeth may be used for catching and holding insect prey, in a similar manner to shrews.

Uranomys ruddi

Foraging and Food Primarily insectivorous: the contents of two stomachs contained adult insect remains, dipteran larvae and ant pupae (Malawi; Hanney 1965). Vegetable foods include cassava 259

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(manioc) and bulbs of savanna plants (Bellier 1968). Food has not been found in burrows. Social and Reproductive Behaviour Little information. Burrows in Côte d’Ivoire were inhabited by two adults (Bellier 1968). In captivity several individuals live together amicably. Reproduction and Population Structure In Côte d’Ivoire (Bellier 1968), pregnancies recorded in all months except Jan (early dry season) and juveniles in all months except Oct and Feb. Peak of reproductive activity in late wet season (Aug–Dec incl.). Seasonal variation in mean number of embryos: 4.3–5.7 (late wet season), 2.6–3.7 (dry season and early wet season) (Bellier 1968). Minimum weight at pregnancy: 22 g (Happold 1974). Sex ratio (live trapping) was 3 : 1 (Nigeria; D. C. D. Happold unpubl.). Population structure unknown. At Ibadan, Nigeria, during a 5-month study, 72% of individuals were caught once and 18% were caught twice (n = 39 individuals), suggesting a rapid turnover of the population (D. C. D. Happold unpubl.). Predators, Parasites and Diseases Preyed upon by Barn Owls Tyto alba (Rosevear 1969).

Conservation IUCN Category: Least Concern. The widespread distribution suggests that the species is not threatened. However, its rarity (in most habitats) and loss of suitable habitat may be cause for concern. Measurements Uranomys ruddi HB: 108 (101–119) mm, n = 15 T: 63 (55–68) mm, n = 15 HF: 17 (16–18) mm, n = 15 E: 13 (12–14) mm, n = 15 WT (""): 37 (31–51) g, n = 5 WT (!!): 30 (22–31) g, n = 5 GLS: 25.1 (24.0–26.4) mm, n = 15 GWS: 12.3 (11.3–13. 4) mm, n = 15 M1–M3: 4.1 (3.8–4.3) mm, n = 15 Measurements: Côte d’Ivoire (MNHN) Weight: Nigeria (Happold 1987) Key References Bellier 1968; Happold 1974, 1987; Rosevear 1969; Verheyen 1964a. D. C. D. Happold

Subfamily GERBILLINAE – Gerbils and Jirds Gerbillinae Gray, 1825. Ann. Philos., n. s., 10: 342. Ammodillus (1 species) Desmodilliscus (1 species) Desmodillus (1 species) Gerbilliscus (12 species)* Gerbillurus (4 species) Gerbillus (36 species) Meriones (3 species) Microdillus (1 species) Pachyuromys (1 species) Psammomys (2 species) Sekeetamys (1 species) Taterillus (8 species)

Ammodile Dwarf Gerbil Short-tailed Gerbil Gerbils Hairy-footed Gerbils Gerbils Jirds Pygmy Gerbil Fat-tailed Gerbil Sand Rats Bushy-tailed Jird Gerbils

p. 262 p. 264 p. 266 p. 268 p. 287 p. 295 p. 333 p. 339 p. 341 p. 343 p. 347 p. 349

*Formerly Tatera

Members of this large subfamily, numbering 16 genera and about 101 species (Musser & Carleton 2005), occur throughout much of Africa and across the Palaearctic desert and steppe, from Asia Minor and the Middle East to southern Mongolia and northern China. In Africa, there are 12 genera and 71 species (see list above). Gerbils mostly live in arid and semi-arid environments, mostly in areas of sparse vegetation. Within Africa, they occur in many biotic zones except the Rainforest and Afromontane–Afroalpine BZs. Typical habitats include sandy and clay deserts, dunes and alluvium with meagre grass or brush cover, gravelly plains and semi-deserts, and a wide mixture of grasslands and woodlands from very dry and open to moderately moist. Certain forms, such as Pachyuromys and some Gerbillus, thrive in some of Africa’s bleakest, seemingly inhospitable, habitats. In such environments, gerbils fill a terrestrial and largely granivorous niche

(the herbivorous Psammomys is a notable exception); most species are nocturnal and a few are diurnal (Psammomys and some Meriones). Most gerbilline genera (12 of 16) contain species with distributions in Africa, and many are endemic to the continent (Ammodillus, Desmodilliscus, Desmodillus, Gerbilliscus [formerly Tatera], Gerbillurus, Microdillus, Pachyuromys, Taterillus). Three genera have distributions that include Africa and parts of the Middle East or Asia (Gerbillus [including Dipodillus], Meriones, Psammomys), and one of these has its greatest number of species within Africa (Gerbillus – 39 of 51 spp.). Meriones is the only genus that contains more species with ranges outside of the continent (13 of 17); the four species distributed entirely or partially in Africa occur only along the Mediterranean coast. Psammomys, too, has a predominantly North African distribution but also reaches to the near Middle East and Arabian Peninsula. Clearly, African environments have figured prominently in the evolutionary diversification of the subfamily, and the continent is considered by some to be its place of origin (e.g. Lay 1972). The present-day concentration of the earlybranching clades of the Gerbillinae within the sub-Saharan region (namely Gerbillurina, Taterillina and Ammodillini (Pavlinov et al. 1990) is consistent with this biogeographic interpretation. Most African gerbils are small to medium in body size, but extremes of very small (Desmodilliscus, some Gerbillus) and large (Gerbilliscus, Psammomys, some Meriones) are also represented. Body form ranges from stout and compact (Pachyuromys) to slender and gracile (Gerbillus, Taterillus). Counter-shading is generally pronounced, the dorsal pelage is pale sandy to saturated brown, and the ventral pelage is white. The expanded auditory bullae impart a relatively large and wide shape to the head. Pinnae are small, rounded and well furred in many species,

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or larger, ovate and sparsely covered in some, especially the taterillines Gerbilliscus and Taterillus; conspicuous white to buffy postauricular patches occur in some (Desmodillus, Pachyuromys). The elongated metatarsal bones of the hindfoot reflect the running and semi-saltatory locomotion of most species, although none is strictly bipedal as in the dipodids Allactaga and Jaculus. The plantar surface is naked, moderately furred, or so densely furred as to obscure the plantar pads.The tail may be longer than, or about equal to the head and body, or notably shorter (as in Desmodilliscus and Pachyuromys); it is moderately to densely haired in most forms, with a terminal black pencil or tuft in some species (Taterillus and Meriones). A mid-ventral sebaceous gland is present in many species, and is especially well-developed in "". There are eight nipples (arranged as pectoral, post-axillary and two inguinal pairs) or six nipples (pectoral pair absent). The head and postcranial skeleton exhibits diagnostic traits of the subfamily: supraorbital shelf present and well developed; zygomatic plate broad and dorsal notch deep; lacrimal enlarged, forming a conspicuous ledge over the anterior orbit; mesopterygoid fossa narrowly V-shaped and parapterygoid fossae compressed and cavernous; anterior palatal foramina and posterior palatine foramina extremely long and slit-like; optic foramen as large as sphenoidal fissure; stapedial foramen present, penetrating the wall of the tympanic bulla rather than the petrotympanic fissure; sphenofrontal foramen and accessory foramen ovale absent (except Tatera); angular process of mandible deflected laterally; auditory bullae inflated, both ectotympanic and mastoid chambers, and malleus of the perpendicular type; entepicondylar foramen present (except Tatera); scapula with third scapular fossa; vertebral column with 12 thoracic vertebrae and seven lumbar vertebrae (Lay 1972, Carleton 1980, Pavlinov 1980, Carleton & Musser 1984, Pavlinov et al. 1990). In all African species, the molars are rooted, anchored by accessory rootlets in most species, and their occlusal pattern is lophate, planar or prismatic; protoconulid present on M1, distinct from or fused with anteroconid; M3 greatly reduced and cylindriform, the posterior lamina (hypoconid and entoconid) of M3 absent; upper incisors with single groove (smooth in Psammomys) (Ellerman 1941, Rosevear 1969, Pavlinov 2001). Other notable characters of those Gerbillinae studied include: the glans penis is of the complex type; preputial glands are generally absent, remainder of accessory reproductive gland of " is complete; the stomach is single-chambered and hemiglandular; and the stapedial artery lacks supraorbital and mandibular branches, the orbital blood supply instead is formed by the infraorbital branch (Arata 1964, Vorontsov 1967, Bugge 1970). Morphology of the auditory bullae has figured prominently in the taxonomic and phylogenetic understanding of gerbilline rodents (Lay 1972, Pavlinov 1980, Pavlinov et al. 1990). Evolutionary increases in pneumatization of the mastoid portion are especially significant and have given rise to spectacular middle ear anatomies, such as those that characterize Pachyuromys duprasi and Meriones crassus. Enlargement of the middle ear chambers lessens impedance of air space behind the tympanic membrane and thereby enhances sensitivity to relatively low sound frequencies (Lay 1972, 1993, Webster & Webster 1984). These volumetric adaptations – coupled with modifications in the ossicular lever system, expanded surface area of the tympanic and accessory tympanic membranes, and acoustic specializations of the inner ear – constitute a clever predator-detection system for animals

a

b

c

d

e

f

g

h

Figure 44. Structure of the auditory bulla in several genera of Gerbillinae to show relative and comparative sizes of tympanic bulla and mastoid. (a) Arvicanthis niloticus (Murinae) without any enlargement of auditory bulla, (b) Gerbillurus, (c) Microdillus, (d) Gerbillus, (e) Meriones, (f) Sekeetamys, (g) Desmodilliscus, (h) Pachyuromys. Dark stipple = tympanic bulla. Light stipple = mastoid. Arrows indicate position of occipital condyles (not visible in lateral view). Variations between genera include size of tympanic bulla and mastoid (relatively and comparatively), size of bulla in relation to GLS, and position of posterior end of auditory bulla to the posterior end of the occipital condyles (see text for further details). Species within a genus may show slight variation to the generalized genus condition shown here. The condition in Dipodidae (Allactaga, Jaculus) is similar to that of Desmodilliscus. (Based on original illustration by D. C. D. Happold.)

living in open environments and must have contributed importantly to the ecological successes of the subfamily (Lay 1972). Indeed, the Gerbillinae contains more species indigenous to the great SaharoGobian realm of Africa and Asia than any other group of Rodentia (Petter 1961, Lay 1991, Shenbrot et al. 1999b). Parallel changes in middle ear anatomy characterize other rodent lineages that have radiated within desert and semi-arid biomes, in both North America (Heteromyidae) and Afro-Asia (Dipodidae) (Figure 44). Although morphological evidence supporting the monophyly of Gerbillinae is substantial (Carleton & Musser 1984, Pavlinov et al. 1990), their phylogenetic stature as a subfamily of Muridae is a relatively recent apprehension, stemming from both palaeontological and molecular investigations. Gerbils were traditionally grouped with the Cricetidae in those classifications that maintained Muridae and Cricetidae as separate families (e.g. Miller & Gidley 1918, Simpson 1945, Misonne 1974), or accorded their own family status as Gerbillidae (Tullberg 1899, Chaline et al. 1977, Pavlinov et al. 1990). Phyletic interpretations of nuclear and mitochondrial DNA sequences, however, cladistically affiliate gerbillines with murines and deomyines (Martin et al. 2000, Michaux & Catzeflis 2000, Michaux et al., 2001). Using molecular-clock estimates, Michaux et al. (2001) suggested that the divergence of Gerbillinae occurred during the early to middle Miocene, about 18 to 16 mya years ago. 261

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Such divergence times are wholly reasonable in the context of the rich fossil history of gerbils. Their origin is convincingly linked to myocricetodontines, an extinct group known from the Miocene of both Africa and Asia and believed to represent the ancestral stock of the subfamily (Jaeger 1977b, Tong 1989, Lindsay 1994, Wessels 1996). Tong & Jaeger (1993) suggested that the Murinae were also derived from an early myocricetodontine, with the split between the two groups transpiring around 16 mya years ago. The oldest true gerbilline so far known from Africa is Protatera, from the late Miocene of Morocco (Jaeger 1977b); representatives of living genera appear in the late Pliocene and are commonplace in Pleistocene faunas (e.g. Lavocat 1978, Denys 1987a, 1989a, Senut et al. 1992, Avery 1998). Generic interrelationships are relatively well understood, as set forth in the comprehensive monograph of Pavlinov et al. (1990, see also Pavlinov 2001), which serves as the basis for recognizing the following tribes (-ini) and subtribes (-ina) among African genera

(ranks as adapted by McKenna & Bell 1997). Here (in contrast to Musser & Carleton 2005), Dipodillus is considered as a synonym of Gerbillus (see Gerbillus profile) and hence 12 genera (Table 22) are recognized as follows: Taterillini: Gerbillurina (Desmodillus, Gerbillurus); Taterillina (Gerbilliscus, Taterillus). Ammodillini: (Ammodillus). Gerbillini: Desmodilliscina (Desmodilliscus); Gerbillina (Gerbillus [including Dipodillus], Microdillus); Pachyuromyina (Pachyuromys); Rhombomyina (Meriones, Psammomys, Sekeetamys). The profiles for each genus, and for each species within each genus, are given alphabetically, not by their tribal and subtribal affiliations. Michael D. Carleton & Guy G. Musser

GENUS Ammodillus Ammodile Ammodillus Thomas, 1904. Ann. Mag. Nat. Hist., ser. 7, 14: 102. Type species: Gerbillus imbellis de Winton, 1898.

Ammodillus is a monotypic genus found only in Somalia. It is very similar to Gerbillus, and distinguished from Gerbillus by special skull characters (especially the lack of a coronoid process on the mandible, and the posterior convergence of the upper cheekteeth). Characters

of the genus are given in the species account below. The only species is Ammodillus imbellis. D. C. D. Happold

Table 22. Genera in the subfamily Gerbillinae. Arranged in order of increasing head and body length. Genera

HB (mean or range) (mm)

Tail (% of HB)

Number of cheekteeth

Bullae (% of GLS)

Molar formb

Position of posterior palatine foramina

Desmodillicus (1 sp.) Microdillus (1 sp.)

55 72

75–80% 80%

3/2 3/3

ca. 40% 39%

Cuspidate Cuspidate

Mid M2 to anterior of M1 Mid M2 to front margin of M1

Gerbillus (36 spp.)

Mostly 80–129a

105–160%

3/3

29–36%

Cuspidate

Posterior M2 to first row of cusps of M1

Gerbillurus (4 spp.)

96–105

120–140%

3/3

ca. 30%

Cuspidate

Posterior M2 to mid M1

Ammodillus (1 sp.)

99

145%

3/3

33–37%

Front margin to hind margin of M2. Small

Pachyuromys (1 sp.)

108.3 (93–121)

54%

3/3

47%

Desmodillus (1 sp.)

110

74–80%

3/3

41%

Taterillus (8 spp.)

107–116

130–150%

3/3

26–30%

Cuspidate Cuspisdate to laminate Cuspidate Cuspidate to laminate

Sekeetamys (1 sp.)

118

120%

3/3

33%

Prismatic

Mid M2 to mid M1

Psammomys (2 spp.) Gerbilliscus (12 spp.)c

122–160

70–86%

3/3

32%

Prismatic

Anterior M2 to posterior M1 (very short)d

128–185

95–ca. 130%

3/3

24–31%

Cuspidate

Posterior M2 to mid M1

136

95–100%

3/3

32–40%

Prismatic

Mid M2 to mid M1

Meriones (3 spp.)

Mid M2 to mid M1 Posterior M3 to mid M1 Posterior M2 to front margin M1

a

A few species of Gerbillus have a mean HB of less than 80 mm (G. brockmani 78 mm; G. juliani 63 mm; G. henleyi 65 mm; G. nanus 72 mm). Cuspidate but often laminate in older animals, with transverse sets of cusps joining to form single transverse occlusal surface. c Formerly Tatera. d Shallow grooves may extend anteriorly to posterior end of anterior palatal foramina. b

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Ammodillus imbellis AMMODILE (WALO) Fr. Ammodile; Ger. Walo Ammodillus imbellis (de Winton, 1898). Ann. Mag. Nat. Hist., ser. 7, 1: 249. Goodar, Somalia.

of forelimbs tipped with grey. White supraorbital and postauricular patches. Fore- and hindfeet white. Soles of hindfeet naked. Tail very long (ca. 145% of HB), dark above, paler below, slightly haired; brown hairs (8–10 mm long) on terminal one-third of upper surface form conspicuous pencil. Two skull characters are unique: the molar rows converge posteriorly (more so than in any other genus of the gerbils) and tend to be laminate rather than cuspidate, and there is no coronoid process on the mandible (Roche & Petter 1968, Funaioli 1971 in Nowak 1999). Incisors strongly opisthodont (Figure 45). Nipples: 1 + 2 = 6. Geographic Variation

Similar Species Gerbillus brockmani. Smaller (HB: 71–84 mm). G. somalicus. HF shorter (24–25 mm. Neither of these species has the skull characters of Ammodillus noted above.

Ammodillus imbellis.

Taxonomy Originally described in the genus Gerbillus, but later placed in a new genus, Ammodillus (see below). Synonyms: none. Chromosome number: 2n = 18 (Capanna & Merani 1981). Description Small gerbil with a long tail ending in a brown pencil. Dorsal pelage reddish-fawn to brownish-yellow; hairs grey at base, with reddish fawn terminal band and black tip. Flanks paler and clearer. Ventral pelage white. Hairs of eyebrows, cheeks and upper surface Pencil at tip of tail

Sahel Savanna Zone; Mauritania to Chad Somalia

Absent

North Africa

Absent

South-West Arid BZ Widespread in savannas of West, central and East Africa north of Rainforest Zone

Very welldeveloped Present Absent to well-developed Small

Distribution Endemic to Africa. Somalia–Masai Bushland BZ. Recorded only from Somalia and E Ethiopia. Habitat In S Somalia, found in coastal steppe (Capanna & Merani 1981). In E Ethiopia, found ‘in sandy soil close to wells’ at Gerlogobi (Thomas 1904a). Digs burrows in sandhills (near El Bur).

Notes

Absent Absent Small to well-developed Absent to well-developed Well-developed

Well-developed

None recorded.

Widespread in North, West and East Africa South Africa, Namibia Somalia, Ethiopia

NE Africa North Africa Widespread in semi-arid and savanna habitats south of Sahara. Slight groove on each upper incisor North Africa. Slight groove on each upper incisor Figure 45. Skull and mandible of Ammodillus imbellis (MZUF M4940).

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Abundance Very rare. Only known from a few specimens from seven localities. Remarks The lack of a coronoid process on the mandible results in a weak bite (Nowak 1999); this suggests that Ammodiles can only eat soft foods. No information on diet. Found in small colonies on sandhills (A. Simonetta pers. comm.). Individuals are recorded to fight amongst themselves (Nowak 1999). One specimen was obtained from the stomach of a viper (Bitis sp.). Conservation

IUCN Category: Data Deficient.

Measurements Ammodillus imbellis HB: 99.3 (84–111) mm, n = 15 T: 145.1 (134–160) mm, n = 11 HF: 27.8 (26–29) mm, n = 15 E: 15.8 (14–18) mm, n = 12 WT: n. d. GLS: 31.4 (30.1–32.7) mm, n = 12 GWS: 15.3 (14.7–16.5) mm, n = 12 M1–M3: 4.4 (4.1–4.6) mm, n = 13 Auditory bulla: 10.5 mm, n = 1* Somalia (Roche & Petter 1968) *BMNH Key Reference

Ammodillus imbellis

Roche & Petter 1968. D. C. D. Happold

GENUS Desmodilliscus Brauer’s Dwarf Gerbil Desmodilliscus Wettstein, 1916. Anz. Akad. Wiss. Wien 53: 153. Type species: Desmodilliscus braueri Wettstein, 1916.

Desmodilliscus is a monotypic genus endemic to Africa, distributed throughout the Sahel Savanna BZ from Mauritania and Senegal to Sudan. The characters of the genus are those given below in the species account. The small overall size and the relatively short tail are the main external characters, but skull shape and reduction in the

number of cheekteeth (3/2) are unique among the Muroidea. Dental formula: I 1/1, C 0/0, P 0/0, M 3/2 = 14. L. Granjon

Desmodilliscus braueri BRAUER’S DWARF GERBIL (POUCHED GERBIL) Fr. Gerbille naine de Brauer; Ger. Brauers Zwergrenmaus Desmodilliscus braueri Wettstein, 1916. Anz. Akad. Wiss. Wien 53: 153. South of El Obeid, Sudan.

Taxonomy Setzer (1969) recognized three subspecies, but Rosevear (1969) and Hutterer & Dieterlen (1986) stated there was morphological homogeneity throughout its range. Synonyms: buchanani, fuscus. Subspecies: none. Setzer (1969) recognized three subspecies, but Rosevear (1969) and Hutterer & Dieterlen (1986) indicated that there is morphological homogeneity throughout the range and therefore no subspecies are recognized here. Chromosome number: 2n = 78, aFN = 104 (Senegal; Granjon et al. 1992).

Description Very small gerbil. Dorsal pelage sandy-grey; hairs blackish-grey at base, with sandy subterminal band and minute dark tip, giving a speckled and relatively dark effect. Hairs of the flanks white at base with sandy tip. Ventral pelage entirely white. Head relatively large with short ears, big eyes and white cheeks. White stripe above and behind the eyes, and white postauricular patch. Hindfeet slender; forefeet with a slight covering of short white hairs; soles of hindfeet naked. Tail short (ca. 65–75% of HB), covered with short hairs, but without any pencil. Skull: incisor teeth with curbed

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Desmodilliscus braueri

Desmodilliscus braueri.

front face, each with single deep groove situated laterally; very inflated tympanic bullae and mastoids that extend well posteriorly to the occipital condyles; palate broad; posterior palatal foramina very wide extending anteriorly to M1 and to within about 1 mm of the anterior palatal foramina. Unique in possessing only two molars on the mandible (see genus profile above) (Figure 46). Nipples: 2 + 2 = 8. Geographic Variation None recorded. Similar Species No other species of gerbil has such a small head and body length (Table 22), and no other species of gerbil has only two cheekteeth in each ramus of the mandible.

Figure 46. Skull and mandible of Desmodilliscus braueri (BMNH number not recorded).

Distribution Endemic to Africa. Sahel Savanna BZ. Widespread, most records being between 12° N and 18° N. Recorded from Mauritania and Senegal to Sudan, through Mali, Burkina, Niger, Nigeria, N Cameroon and possibly Chad (Heim de Balsac 1967b, Hutterer & Dieterlen 1986). Geographic range may be extending southwards, due to desertification (Duplantier et al. 1997). Habitat Scrub savannas with sparse vegetation, especially on indurate sandy or sandy-clay soils, often with gravel. In Senegal, found in the northern Acacia-savanna, receiving an annual rainfall of 200–500 mm (Heim de Balsac 1967b, Poulet 1984). Abundance Probably common to very abundant in suitable habitats (although difficult to catch in traps). Common in owl pellets in some localities of N Mali (Heim de Balsac 1967b). Dozens of these gerbils seen during a few hours of night driving in NW Mali in Nov 1999 (L. Granjon & B. Sicard unpubl.). Said to be much less abundant than syntopic Taterillus in N Senegal, with an estimate of 2–4 individuals/ha (at maximum) (Poulet 1984); in the same region, its abundance may vary inversely with that of other rodents, i.e. high during droughts and low after heavy rainfalls (Poulet 1978). Desmodilliscus braueri

Adaptations Terrestrial and nocturnal. Lives in a small, shallow, but complex burrow with up to 13 entrances, possibly inhabited by groups of individuals (N Senegal; Poulet, 1984). Foraging and Food No information.

Reproduction and Population Structure In captivity, reproduction observed only between Jun and Aug in Senegal (corresponding to the end of dry season and beginning of wet season). Gestation: 26 days. Litter-size: 2 and 3 (n = 8 litters).Weight at birth around 0.9 g. Weaned at 28 days, at a weight of 3–4 g (Poulet 1984).

Social and Reproductive Behaviour Most probably a social, or even colonial, species (Poulet 1984). 265

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Predators, Parasites and Diseases Locally preyed on extensively by Barn Owls Tyto alba (Heim de Balsac 1967b, Poulet 1984). Also preyed on by wild cats, foxes and snakes (e.g. Eryx mulleri; Poulet 1984). Conservation

IUCN Category: Least Concern.

Measurements Desmodilliscus braueri HB: 55.4 (41–74) mm, n = 29 T: 40.1 (33–49) mm, n = 29 HF: 14.1 (13.5–15) mm, n = 23

E: 8.6 (7–11) mm, n = 25 WT: 9.6 (6–14) g, n = 19 GLS: 22.1 (20.4–23.1) mm, n = 17 GWS: 12.8 (11.7–13.3) mm, n = 14 M1–M3: 3.2 (2.9–3.4) mm, n = 18 Auditory bulla: 9.0, 9.9 mm, n = 2* Burkina, Niger, Cameroon, Sudan (Hutterer & Dieterlen 1986) *BMNH Key References

Hutterer & Dieterlen 1986; Poulet 1984. L. Granjon

GENUS Desmodillus Cape Short-tailed Gerbil Desmodillus Thomas and Schwann, 1904. Abstr. Proc. Zool. Soc. Lond. 1904 (2): 6. Type species: Gerbillus auricularis Smith, 1834.

Desmodillus auricularis.

Monotypic genus widespread in the South-West Arid BZ. The genus is characterized by small size (mean HB 110 mm), short tail (mean 84 mm and shorter than head and body) without any pencil or tuft, and short ears. Skull characters include very large tympanic bullae that extend posteriorly to the occiput, short zygomatic plate, long slender incisors each with a shallow groove. M1 has three rows of cusps, M2 has two rows of cusps, M3 has single cusp (Figure 47). The form of the bullae is quite different to that of Gerbillurus and Gerbilliscus (formerly Tatera) (see Figure 44). Although larger than sympatric Gerbillurus, it is smaller than Gerbilliscus (formerly Tatera) in southern Africa. Fossil forms have been found back to the Pleistocene

Figure 47. Skull and mandible of Desmodillus auricularis (BMNH 25.1.2.76).

in South Africa (Avery 1998) and Namibia (Senut et al. 1992, Musser & Carleton 2005). The single species is Desmodillus auricularis. Jan A. J. Nel

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Desmodillus auricularis

Desmodillus auricularis CAPE SHORT-TAILED GERBIL (NAMAQUA GERBIL) Fr. Gerbille à queue courte; Ger. Kurzschwanz-Rennmaus Desmodillus auricularis (Smith, 1834). S. Afr. Quart. J., ser. 2, 2: 160. Kamiesberg, South Africa.

Taxonomy Originally described in the genus Gerbillus. Synonyms: brevicaudatus, caffer, hoeschi, pudicus, robertsi, shortridgei, wolfi. Subspecies: none. Chromosome number: 2n = 52. Description Small, stockily built gerbil with fine, soft, dense pelage. Dorsal pelage variable, ochrous-orange to tawny-brown; hairs slate-grey at base, ochrous-yellow in middle, with dark tip on some hairs. Ventral pelage, including chin and throat, pure white. Head large, with thickened nose, long black vibrissae and large eyes. Ears small, oval and flesh-coloured. Distinctive white postauricular patch; smaller less conspicuous supraorbital and suborbital patches. Fore- and hindlimbs short and thick, with short white hairs on upper surfaces of feet; four digits on forefeet, five on hindfeet; hindfeet with hairy soles. Tail of moderate length (ca. 75–80% of HB), same colour as dorsal pelage (with dark tip in darker coloured individuals); without pencil. Skull characterized by the greatly enlarged auditory bullae (ca. 41% of GLS, larger as percentage of GLS than in most other gerbillines). Nipples: 2 + 2 = 8. Females weigh on average 20% less than "". Geographic Variation

None recorded.

Similar Species Gerbillurus spp. Smaller mean head and body length; tail longer (absolute and as percentage of HB length); pencil at end of tail (in some species); smaller bullae (ca. 30% of GLS). Distribution Endemic to Africa. South-West Arid BZ (Kalahari and Namib Deserts, Karoo) and peripherally in the extreme south of the Zambezian Woodland and South-West Cape BZs. Widely distributed in Namibia (except NE), Botswana (except N and parts of E), arid parts of South Africa, and marginally in SW Angola. Recorded from near sea level to ca. 1600 m (extrapolated from distribution map in De Graaff 1981). Habitat Favours calcareous ground, fine soils or consolidated sand (sometimes covered in pebbles) with a sparse cover of grass or low shrub. In the southern parts of the Kalahari and Little Namaqualand, more common in calcareous ground or outcrops and fringes of pans. Avoids dense grassland or thick scrub. Abundance Common in suitable habitat. Comprises 4.95–5.5% of small mammals in SW Kalahari (Nel & Rautenbach 1975, J. A. J. Nel unpubl.), but with clear differences in abundance in different micro-habitats and at different times (Nel 1978). Large fluctuations in numbers rarely occur. Adaptations Nocturnal and terrestrial. Locomotion is nonsaltatorial. The greatly enlarged auditory bullae afford acute hearing and probably allow gliding owls to be located (Lay 1972). Burrows are extensive and complicated with 1–7 openings, blind alleys and storage chambers; they are 300–600 mm deep (sometimes deeper) and burrow diameter is ca. 53 mm (Nel 1967, Smithers 1971). At

Desmodillus auricularis

ambient temperatures of 11–30 °C, deep body temperature (Tb) can be kept nearly constant (varies by 0.5 °C in "", 0.1 °C in !!) (Nel & Rautenbach 1977). Renal concentrating ability is very good (urine concentration up to 6.1 mOsmol/kg), and evaporative water loss at low relative humidity is very low (Christian 1978, 1979b, Buffenstein et al. 1985). These gerbils are independent of free water, and this allows breeding at times when other sympatric species are reproductively inactive (Christian 1979b). They can also store fat in the tail when conditions are good; hence some older individuals are very large, perhaps being in their second year. Foraging and Food Omnivorous. Food includes seeds, annuals, seeds of wild melons, and insects. In winter, diet is mostly seeds and in summer it changes to insects and green leaves (in nearly equal amounts). Gerbils forage and feed up to 30 m from burrows. Food may be stored in burrows (‘larder hoarding’) and also other locations within the home-range (‘scatter-hoarding’). Social and Reproductive Behaviour Asocial and solitary, although burrows can be close together and linked with pathways. In captivity, !! are dominant over "" and may kill and consume them, as well as other !!. Males are tolerated by !!, for a short period only, at times of copulation. Reproduction and Population Structure Under favourable conditions, reproduction can occur throughout the year, but births are mostly during the hot wet season. Gestation: 21 days. Mean litter-size for !! caught in the wild: 2 (n = 5) in S Kalahari (Nel 267

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Family MURIDAE

& Stutterheim 1973) and 4 (n = 19) elsewhere (Keogh 1973). Mean weight of young at birth 1.84 g (n = 10) in the Kalahari and 4.4 g (n = 19) elsewhere. Nipple-clinging absent. Young naked and blind at birth. Ear pinnae free at Day 12. Eyes open Day 21. External auditory meatus open Day 23. Crawling starts Day 2; walking Day 15; grooming Day 24; digging and sand-bathing Day 30. Sucking ceases at Day 33 (Nel & Stutterheim 1973). In SW Kalahari, young enter the population towards the end of the wet season (Feb–Apr). Populations peak in mid-winter (Jun–Aug) and decline thereafter. Predators, Parasites and Diseases Main predators include Barn Owls Tyto alba, Spotted Eagle-owls Bubo africanus and snakes (e.g. cobra, Naja nivea). Ectoparasites include 24 species of fleas (many involved in transmitting plague to man), mites and ticks (details in De Graaff 1981). Dwarf Gerbils are vectors of bubonic plague, and in the laboratory can become infected with listeriosis, louse typhus, murine or rat typhus and tick-bite fever (De Graaff 1981). Conservation

Measurements Desmodillus auricularis HB: 110.4 (86–129) mm, n = 64 T: 84.8 (70–98) mm, n = 64 HF (c.u): 25.3 (21–29) mm, n = 62 E: 11.6 (10–14) mm, n = 62 WT: 46.1 (29–82) g, n = 71 GLS: 35.7 (34.2–38.1) mm, n = 10 GWS: 19.7 (18.7–22.3) mm, n = 10 M1–M3: 4.95 (4.6–5.6) mm, n = 10 Auditory bulla: 14.5 (13.1–15.5) mm, n = 10 Body measurements and weight: south-western Kalahari (J. A. J. Nel unpubl.) Skull measurements: Namibia (C. G. Coetzee unpubl.) Auditory bulla measurements: Namibia (BMNH) The range of weights probably results from subadults being included, as sampling took place during all seasons Key References De Graaff 1981; Nel 1978; Nel & Rautenbach 1975; Skinner & Smithers 1990.

IUCN Category: Least Concern.

Jan A. J. Nel

GENUS Gerbilliscus Gerbils Gerbilliscus Thomas, 1897. Proc. Zool. Soc. Lond. 1897: 433. Type species: Tatera boehmi (Noack, 1897).

The genus Gerbilliscus comprises 12 species, which are widely distributed in all sub-Saharan Africa with the exception of the Rainforest BZ, and they occupy a variety of habitats in both northern and southern savannas. Gerbilliscus leucogaster, G. brantsii and

Figure 48. Skull and mandible of Gerbilliscus kempi (HC 1321).

G. validus are widespread and common in southern and eastern Africa; other species have less extensive distributions (G. afra, G. gambiana) and one species is very restricted (G. phillipsi). Gerbilliscus, previously considered as a subgenus of Tatera, is now elevated to genus rank, to account for differences between true Tatera (represented by the sole Asian species T. indica) and the African species (Pavlinov et al. 1990, Pavlinov 2001, G. Musser pers. comm.). These differences are mainly in dental pattern and mastoid bone structure (more derived in Gerbilliscus than in Tatera), but also include humerus morphology and diploid number of chromosomes (2n = 68 in Tatera, 2n = 36 to 52 in Gerbilliscus). The species of Gerbilliscus are on average more powerfully built than species from the other genera of Gerbilline rodents; they generally have a darker pelage, although there is considerable geographic variation in colour of the dorsal pelage. Populations in arid to semiarid habitats have paler colouration than populations in moister habitats. They have comparatively long hindfeet, and the soles are naked. The skull is robustly built, and the molar row is larger than in Taterillus. The auditory bullae are inflated anteriorly with a small posterior section, in contrast to Gerbillurus and Desmodillus in which the posterior section is also inflated. Posterior palatal foramina are short (no longer than 3 mm, and considerably shorter than in Taterillus), a feature considered diagnostic among the Gerbillinae by Davis (1975a) (Figure 48). The species of Gerbilliscus are physiologically, morphologically and behaviourally adapted to living in dry environments. They are terrestrial and nocturnal, spending the daytime in burrows that can be complex and deep. They are predominantly granivorous, but may

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Table 23. Species in the genus Gerbilliscus. Arranged in order of increasing mean head and body length. All measurements in mm. (n. d. = no data.) Species

G. leucogaster

G. brantsii

G. afra

G. phillipsi

G. guineae

G. robustus

G. inclusus

G. kempi

G. boehmi

G. validus

G. gambianus

G. nigricaudus

Tail colour Dark brown above, white below; dark pencil Proximal half brown; distal half white above, white below Reddish-brown; evenly coloured throughout Pale orange-brown above, white below; some black hairs above towards terminal end; without pencil Dark above, white below; small black terminal tuft Mostly brown above, usually pale or brown below; some individuals with black at tip or along most of length Dark above, white below; occasionally with white tip Dark above, white below; small dark pencil Upper half/twothirds dark brown, terminal half white; with pencil Brown above, pale below; without terminal pencil Dark above, orange-brown on sides, white below; without pencil Black above, black below; white hairs below near base of tail in some individuals

HB mean (range) (mm)

T % of HB

GLS mean (range) (mm)

Upper incisor teeth

Chromosome number

Notes

128.6 (89.0–155)

115%

37.3 (33.3–40.5)

Opisthodont; single groove

2n = 40, FN = 66

Eastern Africa; Tanzania to N South Africa

134.6 (96.0–164)

106%

38.7 (35.6–42.2)

n. d.

2n = 44, FN = 66

Angola, Namibia, Zimbabwe

141.3 (124–157)

110%

39.1 (34.3–42.0)

Opisthodont; single groove

2n = 44, FN = 66

SW South Africa only

144.0 (143–145)

130%

38.9 (37.3–40.9)

Opisthodont; single groove

n. d.

149.8 (128–178)

110–140%

36.2 (33.1–38.7)

Opisthodont; single groove

2n = 50, FN = 64

Senegal to Togo

152.2 (120–190)

115%

41.9 (39.0–44.7)

Opisthodont; single welldefined groove

2n = 40, FN = 70

NE Africa

156.0 (152–161)

115%

41.1 (36.7–44.3)

Opisthodont; deep single groove

n. d.

Tanzania, Mozambique, Zimbabwe

158 (140–190)

100%

40.2 (38.7–41.6)

Opisthodont; single groove

2n = 36

Senegal to Cameroon

162.3 (139–179)

130%

43.5 (42.0–45. 2)

Opisthodont; two faint grooves

n. d.

East-central Africa

n. d.

Angola, S and E DR Congo, Zambia, Tanzania, Uganda, Sudan and W Ethiopia.

N Kenya, Ethiopia

167 (135–195)

95%

41.7 (38.5–44.7)

Opisthodont; usually smooth without groove

168.2 (148–196)

80–100%

37.2 (34.0–40.7)

Opisthodont; single groove

2n = 52, FN = 64

Senegal, Mali, Niger, Chad

185.8 (178–193)

110%

48.7 (47.0–50.5)

Opisthodont; single welldefined groove

n. d.

NE Africa

eat a variety of food including insects, according to availability. They can be locally abundant and, in agricultural areas, may cause damage to crops. They are important vectors of fleas that carry plague.

The taxonomy of the genus is uncertain, because important intraspecific morphological variation masks the distinction between species. One species, G. boehmi, has been placed in the subgenus 269

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Gerbilliscus gerbils.

Gerbilliscus on account of its double-grooved incisors and fringed, white-tipped tail; all the other African species are included in the subgenus Taterona. In the subgenus Taterona, the species have sometimes been distributed between an ‘afra’ and a ‘robusta’ group, the systematic status of which is questionable. Gerbilliscus afra, G. brantsii and G. inclusus are placed in the ‘afra’ group, and are distinguished from the ‘robusta’ group on the grounds of the quality of pelage, and morphological and craniological characters; however, there is some overlap between these subgeneric characters. Chromosomal and molecular data support the close relationship between G. afra and G. brantsii, but (on present evidence) do not support a monophyletic ‘robusta’ group.

Criteria used to distinguish between species include morphological and biometrical characters (colour, overall size, tail length, presence and size of terminal pencil on the tail, skull size and characters), as well as differences in karyotypes. Considerable geographic variation within species, and overlap in distinguishing characters between species makes precise identification difficult in some cases. Geographic locality aids identification, but two or more species of Gerbilliscus may occur sympatrically and syntopically in parts of the distribution range. Twelve species recognized: G. afra, G. boehmi, G. brantsii, G. gambianus, G. guineae, G. inclusus, G. kempi, G. leucogaster, G. nigricaudus, G. phillipsi, G. robusta and G. validus (Table 23). L. Granjon & Edith R. Dempster

Gerbilliscus afra CAPE GERBIL Fr. Gerbille du Cap; Ger. Kap-Nacktsohlen-Rennmaus Gerbilliscus afra (Gray, 1830). Spicilegia Zool. 2: 10. Cape of Good Hope, South Africa.

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). Synonyms: africanus, caffer, gilli, schlegelii.Subspecies: none. Chromosome number: 2n = 44, FN = 66 (as T. afra); karyotype identical to that of G. brantsii (Qumsiyeh 1986). Description Medium-sized gerbil with white underparts, darker dorsal surface and long tail. Dorsal pelage reddish-orange or pale buffy, faintly grizzled with dark brown; hairs dull lavender at base, reddish-orange or pale buff at tip. Hairs fairly long and broad, with chevron scale pattern. Flanks similar to dorsal pelage. Ventral pelage and inside of limbs white, with clear delineation between colour of flanks and ventral pelage. Head narrow, with pointed nose, long vibrissae, sides of muzzle white. Large eyes. Ears elongated, pale flesh colour inside, dark brown outside, rounded at tips. Outer surface of limbs reddish-orange; hindlimbs much longer than forelimbs; hindfeet elongated. Fore- and hindfeet white, five digits each, Digit 5 on forefeet reduced. Tail long (ca. 110% of HB),

covered with dense short hairs, same colour or slightly paler than dorsal pelage and coloured evenly to tip. Nipples: 2 + 2 = 8, but considerable variation. Front face of each upper incisor with groove, lower incisors ungrooved; molar teeth broader and heavier than G. leucogaster. Auditory bullae not particularly enlarged. Males on average larger than !!. Geographic Variation

None recorded.

Similar Species G. brantsii. Ventral pelage white; tip of tail white; more widespread; allopatric. G. leucogaster. Molar teeth narrower; auditory bullae not markedly inflated; allopatric. Distribution Endemic to Africa. South-West Cape BZ. Recorded in Western and Northern Cape Provinces of South Africa, from

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Gerbilliscus afra

Niewoudtville in Northern Cape southwards to Cape Peninsula and eastwards coastally to Herold’s Bay. Habitat Confined to areas of loose, sandy soils or sandy alluvium. Common in cultivated lands. Abundance Common in suitable habitats. Adaptations Nocturnal and terrestrial. Moves by quadrupedal saltation. Excavates extensive burrows in sandy places. Numerous interconnecting tunnels end in a chamber containing nest of shredded vegetation. Body temperature maintained at 34–36 °C at Ta of 10–30 °C, but susceptible to hyperthermia at Ta above 30 °C. No significant drop in Tb at 5 °C, indicating good tolerance of low temperatures. Water turnover rate higher than other species of southern African gerbils and probably associated with living in a mesic environment. Basal metabolic rate higher than average for gerbilline rodents, a condition that is probably related to the mesic environment, herbivory and tolerance of a relatively low ambient temperature (Duxbury & Perrin 1992). Foraging and Food Herbivorous, and occasionally insectivorous. Cape Gerbils eat grass, bulbs, roots and seeds. Captive animals also eat insects. Social and Reproductive Behaviour Social structure unknown. Adults rarely aggressive in laboratory encounters (Dempster et al. 1993). Copulation consists of series of mounts with and without intromission, culminating in intromission with ejaculation. No lock; copulatory plug deposited after ejaculation. Several bouts of mounting with and without ejaculation occur over about one hour. Vocalizes audibly with calls of 1.6–5.3 kHz, with second to sixth harmonics. Ultrasonic whistles emitted in frequency range 20–34 kHz. Whistles are short (duration 225 msec) (Dempster & Perrin 1994). Reproduction and Population Structure In SW Cape Province, South Africa, !! have a breeding season of eight months (Aug to end Mar) followed by four months of anoestrus when there are no pregnancies and during which none of the young !! reach puberty (Measroch 1953). Males cease spermatogenesis and !! are anoestrus during the cool wet season. Breeding strategy is similar to other gerbil species: short gestation, large litter-size, altricial young, iteroparous. Embryo number: 4.0 (2–6). Litter-size: 4 (range 3–5). Mean weight at birth: 4.1 g. Growth rate in first 28 days: 0.8 g/day. Pups cling to nipples from 1 to 4 days old. Incisors erupt Day 10. Dorsal pelage visible Day 6–8. Eyes open Day 18–21.Weaned by Day 22–28. Females experience a postpartum oestrus (Dempster et al. 1992). The population is mostly subadults Jan–Mar, mostly adults from Jul– Nov. Males and !! may survive to a second breeding season, with an estimated life-span of 12–17 months. During the breeding season, 60% of !! are pregnant. Females in the wild may have six to seven litters in one year (Measroch 1953). Testes in adult "" are unusually large, comprising over 8% of adult body weight (Allanson 1958). Predators, Parasites and Diseases No information available on predators. Susceptible to infections of Mycobacterium tuberculosis, louse

Gerbilliscus afra

typhus caused by Rickettsia prowazekii, rat typhus caused by R. typhi, and tick-bite fever by Rickettsia conorii. Ectoparasites include mites of the families Laelaptidae (9 spp.), Myobiidae (1 sp.), Trombiculidae (2 spp.) and Listrophoridae (1 sp.); and fleas of the families Pulicidae (9 spp.), Hystricopsyllidae (1 sp.) and Chimaeropsyllidae (1 sp.). Like G. brantsii, afflicted by Yersinia pestis, which may lead to local outbreaks of bubonic plague (details in De Graaff 1981). Conservation IUCN Category: Least Concern. Although formerly listed in the South African Red Data Books, Smithers (1986b) recommended that it should be removed because it is not uncommon, adapts well to changing land use, and population numbers do not seem to have declined. Measurements Gerbilliscus afra HB: 141.3 (124–157) mm, n = 44 T: 152.2 (133–175) mm, n = 44 HF: 37.5 (28–44) mm, n = 44 E: 24.5 (20–28) mm, n = 44 WT: 95.1 (78–113) g, n = 15 GLS: 39.1 (34.3–42.0) mm, n = 7 GWS: 19.9 (18.2–21.6) mm, n = 7 M1–M3: 6.6 (6.2–7.0) mm, n = 5 Auditory bulla: 9.7 (9.2–10.3) mm, n = 5 Body measurements and weight: Western Cape (De Graaff 1981, as Tatera afra) Skull measurements: Western Cape (P. J. Taylor unpubl.) Key References Dempster & Perrin 1994; Dempster et al. 1992; Duxbury & Perrin 1992. Edith R. Dempster 271

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Gerbilliscus boehmi BOEHM’S GERBIL Fr. Gerbille de Boehm; Ger. Boehms Nacktsohlen-Rennmaus Gerbilliscus boehmi (Noack, 1887). Zool. Jahrb. Syst., 2: 241. Qua Mpala, Marungu, S DR Congo. This locality has been stated to be in N Zambia (see Ansell 1978, Bates 1988).

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). Synonyms: fallax, fraterculus, varia. Subspecies: none. Chromosome number: not known. Description Large dark-coloured gerbil with very long whitetipped tail. Dorsal pelage medium brown, flecked with black and ochre, darker on midline than on flanks; hairs medium grey at base, with black tip (mid-dorsally) or ochre tip (flanks). Ventral pelage white; sharp delineation between colour of flanks and ventral pelage. Hairs of cheeks and shoulders often tipped with cinnamon. Forehead and nasal region dark brownish-black. Eyes large. Ears large, rounded, with short black hairs. Chin, inner surface of limbs white. Soles of hindfeet naked and darkly pigmented. Tail very long (ca. 130% of HB), thin; proximal half or two-thirds with short hairs, dark above, white below; terminal half or third pure white above and below, often with small pencil of white hairs. Skull: large and deep; incisors orthodont, sometimes with two faint longitudinal grooves on each upper incisor tooth; cheekteeth broad, laminate and relatively long. Nipples: 2 + 1 = 6 or 2 + 2 = 8. Geographic Variation None recorded. Gerbilliscus boehmi

Similar Species Gerbilliscus leucogaster, G. validus, G. robustus, G. nigricaudus. All have shorter tails (actual and relative to HB) without white tip. Distribution Endemic to Africa. Zambezian Woodland BZ, especially northern part. Widespread in savanna woodlands (mostly at 1000–2000 m) from SW Kenya and S Uganda to N Mozambique, Malawi, S DR Congo and W Zambia. Probably also present in Moxico Province of E Angola. Not known from east of L. Malawi or from the lowland areas of the Luangwa Valley in Zambia.

Social and Reproductive Behaviour Mostly unknown. May forage over a large area (Vesey-Fitzgerald 1966). In some localities, lives parapatrically with G. validus (Tanzania; Kingdon 1974), G. leucogaster (Malawi: Hanney 1965), and with Lemniscomys sp. and sengis (Kagera N. P., Rwanda; Misonne 1965a). Reproduction and Population Structure Breeding recorded in early wet season (Nov) and end of wet season (May). Lactating !! in Nov (Zambia) and May (Malawi); pregnant ! in May (Rwanda). Embryo number: 5 (n = 1; Hanney 1965, Misonne 1965).

Habitat Brachystegia woodland, mostly at higher altitudes, where there is good cover of grass and herbaceous plants; also in grassy Predators, Parasites and Diseases Preyed upon by owls at plains (Uganda) and ‘bush’ habitats. Prefers moister habitats (cf. G. several locations in Kagera N. P., Rwanda (Misonne 1965) and at leucogaster). Individuals found in old banana plantations (Misonne Dedza, Malawi (Hanney 1965). 1965a) and old millet fields (Kingdon 1974) in some parts of range. Conservation IUCN Category: Least Concern. Abundance Although widespread, mostly uncommon or rare. Reasons for rarity not known. Measurements Gerbilliscus boehmi Adaptations Nocturnal. Burrows have one or two entrances and HB: 162.3 (139–179) mm, n = 12 are not marked by a pile of excavated soil. Sometimes use burrows of T: 215.5 (190–234) mm, n = 12 other Gerbilliscus spp., or those of mole-rats (Cryptomys spp.) (Vesey- HF: 40.8 (38–47) mm, n = 11 Fitzgerald 1966). Habitat and altitudinal range suggest that moderate E: 24.3 (21–26) mm, n = 12 climatic temperatures are required for survival. WT: 146 g, n = 1 GLS: 43.5 (42.0–45.2) mm, n = 10 Foraging and Food Omnivorous, primarily vegetable material GWS: 23.3 (22.0–24.3) mm, n = 8 and insects (Hanney 1965, Misonne 1965a). M1–M3 (alveolar): 7.4 (6.8–7.8) mm, n = 15 272

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Gerbilliscus brantsii

Auditory bulla: 12.9 (12.0–14.5) mm, n = 14 Throughout geographic range (as Tatera boehmi; Bates 1988) Females only (limited data suggest that "" are slightly larger)

Key References

Bates 1988; Vesey-Fitzgerald 1966. D. C. D. Happold

Gerbilliscus brantsii HIGHVELD GERBIL Fr. Gerbille du Veld; Ger. Brants Nacktsohlen-Rennmaus Gerbilliscus brantsii (Smith, 1836). Rept. Exped. Exploring Central Africa, p. 43. Ladybrand, E Free State, South Africa. ‘Tops of hills near sources of Caledon River’ near Lesotho border (see Meester et al. 1986).

Taxonomy Originally desribed in the genus Tatera (see profile Genus Gerbilliscus). Synonyms: breyeri, draco, griquae, humpatensis, joanae, maccalinus, maputa, miliaria, montanus, namaquensis, natalensis, perpallida, ruddi, tongensis. Subspecies: three. Chromosome number: 2n = 44, FN = 66 (as Tatera brantsii); karyotype identical to that of G. afra (Qumsiyeh 1986). Description Medium-sized gerbil with darker dorsal surface, pale underparts and long tail. Dorsal pelage pale rufous-brown to pale reddish, with faint, uneven brown wash. Pelage soft and fluffy; hairs long and broad with narrow base and typical chevron pattern. Ventral pelage pure white to buffy-grey. Head narrow, with pointed nose and long vibrissae. Large eyes. Chin white. Ears elongated, dark brown, rounded at tips. Hindlimbs much longer than forelimbs, hindfeet long. Fore- and hindfeet pale, five digits each, Digit 5 on forefeet reduced.Tail long (ca. 106% of HB, shorter than HB in some individuals), similar in colour to dorsal pelage, or slightly darker for at least the proximal half; distal half white; white below. Auditory bullae well-developed. Nipples: 1 + 2 = 6 or 2 + 2 = 8. Geographic Variation Dorsal pelage varies from pale in C Botswana, pale or distinctly reddish in south-western parts of distribution, to darker in south-east parts of distribution. Meester et al. (1986) recognize three subspecies: G. b. brantsii: Lesotho, Eastern Cape and KwaZulu–Natal Provinces, South Africa and westwards to edge of Kalahari Desert. Buffygrey patches on chest; heavier molars. G. b. griquae: Kalahari northwards to S Angola and W Zambia. Pure white ventral pelage; narrower molars; pale dorsal pelage. G. b. ruddi: N KwaZulu–Natal Province, South Africa. Buffy-grey ventral pelage; relatively long white-tipped tail; long hindfoot. Similar Species G. leucogaster. Brighter, sleeker fur; sharp line of delineation between flanks and ventral pelage; tail has distinct dark line on dorsal surface, never white-tipped. G. afra.Ventral pelage pure white, tail evenly coloured to tip; confined to Western Cape Province, South Africa. Distribution Endemic to Africa. South-West Arid and Highveld BZs, with marginal extension to southern Zambezian Woodland BZ. Recorded from S Angola,W Zambia, Botswana, E Namibia and South Africa. Limited distribution in N Zimbabwe and C Mozambique. Absent from extremely arid parts of W Namibia and W South Africa.

Gerbilliscus brantsii

Habitat Associated with sandy soils and sandy alluvium, with some cover of grass, scrub or open woodland. Also found in peaty soils around marshes and pans, sometimes using tunnels of mole-rats (Cryptomys spp.). Not normally found on heavy consolidated soils or very loose sandy soils. Abundance Common. In N Transvaal Province, South Africa, Brant’s Gerbils were the commonest species in the dry season (May– Jun): in ‘old field’, density was 16/ha and biomass was 1280 g/ha (ca. 80% of total numbers and biomass; n = 5 spp. of small terrestrial rodents) and in Burkea woodland, density was 12/ha and biomass was 960 g/ha (ca. 75% of total, n = 6 spp.) (Korn 1987). In southern African highlands, estimated density was 14.8/ha (Feb) and 27.1/ha (May) (De Moor 1969). Adaptations Highveld Gerbils are strictly nocturnal with crepuscular peaks of activity. Move by quadrupedal saltation. They excavate complex burrows in loose, sandy soil. A burrow has many entrances, and tunnels (ca. 45–60 mm in diameter) that interconnect underground. Maximum depth of burrows about 20 cm; total length of burrow systems about 6 m. There is one nesting chamber, sealed with loose sand, in each burrow system. 273

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Family MURIDAE

Foraging and Food Highveld Gerbils eat mostly plants, including roots and green parts, with insects comprising about 5% of diet in southern Kalahari. More green than white plant material is eaten in hot season, depending on rainfall (Nel 1978). Gerbils may have significant effects on local vegetation: plant biomass, root biomass and vegetation height is significantly lower, and species diversity and evenness of plants significantly higher, near old colonies than in adjacent, undisturbed areas. Highveld Gerbils maintain high plant diversity in savanna habitats (Korn & Korn 1989). Gut morphology is simple and unspecialized suggesting a general opportunistic diet (Perrin & Curtis 1980).

Predators, Parasites and Diseases Preyed on by snakes, small mammalian carnivores, Barn Owls Tyto alba and African Grassowls Tyto capensis. Wild populations sometimes afflicted by Yersinia pestis, leading to local outbreaks of bubonic plague. Susceptible to infection by Pseudomonas pseudomallei, Listeria monocytogenes and Mycobacterium tuberculosis in the laboratory. Endoparasites include the cestodes Hymenolepis microcantha, H. taterae and Raillietina trapezoides. Ectoparasites include 29 species of fleas and nine species of ticks (details in De Graaff 1981). Widely used as a laboratory animal for medical research. Conservation

Social and Reproductive Behaviour Colonial, with several individuals living in close proximity to others. Adults are rarely aggressive towards each other in laboratory encounters. If aggression does occur, both animals stand up on hindfeet, and strike at each other with their forepaws. Copulation consists of a series of mounts with and without intromission, culminating in intromission with ejaculation. No lock; copulatory plug deposited after ejaculation. Multiple copulations and ejaculations occur over about an hour. Vocalizations include two kinds of ultrasonic whistles: short whistles of 17–27 kHz lasting about 157 msec, and long whistles of 17–31 kHz, lasting about 480 msec (Dempster & Perrin 1991b). Reproduction and Population Structure Pregnant !! recorded throughout year, with peak of reproductive activity at onset of cool dry season (Gauteng Province, South Africa). Males show active spermatogenesis throughout year, with peak in warm wet season. Breeding strategy similar to that of other gerbils: short gestation, altricial young, iteroparous, but small litter-size. Gestation: 22 days. Embryo number: 2.8 (1–5). Mean litter-size: 2.8. Mean weight at birth: 4.6 g. Growth rate in first 28 days: 1.1 g/ day. Pups cling to nipples from Day 1–4. Incisors erupt Day 6. Eyes open Day 16–20. Weaned by Day 28 (Scott 1979). Postpartum oestrus indicated by 40% of !! in the wild being both lactating and pregnant. Females may have five or six litters/year (Measroch 1953). Testes in adult "" are large, comprising ca. 5% of adult body weight (Allanson 1958; cf. G. afra).

IUCN Category: Least Concern.

Measurements Gerbilliscus brantsii HB: 134.6 (96–164) mm, n = 237* T: 143.1 (103–186) mm, n = 237 HF: 35.0 (19–47) mm, n = 237 E: 21.5 (12–34) mm, n = 236 WT: 79.9 (25–126) g, n = 130 GLS: 38.7 (35.6–42.2) mm, n = 15 GWS: 21.6 (19.0–23.0) mm, n = 14 M1–M3: 7.2 (6.7–7.9) mm, n = 15 Auditory bulla: 10.6 (9.6–11.7) mm, n = 13** Body measurements and weight: Transvaal (Rautenbach 1978; recalculated – "" and !! combined; as Tatera brantsii) Skull measurements: Drakensberg Mts, KwaZulu–Natal Province, South Africa (P. J. Taylor unpubl.) *Recalculated; original data as total length **P. J. Taylor unpubl. Mean weight of "" in southern Kalahari is lower than in Transvaal: 64.9 g, n = 67 (Nel & Rautenbach 1975) Key References Measroch 1953.

Dempster & Perrin 1991b; Korn & Korn 1989; Edith R. Dempster

Gerbilliscus gambianus GAMBIAN GERBIL Fr. Gerbille de Gambie; Ger. Gambische Nacktsohlen-Rennmaus Gerbilliscus gambianus (Thomas, 1910). Ann. Mag. Nat. Hist., ser. 8, 6: 428. Marakissa, Upper Gambia, Senegal.

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). Often considered to be a synonym of G. kempi (e.g. Rosevear 1969, Musser & Carleton 1993, 2005) or G. validus kempi (Bates 1988). Re-established as a distinct species by Hubert et al. (1973) following Matthey & Petter (1970). Distinguished by karyotype from other Gerbilliscus. Synonyms: hopkinsoni. (The form hopkinsoni was described later from the same region as gambianus [Gambia R], but this name was given improperly by various authors to specimens with 2n = 48, a karyotype never found in this region.) Subspecies: none. Chromosome number: 2n = 52, aFN = 64 (Matthey & Petter 1970; as T. gambianus).

Description Large robust gerbil. Dorsal pelage grey-brown; hairs dark grey at base, with wide brown or orange central zone, black at tip. Flanks paler; hairs without black tip.Ventral pelage white, clearly delineated from colour of flanks. Head rounded, similar in colour to flanks. Moderately pointed nose. Large eyes; moderately elongated ears. Chin, throat, chest and inner sides of limbs white. Hindfeet white above, dark below; forefeet entirely white. Tail moderately long (ca. 80–100% of HB), well haired, dark above, orange to brown on sides, white below, without any marked pencil of hairs at terminal end. Nipples: 2 + 2 = 8.

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Gerbilliscus gambianus

Geographic Variation None recorded. Similar Species G. kempi. Ear on average longer; chromosome number: 2n = 48. More southern distribution; other species of this short-tailed group occur only in eastern and southern Africa. G. guineae. Tail on average longer (110–140% of HB) with pencil. 2n = 50. Similar distribution. Distribution Endemic to Africa. Sahel Savanna and Sudan Savanna BZs of Senegal (Hubert et al. 1973) and Mali (B. Sicard & L. Granjon unpubl.). Recent captures in SE Niger (Kojimairi, 40 km south of Goudoumaria; Dobigny et al. 2002b) and at the southern edge of L. Chad (Granjon & Dobigny 2003) suggest a much larger range. Habitat In Senegal, found in Combretum woodlands (where sympatric with G. guineae), but on soils that are more sandy. Also trapped in fallow lands and traditionally cultivated fields (Hubert 1977, Hubert et al. 1977). The only species of gerbil found on sandy and mangrove islands in the Saloum Delta, Senegal (Granjon & Duplantier 1989). Gerbilliscus gambianus

Abundance Very varied according to habitat. Density up to 15/ha in sahelo-sudanian woodland at Bandia, Senegal, where may represent up to 17% of the rodents in favourable habitats (Hubert 1977). In the Saloum Delta, abundance varied from 3.5 to 24 individuals/ha on a 2 ha study area during a two-year survey (Granjon et al. 1994). This species seems to show more pronounced variations in abundance than G. guineae in mainland Senegal. Adaptations Terrestrial and nocturnal. Digs complex and moderately deep burrows (average 30 cm), with many entrances. Up to ten burrows can be found within the home-range of one individual (Hubert et al. 1977). Poor swimming abilities (Duplantier & Bâ 2001). Foraging and Food Consumes seeds, but also large amounts of insects especially at the beginning of the dry season (Moro & Hubert 1983). Social and Reproductive Behaviour Little information. Home-range 600–800 m2. Mean successive recapture distance 12.5 m (max 21.5 m) during a period of 10–12 days; recapture distances smaller than for sympatric G. guineae in W Senegal. Considerable overlap in home-ranges between individuals of both sexes. Does not show agonistic or amicable behaviour towards other species (Hubert 1977). Reproduction and Population Structure In W Senegal, pregnant !! recorded at end of wet season and first half of dry season (Sep–Feb) but no pregnancies recorded during second half of dry season and beginning of wet season (Mar–Aug). Gestation: 25 days. Litter-size: 2–6. Sexual maturity: 11–15 weeks (Hubert &

Adam 1975). Monthly mortality rate up to 50% when population is decreasing (Hubert 1977). Predators, Parasites and Diseases Remains found in pellets of Barn Owls Tyto alba from various localities in W Senegal (J.-M. Duplantier & L. Granjon unpubl.). Protozoan parasites include Leishmania major (Dedet et al. 1981) and Borrelia crocidurae (Trape et al. 1991). Arboviruses include Touré, Keuraliba, Gabek Forest and Koutango strains (Annual Reports of Pasteur Institute, Dakar). Conservation

IUCN Category: Least Concern.

Measurements Gerbilliscus gambianus HB: 168.2 (148–196) mm, n = 24 T: 149.2 (130–175) mm, n = 24 HF: 33.2 (31–35) mm, n = 25 E: 18 (17–20) mm, n = 24 WT: 94.1 (66–140) g, n = 25 GLS: 37.2 (34.0–40.7) mm, n = 24 GWS: 17.9 (16.0–19.6) mm, n = 21 M1–M3: 5.9 (5.0–6.3) mm, n = 24 Auditory bulla: n. d. Senegal (Diambour, Terres Neuves Region; MNHN) Key References 1969.

Hubert 1977; Hubert et al. 1977; Rosevear J.-M. Duplantier & L. Granjon

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Gerbilliscus guineae GUINEA GERBIL Fr. Gerbille de Guinée; Ger. Guinea-Nacktsohlen-Rennmaus Gerbilliscus guineae (Thomas, 1910). Ann. Mag. Nat. Hist., ser. 8, 5: 351. Gunnal, Guinea-Bissau.

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). Referred by Davis (1975a) to T. robusta (now Gerbilliscus robustus), but generally considered as a valid species (Rosevear 1969, Bates 1985). Synonyms: picta. Subspecies: none. Chromosome number: 2n = 50, aFN = 64 (Matthey & Petter 1970, as T. guineae). Description Medium to large-sized robust rodent. Dorsal pelage grey-brown; hairs dark grey at base, brown to orange central zone, and usually with short black tip. Flanks and head paler; hairs mostly without black tip. Ventral pelage and inner sides of limbs white; ventral colour clearly delineated on lower flanks. Chin, throat and chest white. Head rounded with moderately pointed nose. Large eyes; relatively elongated ears. Hindfeet white above, dark below; forefeet entirely white. Tail relatively long (110–140% of HB), well haired, dark above, white below, with marked pencil of darkish hairs at terminal end. Nipples: 2 + 2 = 8. Geographic Variation None recorded. Similar Species G. kempi.Tail relatively shorter without terminal pencil; chromosome Gerbilliscus guineae number: 2n = 48; similar distribution. G. gambianus.Tail relatively shorter without terminal pencil; 2n = 52; overlapping distribution (in the northern part of the range of G. 1500 m2. Mean recapture distance 21 m (max 38 m during 10–12 days); larger than for sympatric G. gambianus in W. Senegal (Hubert 1977). guineae). Distribution Endemic to Africa. Sudan and Guinea Savanna BZs, and Northern Rainforest–Savanna Mosaic. Recorded from Senegal (Hubert et al. 1973), Guinea-Bissau and Guinea (Ziegler et al. 2002), Burkina (Matthey & Petter 1970), Sierra Leone (Grubb et al. 1998), Côte d’Ivoire (Gautun & Petter 1972), Ghana (Rosevear 1969) and Togo (Robbins & Van der Straeten 1996). Not recorded from Gambia (Grubb et al. 1998).

Reproduction and Population Structure In a monthly survey from Jun 1971 to Mar 1973 in W Senegal (Hubert 1977), pregnant !! were found mainly during the wet season (Aug–Oct), but reproduction continued until the middle of the dry season (Jan– Feb) in 1972. Embryo number: 4–5. Monthly mortality rate up to 30%, less variable than in G. gambianus. Predators, Parasites and Diseases No information.

Habitat In Senegal, mostly found in Combretum woodlands on lateritic to clay, hydromorphous soils where there is a dense shrub layer and variable herbaceous cover (Hubert et al. 1977). Also trapped in cultivated areas (Senegal, Hubert et al. 1977; Mali, B. Sicard unpubl.) and on bare ironstone hills (Gambia; Rosevear 1969). Abundance Little information. Uncommon in W Senegal. Density of ca. 4/ha in sahelo-sudanian woodland at Bandia, Senegal; comprised 7.7–21% of rodents in favourable habitats (Hubert 1977). Adaptations Terrestrial and nocturnal. Digs relatively deep (average 50 cm) and moderately complex burrows in heavy soils. Entrances are hidden by heaps of excavated earth (Hubert et al. 1977).

Conservation

IUCN Category: Least Concern.

Measurements Gerbilliscus guineae HB: 149.8 (128–178) mm, n = 35 T: 175.3 (156–198) mm, n = 32 HF: 34.9 (32–37) mm, n = 34 E: 20.5 (19–22) mm, n = 34 WT: 73.7 (45–110) g, n = 35 GLS: 36.2 (33.1–38.7) mm, n = 20 GWS: 16.9 (15.4–18.4) mm, n = 18 M1–M3: 5.7 (4.8–6.1) mm, n = 20 Senegal (Diambour, Terres Neuves Region; MNHN)

Foraging and Food No information. Key References Hubert 1977; Hubert et al. 1977; Rosevear 1969. Social and Reproductive Behaviour Home-range 1400– L. Granjon & J.-M. Duplantier 276

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Gerbilliscus inclusus

Gerbilliscus inclusus GORONGOZA GERBIL Fr. Gerbille de Gorongoza; Ger. Gorongoza-Nacktsohlen-Rennmaus Gerbilliscus inclusus Thomas and Wroughton, 1908. Proc. Zool. Soc. Lond. 1908: 169. Tambarara, Mozambique.

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). Synonyms: cosensi, pringlei. Subspecies: three. The form pringlei was described as a valid species (Hubbard 1970b), treated as a subspecies here and by Davis (1975a), and as a synonym by Musser & Carleton (1993, 2005). Chromosome number: not known. Description Large gerbil with dark dorsal pelage, long hindfeet and long tail. Dorsal pelage ochraceous-buff, washed with black; hairs dark slate, ringed with ochraceous and tipped with black. Flanks paler than dorsal pelage. Ventral pelage white from chin to tail tip. Colour varies geographically (see below). Head narrow, with pointed nose, long vibrissae, sides of muzzle darker than head. Large eyes with black mark under each eye. Ears pinkish, almost naked, relatively short. Hindlimbs longer than forelimbs, hindfeet elongated. Forefeet and hindfeet off-white, five digits each, Digit 5 on forefoot reduced. Tail long (ca. 115% of HB), dark above, white below, occasionally white-tipped. Upper incisors deeply grooved, lower incisors ungrooved. Nipples: 1 + 2 = 6 or 2 + 2 = 8. Geographic Variation Meester et al. (1986) recognize three subspecies: G. i. cosensi: N Mozambique and E Tanzania. G. i. pringlei: Muheza, Tanzania (05° 10´ S, 38° 47´ E; Hubbard 1970b). Dorsal pelage black, ventral pelage and feet pure white; tail usually equal to or shorter than head and body length; only known from type locality.

G. i. inclusus: E Zimbabwe and Mozambique south of Zambezi R. Dorsal pelage very dark (from forehead to base of tail); flanks dark ochraceous-buffy or reddish. Similar Species G. leucogaster. Smaller; paler in colour, texture of fur sleeker. Distribution Endemic to Africa. Zambezian Woodland BZ and parts of Coastal Forest Mosaic BZ in Mozambique, E Zimbabwe and S Tanzania (east of L. Malawi). Perhaps occurs in localities between the currently known ranges. Habitat Sandy ground or sandy alluvium, often on fringe areas between dry and riverine woodland, confined to areas with a mean annual rainfall of >800 mm. Often associated with agricultural lands and forest fringes, but not recorded within forests. The subspecies G. i. pringlei recorded in dense grassland with moist, sandy soil. Abundance

Not known.

Adaptations Nocturnal and terrestrial. Moves by quadrupedal saltation. Burrows of G. i. pringlei have an entrance tunnel, one or two chambers, one of which contains the nest, one or two side tunnels and an escape tunnel (which may be 2–2.5 m from the main burrow) (Hubbard 1970b). Food has not been found in burrows. Foraging and Food Seeds, fruit and insects in captivity (as G. i. pringlei; Hubbard 1970b). Social and Reproductive Behaviour Appear to have identical habits to Bushveld Gerbils (G. leucogaster), with which they occur sympatrically, although always in smaller numbers (Smithers 1983). Simple burrow systems indicate a more solitary social structure than Bushveld Gerbils; burrows normally occupied by a single animal or a ! with young. Copulation (in G. i. pringlei) consists of single mount with intromission and ejaculation. Animals footdrum by pattering the hindfeet alternately when alarmed (Hubbard 1970b). Reproduction and Population Structure No information on seasonality of breeding. Gestation: 23–24 days. Litter-size: 2–3. Young not known to nipple-cling. Eyes open at Day 16–20. Predators, Parasites and Diseases Little information. No fleas recorded on T. i. pringlei. Conservation

Gerbilliscus inclusus

IUCN Category: Least Concern.

Measurements Gerbilliscus inclusus HB: 156.0 (152–161) mm, n = 13 T: 164.1 (135–191) mm, n = 13 277

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HF: 41.2 (39–44) mm, n = 13 E: 25 (21–29) mm, n = 13 WT: 115.6 (99–154) g, n = 13 GLS: 41.1 (36.7–44.3) mm, n = 4 GWS: 21.2 (19.2–22.2) mm, n = 4 M1–M3: 6.9 (6.6–7.2) mm, n = 4 Auditory bulla: 9.9 (8.9–10.7) mm, n = 4

Body measurements and weight: E Zimbabwe (recalculated from Smithers & Wilson 1979; as Tatera inclusa) Skull measurements: Mozambique (P. J. Taylor unpubl.) T. i. pringlei: larger body size, with shorter tail, hindfoot and ears Key References

Hubbard 1970; Smithers 1983. Edith R. Dempster

Gerbilliscus kempi KEMP’S GERBIL (NORTHERN SAVANNA GERBIL) Fr. Gerbille de Kemp; Ger. Kemps Nacktsohlen-Rennmaus Gerbilliscus kempi (Wroughton, 1906). Ann. Mag. Nat. Hist., ser. 7, 17: 375. Aguleri, Nigeria.

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). Sometimes included within Tatera valida (now Gerbilliscus validus) (e.g. Davis 1975a, Happold 1987, Bates 1988), but more generally considered to be a valid species (Rosevear 1969, Grubb et al. 1998). Synonyms: beniensis, benvenuta, dichrura, dundasi, flavipes, lucia, nigrita, ruwenzorii, smithi, soror; gambiana, giffardi, hopkinsoni, welmani. All except the last four were included as synonyms under Tatera valida (= Gerbilliscus validus) by Musser & Carleton (1993). Subspecies: none. Chromosome number: 2n = 36 (Matthey & Petter 1980, in Robbins & Baker 1978). Description Medium-sized robust rodent with shaggy hair and tail slightly longer than head and body. Dorsal pelage sandy-grey to sandy-orange; dorsal hairs sandy with dark grey at base and usually with black tip. Flanks similar to dorsal pelage, hairs usually without dark tips. Ventral pelage pure white. Dorsal and ventral colours clearly delineated. Chin, throat, chest and inner sides of limbs white. Eyes large, head rounded with moderately pointed nose. Tail long (ca. 100% of HB), sparsely haired, dark above, white below, with small pencil of darkish hairs (often lost in adults) at terminal end. Juvenile pelage sandy-grey, usually duller and less sandy than in adults. Males tend to be larger than !!. Each upper incisor with single groove. Nipples: not known. Geographic Variation Individuals at southern part of range (within, or close to, the Rainforest BZ) have greater density of blacktipped hairs and appear darker than northern individuals. Similar Species G. validus. On average larger (HB: 167 [135–195] mm, HF: 34.0 [30–39] mm, GLS: 41.7 [38.5–44.7] mm); tail lacks pencil at terminal end; auditory bullae larger; distribution mainly in eastern Africa. G. guineae. Similar size. Hindfoot on average longer; tail on average longer and relatively longer with well-developed pencil; distribution Senegal to Ghana and Togo. Taterillus gracilis. Much smaller and more slender; tail relatively longer (ca. 130% of HB) covered with short bristles and welldeveloped pencil at terminal end; syntopic and sympatric. Distribution Endemic to Africa. Guinea Savanna BZ and Northern and Eastern Rainforest–Savanna Mosaics. Recorded from

Gerbilliscus kempi

Gambia and Sierra Leone to Nigeria, and probably to S Sudan and NW Kenya; may sometimes extend northwards into parts of Sudan Savanna BZ in the east of the range. Sometimes found in savanna-like habitats on edge of rainforest. Recorded from coastal grasslands of the Dahomey Gap in Ghana (Grubb et al. 1998), Togo and Benin (Robbins & Van der Straeten 1996). Extent of geographic range in central and eastern Africa uncertain (see also G. validus). Habitat Savanna grasslands where there is good cover of grasses and/or dense shrubs, and where the soil is sandy and friable for digging burrows. Also occur wherever there are farmlands and plantations, such as maize and cassava fields, cocoa and oil palm plantations, and abandoned farmlands with thick grassy cover. Abundance One of the commonest rodents in suitable savanna habitats where the rainfall is 1000–1500 mm/year and where the wet season lasts for 7–8 months (Apr–Nov). In the Rainforest– Savanna Mosaic of Nigeria (7–8° N), comprised less than 10% of small terrestrial rodents. Likewise on the grasslands of the Shai Hills

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Gerbilliscus leucogaster

(Accra Plains, Ghana, ca. 6° N), formed about 10% of small rodents (n = 27) (Decher & Bahian 1999), and also ca. 10% (n = 3906 trappable small rodents; third commonest of 13 spp.) at Fro-Fro (7°55´ N), Côte d’Ivoire. Relative abundance increases from south to north, and near Kainji N. P., Nigeria (09° 50´ N) comprised nearly 100% of small rodents (Happold 1975b). Density up to 46/ha (= ca. 4.6 kg/ha) in Kainji N. P., Nigeria.

months of the year, and pregnant !! were recorded in Feb, Mar, Jun, Oct and Nov (Gautun 1975). The limited evidence suggests that pregnancies occur at the beginning and end of the wet season, and rarely during the main part of the wet season. Mean litter-size: 2.7 (n = 9) in Feb/Mar and 6.0 (n = 10) in Oct/Nov (Gautun 1975; no ranges given). Predators, Parasites and Diseases No information.

Adaptations Nocturnal and terrestrial. Kemp’s Gerbils live in burrows during the day and forage above ground at night. They have strong limbs and wide hindfeet for digging and burrowing, and large eyes and well-developed auditory bullae for good sensory perception. Foraging and Food Granivorous, herbivorous and oppor tunistic. The diet is primarily seeds, but also leaves, shoots and roots. In the dry season, insects are eaten when succulent foods unavailable. Kemp’s Gerbils also feed on crops and may cause damage in farmlands. They collect and store seeds in their burrows. Stored seeds probably provide food at beginning of the wet season when other seeds are scarce (D. C. D. Happold unpubl.). Social and Reproductive Behaviour Mostly unknown. High density of individuals in suitable habitats suggests overlapping homeranges. Several individuals may be kept together in captivity without any signs of aggressive behaviour. Reproduction and Population Structure Pregnant !! recorded in late wet season (Dec) in S Nigeria. Juveniles (25–50 g) abundant in dry season (Dec–Mar) in Kainji N. P., Nigeria. In C Côte d’Ivoire (07° 55´ N), adult !! were reproductively active in most

Conservation IUCN Category: Least Concern. A common and widespread species in suitable savanna habitats. Measurements Gerbilliscus kempi HB: 158 (140–190) mm, n = 12 T: 156 (142–173) mm, n = 12 HF: 32.7 (27–36) mm, n = 12 E: 21.5 (17–26) mm, n = 12 WT: 97–105 g, n = 13 GLS: 40.2 (38.7–41.6) mm, n = 12 GWS: 20.4 (18.9–21.8) mm, n = 9 M1–M3: 6.4 (6.2–6.6) mm, n = 12 Auditory bulla: 10.9 (10.5–11.3) mm, n = 12 Measurements: Nigeria (BMNH) Weight: Nigeria (Happold 1987, as Tatera valida) Key References

Happold 1975b; Rosevear 1969. D. C. D. Happold

Gerbilliscus leucogaster BUSHVELD GERBIL Fr. Gerbille à ventre blanc; Ger. Weissbauch-Nacktsohlen-Rennmaus Gerbilliscus leucogaster (Peters, 1852). Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin, p. 274. Mesuril and Boror, Mozambique.

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). Synonyms: angolae, bechuanae, beirae, beirensis, kaokoensis, limpopoensis, littoralis, lobengulae, mashonae, mitchelli, ndolae, nigrotibialis, nyasae, panja, pestis, pretoriae, salsa, schinzi, shirensis, stellae, tenuis, tzaneenensis, waterbergensis, zuluensis. Subspecies: none. Previously, 17 of these forms from the southern African region were considered to be subspecies, but their distribution is contiguous and populations of these forms integrate evenly throughout their range. Chromosome number: 2n = 40, FN = 66 (Qumsiyeh, 1986). Description Medium-sized gerbil with white underparts, darker dorsal surface and very long tail. Dorsal pelage reddish-brown to orange-buffy, depending on geographical area. Dorsal hairs slate-grey on basal two-thirds, changing to buffy-white, then pale rufous-buff at tip. Ventral pelage pure white from chin to tail tip. Head narrow, with pointed nose, long vibrissae, sides of muzzle white. Large eyes with white mark above and behind each eye. Ears elongated, dark brown, rounded at tips. Hindlimbs much longer than forelimbs, elongated hindfeet. Fore- and hindfeet pure white, five digits each, Digit 5 on forefoot reduced. Tail relatively long (ca. 115% of HB),

covered with dense short hairs, distinct brownish band down entire length on dorsal surface, white underneath, often darker terminal tuft. Anterior face of incisors grooved, lower incisors ungrooved. Auditory bullae well developed. Nipples: 2 + 2 = 8, but considerable variation. Geographic Variation Dorsal pelage ranges from bright cinnamon-buff in west to reddish-brown in east. Similar Species G. brantsii. Pelage softer and fluffier; tail without distinct dark line, usually white at tip. Distribution Endemic to Africa. Widely distributed in the Zambezian Woodland BZ, parts of the South-West Arid BZ (Kalahari Desert) and Highveld BZ. Northern limits uncertain, probably ca. 6° S. Widespread in Angola, S DR Congo, Zimbabwe, S Tanzania, Malawi, Mozambique, Namibia, Botswana and N South Africa. Southern limit in South Africa about 30° S, including the lowveld of Swaziland and NE KwaZulu–Natal Province. 279

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Habitat Predominantly associated with open grasslands and wooded savannas on light sandy soils or sandy alluvium. Common on sandy plains along dry river courses in C Namib, in miombo woodland in Zimbabwe and woodland savanna in Malawi. Absent from areas of heavy red clay soils or soft sand. Generally restricted to areas with mean annual rainfall above 250 mm, although in Namibia occur in areas with mean annual rainfall less than 100 mm. Occur at altitudes of sea level to ca. 1600 m. Abundance Abundance varies according to locality, vegetation and time of year. At Sengwa, Zimbabwe, number of individuals was 1.9–7.0/ha in Brachystegia woodland, 0.8–2.3/ha in thicket habitats and 0–1/ha in riverine grasslands (Linzey & Kesner 1997a). Percentage abundance was 70–90% (cool dry season) and ca. 86% (other seasons) in Brachystegia woodland (where it was always the most numerous species), 56–100% (cool dry season) and 67% (hot wet season) in thicket habitats, and 0% (most seasons) to 3% (cool dry season) in riverine grasslands. At Nylsvley, South Africa, percentage abundance varied during the dry season from 6% (in old fields, where it was the least common of the five species present) to 61% (in burnt Acacia woodland where it was the commonest of five species present) (Korn 1987). In Liwonde N. P., Malawi, comprised 18% of small terrestrial rodents (n = 106; 7 spp.) and was encountered only during the dry season (Happold & Happold 1990). The frequency of burning, and when burning occurs, has a considerable effect on the numbers of individuals and their percentage occurrence within the community of small mammals; in general formed higher proportions of the small mammal community in habitats that had been burnt at one or more times during the last three years than in unburnt control habitats (Korn 1981). De Graaff (1981) comments that ‘populations are subject to cyclic explosions in numbers’, but does not give quantitative data to support this statement. Adaptations Nocturnal and terrestrial. Moves by quadrupedal saltation. Excavates burrows, which are about 40–45 mm in diameter, with the entrance at the base of small bushes or grass clumps. Burrows are complex with many entrances and tunnels that interconnect underground. Burrow systems include chambers lined with vegetable debris. Bushveld Gerbils have many physiological characters concomitant with survival in hot, arid environments, although they are less aridadapted than other gerbil species. Basal metabolic rate and evaporative water loss is lower than in mesic rodents, minimal wet thermal conductance is about the same as expected for body mass, resting body temperature below thermoneutrality is lower than in mesic rodents, and the thermoneutral zone is broader than in more xeric-adapted gerbil species (Downs & Perrin 1994, Webb & Skinner 1996a).

Gerbilliscus leucogaster

Copulation consists of series of mounts with and without intromission, culminating in intromission with ejaculation; there is no lock and a copulatory plug is deposited after ejaculation. Multiple copulations and ejaculations occur over about an hour. Vocalize audibly between 5.5 kHz and 4.8 kHz; ultrasonic whistles begin at 48 kHz, descending to a trill at about 30 kHz; short ultrasonic ‘peeps’ emitted at about 40 kHz and 63 kHz. Males perform a post-copulatory song, which consists of series of whistles beginning at about 50 kHz, descending to about 15 kHz, and lasts about one minute. Animals footdrum when alarmed. Hindfeet patter alternately in bursts of 6–10 beats (Dempster & Perrin 1994).

Foraging and Food Bushveld Gerbils eat insects, seeds and herbage. Individuals in N South Africa eat mostly insects and seeds during warm wet season, and lose weight in cool dry season when eating mostly herbage. No seasonal variation in diet in Zimbabwe.

Reproduction and Population Structure Breeding occurs throughout the year, with peaks in Dec–Jan and Apr–May, associated with rainfall. Second breeding season in dry season reported in parts of Zimbabwe, elsewhere breeding ceases in cool dry season. Breeding strategy similar to other gerbil species: short gestation, large litter-size, altricial young, iteroparous (Perrin & Swanepoel 1987). Gestation: 28 days. Embryo number: 4.5 (2–9) over range of habitats. Mean littersize: 5.0. Mean birth-weight: 3.6 g. Growth rate in first 32 days: 0.9 g/ day.Young do not nipple-cling. Incisors erupt Day 5–6. Eyes open Day 16–21.Young weaned by Day 28. Annual reproductive capacity of !! is 28 young/year in mixed woodland habitat, and 12 young/year in miombo woodland habitat (Neal 1991). Testis mass of adult "": ca. 6% of total body weight, larger than most other rodent species. Testes of many "" regress during dry season. Sex ratio not significantly different from parity. Juveniles enter the population during the warm wet season, and population numbers decline in dry season; seasonal changes in population structure are variable.

Social and Reproductive Behaviour Social structure unknown, but burrows often clumped, and several individuals trapped from a single burrow system indicate a tolerant social structure. Adults rarely aggressive towards each other in laboratory encounters.

Predators, Parasites and Diseases Recorded as a prey item of the Barn Owl Tyto alba. Susceptible to infection of the neurotropic strain of African horse sickness and Listeria monocytogenes under laboratory conditions. Recorded as a reservoir of the plague bacillus Yersinia pestis

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in South Africa and DR Congo. Fleas include 25 species in the families Hystrichopsyllidae, Chimaeropsyllidae and Pulicidae (details in De Graaff 1981). In East Africa, Bushveld Gerbils carry three species of fleas of their own: Xenopsylla debilis, X. humilis and X. difficilis. Conservation IUCN Category: Least Concern. Abundant throughout its geographic range. Measurements Gerbilliscus leucogaster HB: 128.6 (89–155) mm, n = 1023 T: 148.5 (120–175) mm, n = 1024 HF: 33.5 (24–38) mm, n = 1068 E: 21.0 (18–26) mm, n = 1045 WT: 69.8 (32–114) g, n = 696

GLS: 37.3 (33.3–40.5) mm, n = 23 GWS: 19.0 (17.3–20.7) mm, n = 23 M1–M3: 6.5 (5.6–7.9) mm, n = 23 Auditory bulla: 10.5 (9.7–11.1) mm, n = 18* Body measurements and weight: Former Transvaal (Rautenbach 1978; as Tatera leucogaster) Skull measurements: KwaZulu–Natal and Northern Cape Province, South Africa, and Namibia (P. J. Taylor unpubl.) *A series from Angola (Crawford-Cabral 1988) had slightly smaller auditory bullae (10.3 [9.4–10.9] mm) Key References Dempster & Perrin 1994; Neal 1991; Perrin & Swanepoel 1987; Webb & Skinner 1996a. Edith R. Dempster

Gerbilliscus nigricaudus BLACK-TAILED GERBIL Fr. Gerbille à queue noire; Ger. Schwarzschwanz-Nacktsohlen-Rennmaus Gerbilliscus nigricaudus (Peters, 1878). Monatsber. K. Preuss. Akad. Wiss. Berlin 1879: 200. (publ. 1878). Ndi, Taita, Kenya.

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). This species, G. robustus and G. validus are closely related and sympatric in parts of their ranges. Although the allblack tail is the main character of nigricaudus, the tail of G. robustus is black (or partially black) in some parts of its range, especially in the northern part. The three species are difficult to differentiate with absolute certainty using external characters only; and some individuals ascribed to nigricaudus in the past are probably G. robustus. A combination of external measurements, colour and certain skull characters are necessary to separate these species from each other. Bates (1988) considered that bayeri (Mt Elgon and Karamoja) and bodessae (Ethiopia), both formerly considered to be subspecies of G. nigricaudus, are referable to G. robustus. Synonyms: nyama, percivali. Subspecies: two. Chromosome number: not known. The individuals referred to by Matthey (1969), for which he gave a chromosome number of 2n = 40, are considered to be T. robusta (now Gerbilliscus robustus) by Bates (1988). Description Large brown gerbil with long black tail. Pelage long, soft and sleek. Dorsal pelage brown, suffused with black in some individuals; dorsal hairs pale grey at base, with ochre, brown or black at tip. Flanks paler; hairs white at base, pale ochre or sandy at tip. Ventral pelage white. Dorsal and ventral colours clearly delineated. Head and cheeks similar to dorsal pelage. Chin, chest, inner surfaces of limbs white. Hindfeet white; soles naked and darkly pigmented. Tail long (ca. 110% of HB), thickly covered with very dark brown or black bristles above and below; longer black hairs form small pencil at tip in some individuals; a few pale hairs may occur on undersurface, especially on distal third of tail; colours above and below merge laterally. Skull large; rostrum rounded and wide (cf. G. robustus); upper incisors opisthodont, each incisor with clearly defined single groove. Males larger than !! for most body and cranial measurements. Width of M1 2.4–2.5 mm. Nipples: not known.

Gerbilliscus nigricaudus

GeographicVariation Two subspecies are recognized (Bates 1988): G. n. nigricaudus: NE Tanzania and S Kenya. Larger size; tail black above and below. G. n. nyama (incl. percivali): N Kenya and Somalia. Smaller size; tail black above, with some pale hairs below, especially on distal third. Similar Species G. robustus. Smaller HB; tail brown, often with black at tip or on all of upper surface; often black/brown pencil at terminal end; no white tip to tail; skull on average smaller in all dimensions. 281

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G. validus. HB on average smaller; tail brown, normally without any black, no white tip to tail, no pencil at terminal end; skull usually smaller. G. boehmi. HB smaller; tail brown, usually longer (actual length and relative to HB), white at terminal end; skull usually smaller. Distribution Endemic to Africa. Somalia–Masai Bushland BZ. Savanna woodlands and grasslands of C and N Kenya east of the Rift Valley, extreme NE Tanzania, S Somalia and S Ethiopia. Habitat Savanna woodlands with bushy thickets and scattered Acacia, Stercularia and Terminalia trees; also open grassy plains dominated by Chrysopogan, Sehima and other grasses (Meru N. P., Kenya; Neal 1984b). Habitat shows seasonal changes in cover and productivity; rainfall is bimodal with two wet seasons per year.

early dry seasons and ceased at the end of each dry season (Sep and Mar) (Meru N. P., Kenya; Neal 1982).Weight of testes greater during and immediately after wet season than during dry season; no evidence that "" sexually inactive at any time of year. Embryo number: 5.22 (1–8, n = 37 litters). Litter-size increases with increasing maternal weight, and mean litter-size (regardless of maternal size) is higher in wet season (5.5) than in early dry season (4.2). Pregnancy rate: up to 50% of adult !! pregnant during wet seasons. Females may have three litters/year, and a reproductive capacity of up to 24 young/year (Neal 1982). Young conceived and born during wet seasons are recruited into the trappable population in Jan–Feb and Jul–Aug; maturity is attained by beginning of following wet season. Young born in early dry seasons fail to contribute to the population (perhaps due to early mortality?). Predators, Parasites and Diseases No information.

Abundance Distribution patchy, although probably relatively common in suitable habitats. Adaptations Nocturnal and terrestrial. Individuals exhibit rapid response to seasonal changes in food availability (caused by seasonal changes in rainfall, number of arthropods and plant growth), which, in turn, regulates reproductive cycle (see below). Foraging and Food Omnivorous and opportunistic. Pronounced seasonal changes in diet. In the dry seasons (in Meru N. P., Kenya; Neal 1984b) arthropods (mainly insects) comprise the main proportion of the diet (ca. 90%), with smaller amounts of seeds (1– 9%) and forbs (1–3%). In the wet seasons, the proportion of arthropods drops (50–70%), and the proportions of grasses and sedges (3–13%), forbs and browse (13–20%) and seeds (7–23%) increase. In the period after the wet season and before the dry season begins, when the grasses have set seed, the diet is mainly seeds (53–62%) and insects (30–35%) with hardly any grass or forbs (Neal 1984a). Gerbils utilize the foods that, at each season, are common and provide the best source of energy and protein. The diet was similar to that of parapatric Acomys wilsoni and Gerbilliscus robustus (as Tatera robusta), which suggests possible competition for food between these species (Neal 1984a). Social and Reproductive Behaviour

No information.

Reproduction and Population Structure Seasonal reproduction, with two periods of breeding each year, correlate with the bimodal pattern of rainfall. Reproductive activity (pregnancies, number of corpora lutea in ovaries) was maximum during each of the two wet seasons per year (Nov–Jan, Apr–May), declined during the

Conservation

IUCN Category: Least Concern.

Measurements Gerbilliscus nigricaudus* HB (""): 185.8 (178–193) mm, n = 4 HB (!!): 164.0 (130–164) mm, n = 6 T (""): 200.8 (190–208) mm, n = 4 T (!!): 187.2 (170–204) mm, n = 6 HF (""): 40.5 (40–41) mm, n = 4 HF (!!): 37.3 (34–39) mm, n = 6 E (""): 22.3 (20–24) mm, n = 3 E (!!): 21.8 (21–24) mm, n = 6 WT (""): 132 (80–195) g** WT (!!): 114 (80–161) g** GLS (""): 48.7 (47.0–50.5) mm, n = 3 GLS (!!): 45.3 (43.1–47.9 ) mm n = 6 GWS (""): 25.3 (24.2–26.1) mm, n = 4 GWS (!!): 22.6 (21.9–23.2) mm, n = 5 M1–M3 (""): 7.0 (6.7–7.3) mm, n = 3 M1–M3 (!!): 6.9 (6.7–7.0) mm, n = 4 Auditory bulla (""): 14.5 (13.6–15.1) mm, n = 4 Auditory bulla (!!): 13.3 (12.7–13.8) mm, n = 6 Measurements: Kenya (Bates 1988) Weight: Kenya (Neal 1982) *As Tatera nigricauda nigricauda **No sample size Key References Bates 1985, 1988; Neal 1982, 1984a. D. C. D. Happold

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Gerbilliscus phillipsi

Gerbilliscus phillipsi PHILLIPS’S GERBIL Fr. Gerbille de Phillips; Ger. Phillips Nacktsohlen-Rennmaus Gerbilliscus phillipsi (de Winton, 1898). Ann. Mag. Nat. Hist., ser. 7, 1: 253. Hanka Dadi, Somalia.

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). Often considered to be a subspecies of G. robustus, but raised to specific level by Bates (1985, 1988) on the basis of size (see also Musser & Carleton 1993). Synonyms: bodessana, umbrosa and probably miniscula (see Bates 1988). Subspecies: none. Chromosome number: not known. Description Medium-sized gerbil, similar to G. robustus. Smallest species of Gerbillurus in eastern Africa. Dorsal pelage pale brown to orange-brown, darker on lower back and rump; hairs grey at base, with dull pale orange-brown subterminal band and (especially on rump) black tip. Flanks paler, with fewer black-tipped hairs. Ventral pelage pure white, clearly delineated from colour of flanks. Lips, cheeks, throat and chest pure white. Ears large, slightly pigmented. Forefeet white above; five digits; Digits 1 and 5 short. Hindfeet white above; soles with brown pigment; Digit 1 short, not reaching base of other digits; Digits 2, 3 and 4 long, Digit 5 shorter; all digits with short pointed claws. Tail long (ca. 128% of HB), bicoloured, pale orange-brown above, white below; some black hairs above towards terminal end but without pencil or tuft. Skull: smaller than that of G. robustus (as T. robusta; Bates 1988). Gerbilliscus phillipsi

Geographic Variation Populations in Kenya have darker pelage than those further north in Ethiopia and Somalia (Bates 1985). Similar Species G. robustus. On average larger in body and skull measurements (HB: 152.2 [120–190]) mm; GLS: 41.9 [39.0–44.7] mm; GWS: 21.1 [19.5–22.4] mm). Tail usually with pencil. Distribution Endemic to Africa. Somalia–Masai Bushland BZ. Recorded from NC Kenya, Ethiopia (Rift Valley) and Somalia. Habitat Dry arid savanna, and semi-desert. In the Omo Valley of Ethiopia (Hubert 1978b) populations (recorded as Tatera nigricauda) occur in the treeless grasslands and thickets where the soils are heavy and clay-like; these grasslands are seasonally flooded. On the lower slopes of the valley they live in very arid habitats of tuffs and recently eroded sediments, which support a sparse arid vegetation (e.g. Euphorbia, Adenium). Abundance The species has been recorded at only nine localities (Bates 1988). In the Omo Valley, Ethiopia, comprised 21% of small trappable rodents in tree and shrub habitats near the river (total n = 19, 4 spp.), 20% in grasslands near the river (total n = 53, the second most numerous of 8 spp.) and 6% in the arid habitats (total n = 30; 6 spp.).

Remarks Only found in dry habitats, so evidently well adapted to high temperatures and a lack of free water. Recorded in the same habitat in the Omo Valley as Acomys wilsoni, Saccostomus mearnsi and Mastomys erythroleucus (in grasslands), and Taterillus harringtoni, Arvicanthis somalicus, Gerbillus pusillus and Xerus rutilus (in arid habitats) (Hubert 1978b). Conservation

IUCN Category: Least Concern.

Measurements Gerbilliscus phillipsi HB: 136.3 (116–145) mm, n = 9 T: 174.1 (162–185) mm, n = 9 HF: 34.0 (32–37) mm, n = 10 E: 19.2 (17–21) mm, n = 9 WT: n. d. GLS: 38.4 (37.6–39.2) mm, n = 9 GWS: 17.8, 19.4 mm, n = 2 M1–M3: 5.9 (5.6–6.2) mm, n = 9 Auditory bulla: 11.8 (11.4–12.4) mm, n = 9 Throughout geographic range (Bates 1988, as Tatera phillipsi) Females only; external measurements of "" are larger than for !! Key References

Bates 1985, 1988. D. C. D. Happold 283

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Gerbilliscus robustus FRINGE-TAILED GERBIL Fr. Gerbille robuste; Ger. Fransenschwanz-Nacktsohlen-Rennmaus Gerbilliscus robustus (Cretzschmar, 1826). In: Rüppell, Atlas Reise Nordl. Afrika, Zool. Säugeth. 1: 75. Ambukol, Sudan.

Taxonomy Originally described in the genus Tatera (see profile Genus Gerbilliscus). The species of Gerbilliscus in eastern Africa – G. robustus, G. validus, G. nigricaudus and G. phillipsi – are morphologically similar, and positive identification requires consideration of a number of characteristics (see Table 23). Colouration of tail and length of hindfoot are not good diagnostic characters by themselves. The similarity between this species and G. nigricaudus has resulted in some specimens of G. robustus being incorrectly identified as G. nigricaudus (e.g. Delany 1964a, Hubert 1978b). This species is closely related to G. validus, and the two may, in fact, represent a single polymorphic species (D. Kock pers. comm.). Synonyms: bayeri, bodessae, iconica, loveridgei, macropus, mombasae, muansae, pothae, shoana, swathlingi, taylori, vicina. Subspecies: none. Chromosome number: 2n = 40, FN = 70 (Omo Valley, Ethiopia; Bates 1988 as T. robusta). Description Large dark-coloured gerbil. Pelage long and soft. Dorsal pelage dark brown, flecked with black. Dorsal hairs (12–15 mm long) grey at base, with cinnamon, dark brown or black tip. Flanks paler, orange-brown, with fewer black-tipped hairs.Ventral pelage pure white. Dorsal and ventral colours clearly delineated. Head blackishbrown, with sandy-brown cheeks. Eyes large, sometimes with black eye-rings. Black vibrissae. Ears large, rounded and naked. Chin, throat, neck and chest pure white. Fore- and hindlimbs pale brown above, white below. Soles of hindfeet naked, darkly pigmented. Tail long (ca. 115% of HB), thickly covered with small short hairs; colour varied – brown with varying amounts of black to nearly all black, usually with pencil of hairs on terminal third; paler below; upper and lower colours usually clearly delineated (see details below). Skull: rostrum long and narrow; width of M1: 2.0 (2.0–2.4) mm (cf. G. validus; see Table 23). Sexes similar in size. Nipples: not known. Geographic Variation Dorsal pelage varies in different parts of geographic range. Individuals from semi-arid areas of Sudan are paler than those from Kenya and Tanzania. Tail colour varies geographically. In southern part of range (Tanzania), brown above with some brownish-black or black hairs at terminal end or on terminal half; white, ochre or pale brown below; in northern part of range (Ethiopia), mostly or completely black above; ochre or brown below (and sometimes with black on the terminal third). Individuals intermediate between these extremes in central part of range. Individuals with dark tails appear very similar to G. nigricaudus although, in most individuals, the tail is not pure black above and below as in G. nigricaudus (Bates 1985).The large amount of variation in this species was responsible for each variant being described originally as a new species (see synonyms). Similar Species G. validus. Slightly larger on average; tail ca. 95% of HB, usually without any black colouration; width of M1 usually >2.5 mm. G. nigricaudus. Tail ca. 100% of HB, always black above and below; width of M1 2.4–2.5 mm.

Gerbilliscus robustus

G. phillipsi. On average smaller size; tail ca. 130% of HB, orangebrown above; width of M1 200 ind/ha causing damage to cereal and food crops, and becoming major pests in agricultural fields (Bernard 1969, Zaime & Gautier 1988).

Meriones shawi

Adaptations Nocturnal and partly diurnal. The most hydrophilic jird with a higher water turnover and a lower urine osmotic pressure

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(2040 mOsmol/L) than M. libycus. Also requires a higher iodine intake than M. libycus to maintain thyroid function (Ben ChaouachaChekir 1996). Foraging and Food Herbivorous and opportunistic, eating many different plants and seeds. Diet varies according to season and habitat. In Morocco, preferred plants are species of Graminaceae (Sorghum halepense and Stipa retorta), Papilionaceae (Medicago hispida) and Compositae (Calendula ægyptica). Also eats Salsola vermiculata (Chenopodiaceae) in spring and autumn, and arthropods in Jul (Zaime & Gautier 1989). Seeds are stored in special caches in the burrows. Social and Reproductive Behaviour Colonial. Range length (distance between most distant captures of an individual) varies according to season and sex. Range length in spring (May–Jun) is 40–50 m, and 10–20 m in autumn and winter (Sep–Feb). Range length of males (ca. 70 m) is larger than for females (ca. 30 m), especially in uncultivated areas where food availability is low (Zaime & Gautier 1988). Communication between individuals is by footdrumming and vocalizations (Bridelance 1989). Reproduction and Population Structure Pregnancies recorded during wet season in Morocco (Nov–May; Zaime & Gautier 1988) and during dry season in Tunisia (Mar–Sep; Bernard 1969). Gestation: 21 days. Litter-size: 5.0 (3–8, with a maximum [mean 6.2] in Mar). Weight at birth: 3.5–6 g. Predators, Parasites and Diseases Predators include foxes (Vulpes vulpes) and owls (Tyto alba, Asio otus, Strix aluco, Bubo bubo). Remains found in pellets of eagle-owls, comprised 0.5–3.5% of prey in Morocco (M. Thévenot pers. comm.) and 29% in N Algeria (Boukhamza et al. 1994). Reservoir of protozoan Leishmania major, which causes zoonotic cutaneous leishmaniasis (WHO 1990) (see also Psammomys obsesus). Endoparasites in Tunisia include nematodes (Bernard 1987). Conservation IUCN Category: Least Concern. Shaw’s Jirds are considered as pests in cultivated areas. Control measures may be necessary when population numbers are high.

Measurements Meriones shawi shawi HB: 143 (128–160) mm, n = 31 T: 140 (122–155) mm, n = 25 HF: 35 (32–37) mm, n = 31 E: 19 (17–22) mm, n = 31 WT: 91 (70–120) g, n = 11 GLS: 38.8 (37.1–41.5) mm, n = 20 GWS: 22.2 (20.6–23.5) mm, n = 18 M1–M3: 6.0 (5.6–6.3) mm, n = 22 Egypt (Osborn & Helmy 1980) Meriones shawi grandis HB: 167 (138–200) mm, n = 18 T: 159 (134–185) mm, n = 12 HF: 38 (33–40) mm, n = 18 E: 22 (19–24) mm, n = 17 WT: 242 (230–255) g, n = 3 GLS: 45.1 (39.9–50.4) mm, n = 18 GWS: 25.4 (22.3–28.6) mm, n = 17 M1–M3: 6.2 (5.6–6.8) mm, n = 18 Morocco (MNHN) Meriones shawi trouessarti HB: 131 (115–150) mm, n = 19 T: 124 (112–146) mm, n = 19 HF: 31 (30–34) mm, n = 19 E: 16 (14–17) mm, n = 19 WT: 65 (45–86) g, n = 19 GLS: 35.7 (34.5–36.9) mm, n = 4 GWS: 19.7 (19.2–20.0) mm, n = 4 M1–M3: 5.5 (5.5–5.6) mm, n = 4 Auditory bulla: 13.3 (12.4–14.2) mm, n = 15* Tunisia, Sidi Bouzid (E. Calvet unpubl.) *Egypt (Osborn & Helmy 1980) Key References 1988.

Zaime & Gautier 1988, 1989; Zaime & Pascal E. Fichet-Calvet

GENUS Microdillus Peel’s Pygmy Gerbil Microdillus Thomas, 1910. Ann. Mag. Nat. Hist., ser. 8, 5: 197. Type species: Gerbillus peeli de Winton, 1898.

Small gerbil-like rodents, often considered as a subgenus of Gerbillus but now recognized as a full genus (Petter 1975a, Musser & Carleton 1993, 2005). The genus has many gerbil-like features, but is characterized by small size, short tail and short square skull, which is abnormally bowed with a strong convex cranial profile. Unlike

Gerbillus, the upper third molar has three or four cusps (Petter 1977a). There is only one species, Microdillus peeli. D. C. D. Happold

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Family MURIDAE

Microdillus peeli PEEL’S PYGMY GERBIL (SOMALI PYGMY GERBIL) Fr. Gerbille pygmée de Somalie; Ger. Peels Koboldrennmaus Microdillus peeli (de Winton, 1898). Ann. Mag. Nat. Hist., ser. 7, 1: 250. Eyk, Somalia (see Roche & Petter [1968] for map).

Taxonomy Originally described in the genus Gerbillus (see genus profile). Synonyms: none. Chromosome number: not known. Description Very small gerbil with long pelage and short tail. Dorsal pelage dull orange-brown; dorsal hairs grey at base, with orange-brown tips. Pelage long and shaggy, so that grey bases of hairs may show on surface of pelage. Flanks paler; hairs whitish-grey or white at base. Ventral pelage pure white. Crown of head similar to back. Large eyes. Nose, lower cheeks, chin and throat white. Conspicuous supraorbital and postauricular white patches. Limbs and feet white. Soles of feet naked. Tail short (ca. 80% of HB), rather thick, slightly scaly, covered with brown or blackish bristles; without terminal pencil. Skull short and square; upper incisors opisthodont; zygomatic arch curves deeply downwards to form a large orbit; zygomatic plate small with almost vertical anterior face; auditory bullae (tympanic bullae and mastoids) large (Figure 53). Nipples: not known. Geographic Variation

None recorded.

Similar Species In all other species of gerbils of similar size (except Desmodillicus braueri) the tail is longer than HB (Table 22).

Figure 53. Skull and mandible of Microdillus peeli (BMNH 8.4.9.9).

Distribution Endemic to Africa. Somalia–Masai Bushland BZ. Known only from three localities in N and C Somalia. Not recorded from similar habitats in Ethiopia. Habitat In C Somalia, found in sandy habitats. Other individuals recorded from hilly country (1500 m) in N Somalia. Abundance are scattered.

Probably rare; there are few records and populations

Remarks The tail appears to be used for fat storage in the same way as in Pachyuromys duprasi and Gerbillus simoni. The long shaggy pelage is probably an adaptation to the cool nights in winter (as in G. somalicus). Most specimens were ‘caught on track at night’. Conservation IUCN Category: Least Concern. However, limited geographic range and probable rarity may be cause for concern.

Microdillus peeli

Measurements Microdillus peeli HB: 72.1 (66–80) mm, n = 12 T: 57.4 (50–65) mm, n = 12 HF: 17.8 (16–19) mm, n = 12 E: 11.4 (10–13) mm, n = 12 WT: n. d. GLS: 24.4 (23.7–25.2) mm, n = 6 GWS: 14.5, n = 1

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M1–M3: 3.7 (3.6–3.9) mm, n = 6 Auditory bulla: 9.5 (9.2–9.8) mm, n = 6 Somalia (BMNH) Key Reference

Roche & Petter 1968. D. C. D. Happold

GENUS Pachyuromys Fat-tailed Jird Pachyuromys Lataste, 1880. Le Naturaliste, 2 (40): 313. Type species: Pachyuromys duprasi Lataste, 1880.

A monotypic genus widespread in semi-arid regions of North Africa. The genus is characterized by hairy soles on the hindfeet, relatively short thickened tail without pencil and often club-shaped at terminal end, faint groove on each upper incisor, prismatic molars (but to a lesser extent than in Psammomys, Meriones and Sekeetomys), very

inflated auditory bullae and wide posterior palatal foramina from mid M1 to mid M2 (see also Table 27). Further details are given in the species profile. The single species is Pachyuromys duprasi. D. C. D. Happold

Pachyuromys duprasi FAT-TAILED JIRD Fr. Gerbille à queue grosse; Ger. Fettschwanzmaus Pachyuromys duprasi Lataste, 1880. Le Naturaliste, 2 (40): 313. Laghouat, Algeria.

Taxonomy Synonyms: faroulti, natronensis. Subspecies: none. Chromosome number: 2n = 54 (Qumsiyeh & Schlitter 1991). Description Small pale-coloured rodent with thick short naked tail. Pelage long, fine and soft with a rather shaggy appearance. Dorsal pelage beige, sandy or pale orange; hairs dark grey at base, beige or sandy subterminal band, sometimes with black tip. Dorsal hairs may lie irregularly so black tips form small lines across the body. Ventral hairs pure white. Head similar in colour to dorsal pelage. Eyes large, dark. Ears comparatively small, with sparse longish hairs; ears often partially obscured by pelage of head and neck. Muzzle, lips, chin and throat white. Fore- and hindlimbs small. Forefoot with four digits, each with small claw. Hindfeet with five digits, each with small claw. Feet and digits well covered with long white hairs.Tail short (ca. 54% of HB), thick and club-shaped, naked and without scales; terminal pencil absent; size of tail varies seasonally (see below). Skull with

Pachyuromys duprasi.

single faint groove on each upper incisor, braincase broad, auditory bullae very inflated (ca. 47% of GLS) and extending posteriorly to occiput (see also genus profile) (Figure 54). Nipples: not known. Geographic Variation

None recorded.

Figure 54. Skull and mandible of Pachyuromys duprasi (BMNH 4.11.3.118).

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rainstorms. Burrows are extensive, with numerous entrances (up to 12), which descend almost vertically into the burrow. Burrows are mostly shallow (5–10 cm) and rarely deeper than ca. 30 cm (max depth ca. 100 cm), and contain a nest. Burrows may be used for storage of food (Petter 1961). Fat-tailed Jirds become active at dusk, and may forage considerable distances (up to 2 km) from their burrows (Mermod 1970). One of the unique features of this species is its short fat tail, which is used to store fat; the tail alters in size according to season and to the amount of stored fat. In captivity, they are capable of entering ‘torpor’ for several days when Tb is 32–35 °C (Petter 1961). The auditory bullae are greatly inflated and, as in many species of gerbils and jerboas, provide for very sensitive perception of sounds.

Pachyuromys duprasi

Similar Species Meriones spp. Larger HB (mean >130 mm); tail long (mean >135 mm, at least 95% of HB) with black terminal pencil; upper incisor teeth each with distinct longitudinal groove; auditory bulla smaller (mean 14.6–15.3 mm, ca. 32–40% of GLS). Psammomys spp. Larger HB (mean 122 mm or larger); tail moderately long (mean ca. 115 mm, ca. 75% of HB) with black terminal pencil; upper incisor teeth without longitudinal groove; auditory bulla smaller (mean 13.3 mm, ca. 32–39% of GLS). Distribution Endemic to Africa. Sahara Arid BZ. Recorded from Mauritania, Morocco, Algeria, Tunisia, Libya and Egypt. Distribution is widespread but very localized in semi-arid areas south of the Atlas Mts in Morocco and Algeria, and south of the coastal plain in Libya and Egypt. In Algeria extends southwards to ca. 25° N (as evidenced by remains in owl pellets). One isolated record in N Mali (Heim de Balsac 1968). Habitat Hamadas with coarse pebbles and large boulders where vegetation is sparse, and along the edges of shallow dry watercourses that bisect the hamadas (Libya: Ranck 1968; Algeria: Daly & Daly 1979). Vegetated ‘sand sheets’ among sparse vegetation, and rocky deserts (Egypt: Osborn & Helmy 1980). Abundance Generally rarely encountered. Few specimens are obtained even where other species of rodents are common (Daly & Daly 1979). These jirds were more abundant than usual at BeniAbbès, Algeria, when there were large numbers of crickets (Petter 1961) (see also below). Adaptations Nocturnal and terrestrial. Movement is by walking and running, and not by bounding as in many other gerbils (Petter 1961). Fat-tailed Jirds construct large complex burrows, often on slopes, where the sand is dry and has been compacted by ephemeral

Foraging and Food Omnivorous; mostly herbivorous. Detailed information on diet not available. In Algeria, burrows contained fragments of fruits (Colocynthis vulgaris and Hyoscyamus niger) (Petter 1961), and in Egypt, Fat-tailed Jirds have been observed to feed on Anabasis articulata and A. monosperma (Osborn & Helmy 1980). In captivity, they feed on grains, chopped meat, cheese, lettuce and lucerne; the addition of food containing meat to the diet stimulated reproduction (Petter 1961). The suggestion that snails may be eaten (Setzer 1957) needs confirmation. Social and Reproductive Behaviour

No information.

Reproduction and Population Structure Gestation: ca. 21 days. Litter-size (in captivity): usually 3–5 (also 7 [n = 1], 9 [n = 1]). Births in captivity (Giza, Egypt) in Apr, May, Jul, Oct and Nov (Flower 1932). No data for wild populations, but likely to be opportunistic. Ears open Day 5–8, hair begins to grow Day 7–9; incisors erupt Day 11–15; eyes open Day 20–21; adult weight and body size ca. Week 10 (Petter 1961). Predators, Parasites and Diseases Remains are common in owl pellets (Kowalski & Rzebik-Kowalska 1991). One species of flea, Xenopsylla nubica, recorded in Algeria (Beaucornu & Kowalski 1985); seven species of fleas recorded on animals and in nests within burrows in Egypt: Ctenocephalides felis, Synosternus cleopatrae, Xenopsylla conformis, X. ramesis, Stenopoma tripectinata, Nosopsyllus henleyi, Hopkinsipsylla occulata (Hoogstraal & Traub 1965a). These fleas are also found on many other species of desert gerbils. Conservation

IUCN Category: Least Concern.

Although not common, the species has a widespread distribution and is unlikely to be threatened. Measurements Pachuromys duprasi HB: 108.3 (93–121) mm, n = 4 T: 58.2 (55–62) mm, n = 4 HF: 23.3 (22–24) mm, n = 4 E: 14.0 (12–16) mm, n = 4 WT: 36.5 (22–45) g, n = 3 GLS: 24.9 (32.4–36.5) mm, n = 4 GWS: 19.3 (17.5–20.2) mm, n = 4

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M1–M3: 5.2 (4.8–5.7) mm, n = 4 Auditory bulla: 16.7 (15.6–17.6) mm, n = 10 Egypt (Osborn & Helmy 1980) Auditory bulla: throughout geographic range (BMNH) Key References Kowalski & Rzebik-Kowalska 1991; Osborn & Helmy 1980; Petter 1961. D. C. D. Happold

GENUS Psammomys Sand Rats Psammomys Cretzschmar, 1828. In: Rüppell, Atlas Reise Nordl. Afr., Zool. Säugeth., p. 56. Type species: Psammomys obesus Cretzschmar, 1828.

Psammomys obesus.

The genus contains two species that live in semi-arid regions of N Africa, mostly north of the Sahara Desert. Habitats include salty lowlands, wadis and coastal deserts where saltbushes are prevalent. Species in the genus are large heavily built rat-like gerbils. Dorsal pelage is sandy coloured, and the thick hairy tail is comparatively short (less than ca. 75% of HB) with a black terminal pencil. Hindfeet are broad and slightly hairy.The skull is robust, with well-developed supraorbital, temporal and occipital crests, large inflated auditory bullae (Figure 52), and narrow slit-like anterior and posterior palatal foramina. Upper incisors do not have a longitudinal groove (cf. Meriones); cheekteeth are similar in structure to those of Meriones. Cheekteeth do not have cusps (at any stage of wear) and the laminae are prismatic (as in Meriones and Sekeetamys) (Figure 55, see also Table 27). Species in the genus are adapted for living in arid conditions.Their unique characters are diurnal activity and the production of large quantities of very concentrated urine. These characters are quite different to those of other arid-living small rodents and are associated with their ability to feed primarily on succulent saltbushes, rich in salt and water. The genus Psammomys is placed in the subfamily Rhombomyini, which also includes the genera Sekeetamys, Meriones (and three other non-African genera [Pavlinov et al. 1990]). The number of species

Figure 55. Skull and mandible of Psammomys obesus (BMNH 3.12.8.50).

in the genus is uncertain. Here, following Musser & Carleton (2005), two species are recognized – the larger P. obesus and smaller P. vexillaris. Several forms are considered now as synonyms of P. obesus, and P. vexillaris is also considered as a synonym of P. obesus by some authorities. The two species of Psammomys are distinguished by body size, pelage colour and distribution. E. Fichet-Calvet

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Psammomys obesus FAT SAND RAT Fr. Rat des sables obèse; Ger. Fette Sandratte Psammomys obesus Cretzschmar, 1828. In: Rüppell, Atlas Reise Nordl. Afr., Zool. Säugeth., p. 58, pl. 22. Alexandria, Egypt.

Taxonomy Many forms have been described as species of Psammomys, including algiricus in Algeria, tripolitanus and vexillaris in Libya, nicolli and obesus in Egypt and terræsanctæ in Palestine. All, except vexillaris, are now regarded as synonyms of obesus (Musser & Carleton 1993, 2005).The Fat Sand Rat is one of the best studied and most interesting of North African small rodents. Synonyms: algiricus, dianae, elegans, nicolli, roudairei, terraesanctae, tripolitanus. Subspecies: none. Chromosome number: 2n = 48, FN = 74–78 (Qumsiyeh & Schlitter 1991). Description Large heavily built sand rat with long pelage. Dorsal pelage ochre to tawny with long dark brown guard hairs; hairs dark grey at base, with orange or brown terminal band and black tip; width of bands varies geographically. Flanks and ventral pelage pale ochre. Massive head with large crown; vibrissae very long, pale or brown. Eyes large. Ears small and round, hairy, grey to ochre, set low on side of head. Small white postauricular patch. Fore- and hindlimbs short with buffy hairs on inner surface. Soles of hindfeet partly hairy; claws dark. Tail of moderate length (ca. 70% of HB), very hairy, ochre or brown with black terminal pencil. Skull: well-developed supraorbital ridges; auditory bullae inflated (ca. 32% of GLS); upper incisors smooth, without groove. Males tend to be larger than !!. Nipples: 2 + 2 = 8. Geographic Variation None recorded. Similar Species Psammomys vexillaris. On average smaller; pelage paler; ear smaller; E Algeria to W Libya only. Meriones spp. Smaller (except for M. shawi grandis); dorsal pelage similar, but with white ventral pelage; ear on average larger; sympatric throughout most of geographic range. Distribution Sahara Arid BZ. Widespread but disjunct distribution in salty lowlands, wadis and coastal deserts in Mauritania, Morocco, Algeria, Libya and Egypt where annual rainfall is 250– 300 mm; also in southern part of the Haute Plateaux and Haggier Mts of Algeria. Outlier population on Red Sea coast of Sudan. Southern limit in the Sahara depends on the location of suitable habitats such as salt steppes near an oasis or a chott (= ancient lake). Extralimitally recorded from Syria, Jordan, Israel and parts of Arabian Peninsula. Habitat Preferred habitats are succulent halophytic steppes (called daya in Egypt, and sebkhet in Tunisia) where the soil is moist, muddy and salty (salinity 9–30 g/L) (Petter 1961, Fichet-Calvet et al. 2000).The main plants in these habitats are saltbushes (Arthrocnemum, Atriplex, Halocnemum, Salsola and Sueda; family Chenopodiacae), which are essential for the survival of these sand rats. Abundance Generally common; populations fluctuate seasonally. Abundance in Tunisia ranged from 5 individuals/100 m of trap line

Psammomys obesus

(summer and early autumn) to 25/100 m (spring). In Morocco, densities (in Jul) of 42/ha have been recorded (Zaime & Pascal 1988). Abundance also varies spatially in relation to flooding (in salt steppes). Multi-annual fluctuations have been recorded, but the causes are as yet unknown (Fichet-Calvet et al. 1999b, 2000). Adaptations Fat Sand Rats live in an ecological niche utilized by no other species of rodent. They feed on the succulent leaves of saltbushes, which provide large amounts of water but are low in energy. The leaves of saltbushes contain salt (up to 12% by dry weight) as well as large quantities of water (up to 82%; Petter 1952). The kidneys are large and very efficient compared with those of other rodents, and are capable of producing a highly concentrated urine (salt concentration 2859 mmol/L, which is about four times as concentrated as seawater). Despite the high concentration of the urine, large quantities of urine have to be excreted each day (up to 25 ml/day) because of the high intake of water. This method of maintaining a positive water balance (and keeping cool in the desert during the daytime) is quite different to any other desert animal: it is a ‘large water intake with large water loss’ method, the large water loss being necessary because of the salty diet. This unusual method enables sand rats to exploit food that is inedible to other species, and to be diurnal when the ambient temperature is high (Ben ChaouachaChekir et al. 1983, Kam & Degen 1989). Fat Sand Rats are diurnal and nocturnal. Activity above ground is usually 09:00h to 17:00h in winter, but confined to early morning and late afternoon in summer (Petter 1961, Ilan & Yom-Tov 1990). During the cooler months of the year, they bask in the sun.

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Psammomys obesus

Extensive complex burrows are excavated under bushes. Burrows are mostly less than 0.5 m deep but several metres in length, and have many entrances (usually 5–15 in Tunisia). A typical colony of Fat Sand Rats has a burrow under nearly every bush, with a maze of trails connecting burrows and food plants (Osborn & Helmy 1980). An actively used burrow can be recognized by fragments of saltbushes, faeces and urine marks at the entrance (Fichet-Calvet et al. 1999b). A chamber within the burrow may be used for faeces (Petter 1961). Burrows in salt marshes are abandoned in winter if flooded by water. An animal may use several burrows depending on climate and food resources, and lactating !! sometimes carry their offspring from one nest burrow to another. Foraging and Food Herbivorous. Leaves and twigs of many species of saltbushes are the principal food. Fat Sand Rats forage 10–30 m around their burrows. They climb into low saltbushes and cut little branches, which they carry back to the main entrance of their burrows. The succulent leaves and twigs are eaten at the burrow entrance, where piles of discarded pieces are dropped onto the sand. Fragments of many species of plants are found in and near burrows: in Egypt, one burrow contained nine species of saltbush and another contained five species (Wassif 1953). Although food gathering does not take long, chewing and ingesting may take up to 4 h/day. Preferred plants vary depending on the locality, and include Salsola vermiculata (76.6% of the diet) in Morocco (Zaime & Gautier 1989), Suæda mollis, Traganum nudatum and Salsola fœtida in Algeria (Daly & Daly 1973), Suæda fruticosa, Arthrocnemum glaucum and Salsola tetragona in Tunisia (Fichet-Calvet et al. 2000) and Suæda monoïca, A. glaucum and Anabasis articulata in Egypt (Wassif & Soliman 1979). Fat Sand Rats need to eat large quantities of food each day in order to provide adequate water for their metabolism (see above). Occupied burrows are positively correlated with the presence of green and vigorous Chenopod bushes (Daly & Daly 1974, Zaime & Pascal 1989). Social and Reproductive Behaviour Although Fat Sand Rats live in colonies, individuals tend to be solitary. Females have smaller range lengths (mean RL = 76 m) than "" (mean RL = 190 m). Only a small part of the range is used at a time: on a weekly basis, !! have a mean RL of 12 m and then move to an adjacent part of the range. Males have a mean weekly RL of 68 m; the range of a " overlaps that of several !! so each ! is visited sequentially. Fat Sand Rats have to move (or ‘drift’) from one part of the range to another as they ‘eat out’ one patch of saltbushes and move on to another. Range lengths of juveniles when they are dispersing are larger than those of adults (e.g. juvenile "", mean 233 m; juvenile !!, mean 208 m) (Daly & Daly 1975b). Adult "" tend to be aggressive to juvenile "", chasing them from their home-range (Daly & Daly 1975b). Fat Sand Rats make sonic and ultrasonic squeals, associated sometimes with foot-drumming, when a conspecific is nearby. Predator warning squeals are brief, and produced close to the burrow (Bridelance 1989). Faeces and urine are used for marking the home-range (Fichet-Calvet et al. 1999b). Reproduction and Population Structure Pregnancies occur during the cooler dry months of the year (Sep–Apr) with peaks of births in Oct and Feb (Tunisia: Fichet-Calvet et al. 1999a; Egypt:

Osborn & Helmy 1980). In years of low rainfall and low food availability, reproduction is restricted to Jan–Apr. Gestation: 24 days. Litter-size: 4.8 (2–8, n = 34; Fichet-Calvet et al. 1999a). Mean litter-size shows seasonal variation: 3.6 (n = 26) in Sep, and 6.0 (n = 18) in Jan–Mar. Weight at birth: 6–7 g. Weaned: Day 15. Adult size: Day 120. Sexual maturity: 3–6 months according to the season of birth. Longevity: 14–18 months (in field), 6 years (in captivity) (M. Kam unpubl.). Interval between litters: 35–44 days. In Tunisia, age structure of population varies seasonally. In Sep (beginning of reproductive season), population is composed entirely of adult animals; these live and reproduce until the following Mar. Proportion of young increases from Oct to Mar (to max of 72% juveniles). In Mar, breeding population consists of two cohorts: a few old multiparous adults and some young primiparous young adults (born earlier in the season). Most of multiparous adults die at the end of breeding season (Daly & Daly 1975b, Amirat et al. 1977, Fichet-Calvet et al. 1999a). Predators, Parasites and Diseases Predators include foxes (Vulpes vulpes), dogs, snakes (Malpolon monspessulanus) and raptors (Buteo rufinus, Tyto alba, Bubo bubo, Strix aluco and Athene noctua). Remains of Sand Rats have been found in pellets of Eurasian Eagle-owls Bubo bubo or Desert Eagle-owls B. ascalaphus (0.6–6% occurrence) in Morocco and Algeria (M. Thévenot pers. comm.). Occurrence in owl pellets is sometimes high (50%, S. Aulagnier pers. comm.). Fat Sand Rats are principal reservoir of a protozoa, Leishmania major, causing zoonotic cutaneous leishmaniasis in humans (WHO 1990). During the sylvatic plague cycle, the parasite circulates between the sand rats and sandflies living in the burrows. Bites of sandflies to humans cause cutaneous lesions. Population explosions of Fat Sand Rats increase the risk of transmission of this disease (70,000 cases in Tunisia from 1982 to 2000). Other parasites recorded from Sand Rats include bacteria (Bartonella spp., Borrelia spp.), Protozoa (Babesia spp.) and nematode and cestode worms. It is an important laboratory animal for studying disease such as diabetes. Conservation IUCN Category: Least Concern. Fat Sand Rats are common and widespread and not threatened. Control measures may be necessary when population numbers are high. Measurements Psammomys obesus HB (""): 161.4 (116–185) mm, n = 228 HB (!!): 156.9 (133–183) mm, n = 200 T (""): 116.7 (88–140) mm, n = 219 T (!!): 115.6 (92–135) mm, n = 187 HF (""): 34.5 (33–36) mm, n = 59 HF (!!): 34.0 (32–36) mm, n = 44 E (""): 15.5 (14–17) mm, n = 41 E (!!): 14.9 (13–16) mm, n = 42 WT (""): 157.6 (82–237) g, n = 229 WT (!!): 141.7 (83–220) g, n = 200 GLS (""): 40.9 (36.9–43.0) mm, n = 40 GLS (!!): 40.4 (37.1–43.4) mm, n = 59 GWS (""): 24.2 (21.4–26.2) mm, n = 40 GWS (!!): 23.4 (21.4–25.4) mm, n = 57 345

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M1–M3: 6.6 (6.0–7.4) mm, n = 100 Auditory bulla: 13.3 (12.3–14.4) mm, n = 73 Tunisia (40 km south of Sidi Bouzid; E. Calvet unpubl.) Auditory bulla: Egypt (Osborn & Helmy 1980)

Key References Daly & Daly 1974, 1975b; Fichet-Calvet et al. 1999a, b, 2000; Petter 1961. E. Fichet-Calvet

Psammomys vexillaris PALE SAND RAT (LESSER SAND RAT) Fr. Rat des sables pâle; Ger. Dünne Sandratte Psammomys vexillaris Thomas, 1925. Ann. Mag. Nat. Hist., ser. 9, 16: 198. Bondjem, Libya.

Taxonomy Although vexillaris is sometimes considered synonymous with obesus, Ranck (1968) and Cockrum et al. (1977) consider, on the basis of skull morphology and chromosome numbers, that vexillaris is a valid species. Kowalski & Rzebik-Kowalska (1991) placed vexillaris as a synonym of P. obsesus and therefore did not recognize this species in Algeria. Musser & Carleton (2005) recognize P. vexillaris pending revision of geographic variation in P. obsesus. Synonyms: edusa. Subspecies: none. Chromosome number: 2n = 46, FN = 78.

Geographic Variation

Description Medium-sized gerbil, similar in form to P. obsesus but smaller and paler. Dorsal pelage gold. Flanks and ventral pelage cream (or white). Ears small, hairy; postauricular patch absent. Foreand hindlimbs short with white hairs on the inner surfaces. Soles of feet partly haired, claws dark. Tail long (ca. 86% of HB), hairy with a terminal pencil. Skull comparable to P. obesus but smaller; auditory bullae inflated (ca. 39% of GLS); upper incisors smooth, without groove. Nipples: not known.

Distribution Endemic to Africa. Mediterranean Coastal and Sahara Arid BZs close to the Mediterranean Sea in E Algeria to W Libya. Recorded near Biskra, E Algeria (Thomas 1925); near Tozeur, Tunisia (Cockrum et al. 1977); and near Tripoli, Libya (Thomas 1925, Ranck 1968). Distribution of P. vexillaris is totally within the distribution of P. obesus.

None recorded.

Similar Species Psammomys obesus. Usually larger, ventral pelage pale ochre; hindlimb buffy on inner surface; Mauritania and Morocco to Egypt and Sinai. Meriones spp. Larger; dorsal pelage darker but with white ventral pelage; ear larger; sympatric throughout most of geographic range.

Habitat Poorly known: alluvial soils on roadsides and hillocks (Ranck 1968). Inhabits sandier substrates than P. obesus (Cockrum et al. 1977). Abundance Remarks

Probably rare; known only from a few specimens.

Apparently no other information available.

Conservation

IUCN Category: Data Deficient.

Measurements Psammomys vexillaris HB: 122 (115–130) mm, n = 7 T: 106 (80–120) mm, n = 7 HF: 31 (30–35) mm, n = 7 E: 11 (10–12) mm, n = 7 WT: n. d. GLS: 34.8 (33.0–37.0) mm, n = 7 GWS: 21.5 (19.6–23.1) mm, n = 5 M1–M3: 5.7 (5.2–5.9) mm, n = 7 Auditory bulla: n. d. Nefta, Tunisia (MNHN) Key References Kowalska 1991. Psammomys vexillaris

Cockrum et al. 1977; Kowalski & RzebikE. Fichet-Calvet

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Sekeetamys calurus

GENUS Sekeetamys Bushy-tailed Jird Sekeetamys Ellerman, 1947. Proc. Zool. Soc. Lond. 117: 271. Type species: Gerbillus calurus Thomas, 1892.

A monotypic genus distributed mainly in the Middle East with its western boundary in Egypt.The single species in the genus (S. calurus) occurs only in rocky habitats in semi-arid and arid environments. See species profile for further details.

The status of the genus is uncertain. Although the type for the genus was described as a Gerbillus, it was placed in a new genus, Seeketamys, by Ellerman (1940). Other authorities have considered calurus as a species of Gerbillus, Dipodillus or Meriones.The genus is phylogenetically close to Meriones (Pavlinov et al. 1990) and also to Microdillus and Gerbillus (Tong 1989). The genus shows extensive chromosomal rearrangements, more so than in related genera, as well as special morphological features; on cytogenetic and allele evidence, it is closely related to Psammomys and Meriones, and less closely to Desmodillus (Qumsiyeh & Chesser 1988). The distinguishing characters of the genus are the long very hairy tail, long narrow hindfeet, naked soles, large auditory bullae and narrow interorbital constriction.The teeth are intermediate between Gerbillus and Meriones. The single species is Seeketamys calurus. See also Table 27. Christiane Denys

Sekeetamys calurus.

Sekeetamys calurus BUSHY-TAILED JIRD Fr. Gerbille à queue touffue; Ger. Bilchrennmaus Sekeetamys calurus (Thomas, 1892). Ann. Mag. Nat. Hist., ser. 6, 9: 76. Tor, Sinaï, Egypt.

Taxonomy Originally described in genus Gerbillus. Darker individuals from the Eastern Desert of Egypt described as a separate species, S. mackrami (Setzer 1961), are now recognized as a subspecies (Osborn & Helmy 1980). Synonyms: mackrami. Subspecies: two. Chromosome number: 2n = 38, aFN = 70 (Qumsiyeh & Chesser 1988). Description Medium-sized rodent with long bushy black tail, with white at tip. Pelage long (ca. 18–20 mm), soft, fine and dense. Dorsal pelage pale brownish-yellow to sandy-buff, speckled with black; hairs grey at base, with yellowish subterminal band and black tip. Flanks paler, with yellow or orange line from wrist to ankle. Ventral pelage pure white. Head similar in colour to back. Eyes large, dark. Ears large, darkly pigmented, rounded at tip, with sparse short hairs. Very long coarse vibrissae (up to 60 mm), mostly black, some white. Hindfeet long and narrow; upper surface with dense white hairs; soles naked. Tail very long (ca. 120% of HB), bushy throughout its length, densely covered with long black hairs, white at tip; tail is unlike that of any other species of gerbil. Skull: auditory bullae inflated, supraorbital and cranial ridges conspicuous, zygomatic arches compressed, rostrum long and narrow, interorbital constriction narrow, upper incisors opisthodont each with single longitudinal groove, molar teeth prismatic (Figure 56, see also Table 27). Nipples: 2 + 2 = 8.

Figure 56. Skull and mandible of Sekeetamys calurus (BMNH 12.11.19.1).

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Geographic Variation S. c. calurus: Sinai Peninsula, Egypt. Paler dorsal pelage, sometimes with broad dark mid-dorsal stripe. S. c. mackrami: Eastern Desert (between Nile R. and Red Sea coast), Egypt. Darker dorsal pelage with narrow dark mid-dorsal stripe. Similar Species Eliomys melanurus. Black bushy tail without white tip, black eye-ring and black stripe from back of eye to base of ear, auditory bullae only slightly inflated; mesic and semi-arid habitats. Meriones spp. Sparsely haired pale tail with black pencil at tip, auditory bullae inflated but larger than in Seeketamys; semi-arid and arid habitats. Distribution Sahara Arid BZ. Recorded from Egypt (Eastern Desert and Sinai) and extreme NE Sudan. Extralimitally recorded from S Israel, Jordan and Saudi Arabia. Habitat Rocky areas, sandstone cliffs, crevices in granite and amongst boulders in arid regions; also mountain regions in Sinai. Not recorded from sandy areas. Sekeetamys calurus

Abundance Uncertain; recorded as moderately abundant in Israel (Zahavi & Wahrman 1957). In Negev Highlands, Israel, density is 0.8–3.0 ind/ha; populations tend to be stable throughout the year (Shenbrot et al. 1999a). Adaptations Nocturnal. Very agile; climbs rapidly on boulders and rock faces. During day rests in crevices of rocks and under boulders; does not dig burrows. At night, active for about 10.5 hours (Degan et al. 1986). During locomotion, the tail is held upright and curved forward in a squirrel-like fashion, and at times locomotion is bipedal. In Egypt, coexists with other rock-adapted species including Eliomys quercinus, Acomys cahirinus, A. russatus and Gerbillus dasyurus (Osborn & Helmy 1980). Metabolic rate (as measured by oxygen consumption) is 31% (Degan et al. 1986) to 44% (Haim & Borut 1986) below predicted level at the thermoneutral zone (Ta = 34–36 °C) and variable depending on environmental conditions and temperature in different habitats. Food consumption is low (when compared with non-arid species of similar size). The dense pelage provides insulation against cold when active at night, yet thermal conductance is higher than expected (in spite of the dense pelage) enabling dissipation of body heat when Ta is high. Studies on water relations indicate that water flux was 99% of that predicted, and that S. calurus does not show any special adaptations for water conservation. The combination of a high proportion of insects in the diet, low food intake, relatively low metabolic rate and nocturnal activity (even though Ta is cold at night during some seasons) enables survival in an arid environment where resources are scarce (Degan et al. 1986, Haim & Borut 1986); this strategy for survival is quite different to that of granivorous desert rodents, which eat a low-water diet and exhibit many waterconservation abilities. Foraging and Food Omnivorous. Detailed analysis of diet not available. Individuals trapped in crevices that contained parts of seeds and seed capsules, as well as bits of succulent plants. Insects

probably supply 87% of food in wild-living individuals (Degan et al. 1986), although stomach contents of specimens (from Saudi Arabia) contained only green plant material (Nader 1974 ). Captive individuals ate cockroaches and crickets (Osborn & Helmy 1980). Social and Reproductive Behaviour Mobility in "" greater than !!. In "", mean distance between captures was 57 m (max 332 m); in !! 20 m (max 72 m). Adult "" have larger home-ranges (ca. 10–23 ha) than adult !! (0.7–0.8 ha) (Shenbrot et al. 1999a). Reproduction and Population Structure Flower (1932) reported litters from captive animals every month of the year in Egypt except Sep. Litter-size (in captivity): 2.8 (n = 47 litters; max 6/litter [n = 2]). In the wild, period of reproduction appears to be limited and variable, and dependent on local conditions; longevity is probably high and reproductive success is low in natural conditions (Shenbrot et al. 1999a). Sex ratio of individuals captured in the Eastern Desert was 13 "" : 11 !! (Osborn & Helmy 1980). Predators, Parasites and Diseases Preyed on by Eurasian Eagle-owls Bubo bubo at Gebel Migif, Egypt; remains of ten individuals found in pellets (n = 45 small mammals) (Goodman 1986). Four species of fleas recorded – Xenopsylla nubica, X. dipodilli, X. conformus and Nesopsylla theodori – from individuals near St Catharine’s Monastery, Sinai, Egypt (Hoogstraal & Traub 1965a). Some of these species of fleas are also recorded from Meriones crassus. Conservation

IUCN Category: Least Concern.

Measurements Sekeetamys calurus calurus HB: 118.9 (98–128) mm, n = 25 T: 144.4 (131–164) mm, n = 20 HF: 33.1 (31–35) mm, n = 25

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E: 21.4 (20–23) mm, n = 25 WT: 41.4 (26.6–49.8) g, n = 17* GLS: 35.8 (34.5–37.4) mm, n = 20 GWS: 18.2 (17.2–19.1) mm, n = 10 M1–M3: 5.2 (4.8–5.8) mm, n = 17 Auditory bulla: 11.7 (11.0–12.7) mm, n = 25

Egypt (Osborn & Helmy 1980) * = S. c. makrami Key References

Harrison & Bates 1991; Osborn & Helmy 1980. Christiane Denys

GENUS Taterillus Taterils (Gerbils) Taterillus Thomas, 1910. Ann. Mag. Nat. Hist., ser. 8, 6: 222. Type species: Gerbillus emini Thomas, 1892.

Taterillus emini.

The genus Taterillus is endemic to Africa and contains eight species (Table 28). It has a widespread distribution, and occurs in the Sahel, Sudan and Guinea Savanna BZs from Mauritania to Somalia, from the Tilemsi Valley in N Mali to SW Nigeria, and from C Sudan to C Kenya. Typical habitats range from sandy dunes, thorny scrubs and woodland savannas, as well as fields, gardens and even human dwellings. Three species have fairly wide distributions (T. gracilis in West Africa, T. congicus in central Africa and T. emini in East Africa); the others mostly have very restricted distributions. Species in the genus are small to medium-sized gerbils. Dorsal pelage is pale yellow to reddish-brown; ventral pelage, hands and feet are white. The head is characterized by a pointed muzzle, large eyes and elongated ears. The tail is longer than head and body, with soft hairs on its whole length, and a long terminal pencil of darker hairs. The hindfeet are long, naked and the soles are dark. The genus differs from Gerbilliscus (formerly Tatera) by overall smaller size, a more gracile appearance, elongated anterior palatal foramina, which end well forward of M1, and longer slit-like posterior palatal foramina. It differs from Gerbillus by overall larger size, larger ears and longer and naked soles on the hindfeet; the skull is more robust with longer posterior palatal foramina and laminated molars without longitudinal crests connecting tooth cusps (Figure 57). These gerbils are nocturnal and terrestrial. They dig burrows of various depth and complexity, and feed on seeds, stems, leaves and insects.They are often quite common, and sometimes subject to local

Figure 57. Skull and mandible of Taterillus gracilis (HC 1079).

population explosions that make them potential pests in agricultural fields. Cycles of abundance, as well as some eco-ethological and physiological attributes, are known for T. gracilis and T. pygargus in Senegal and for T. petteri in Burkina. The taxonomy of the genus depends heavily on chromosomal information, as most species are very similar morphologically. Chromosome numbers are species-specific and vary from 2n = 14/15 in T. tranieri to 2n = 54 in T. congicus. One current problem concerns the characterization of T. harringtoni relative to T. emini: a karyotype of 2n = 44 chromosomes has been attributed to both, and further studies are needed to clearly diagnose these two species; here, harringtoni is considered as a synonym of T. emini. Further cytotaxonomic investigations will undoubtedly lead to the discovery of new biological species, as suggested by the very complex chromosomal evolution of this group. The genus can be divided in two distinct lineages as some species are characterized by a strong chromosomal synapomorphy, i.e. a double autosome–gonosome translocation. The males of one lineage (T. arenarius, T. gracilis, T. petteri, T. pygargus, T. tranieri) are 349

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Table 28. Species in the genus Taterillus. Arranged in alphabetical order. Species T. arenarius T. congicus T. emini T. gracilis T. lacustris T. petteri T. pygargus T. tranieri

Chromosome number

Dorsal pelage

Notes

2n = 30 (""), 2n = 31 (!!) 2n = 54 2n = 44 2n = 36 (!!), 2n = 37 ("") 2n = 28 (!!), 2n = 29 ("") 2n = 18 (!!), 2n = 19 ("") 2n = 22 (!!), 2n = 23 ("") 2n = 14 (!!), 2n = 15 ("")

Pale sandy-yellow Chocolate-brown – reddish-brown Reddish-brown – orange Reddish-yellow Tawny Pale orange – cinnamon Yellow, reddish-brown Buffy-brown

Mauritania (Mali, Niger) Central African Republic, Chad (Cameroon, Sudan, Uganda) Sudan and Sahel BZs of eastern Africa, Somalia–Masai BZ Sudan and Sahel BZs of western Africa; widespread Area around L. Chad Sudan Savanna BZ (Burkina, Mali, Niger) Sudan Savanna BZ (Senegal, Niger) Sahel Savanna BZ (Mali)

characterized by a set of three (X,Y1,Y2) sex chromosomes, whereas been said to be of help, a result that was refuted by recent studies on the males of the other lineage (T. congicus, T. emini/harringtoni, T. karyotyped specimens. lacustris) are characterized by the classical XY sex chromosomes. The eight species are distinguished primarily on chromosome (Females retain the usual XX sex chromosomes.) Again, the only number but also on size and distribution (see also Table 28). non-ambiguous character to distinguish between species in the genus is the karyotype. Multivariate analyses of morphometric data have Laurent Granjon & Gauthier Dobigny

Taterillus arenarius SAND TATERIL (ROBBINS’S TATERIL) Fr. Tatérille des sables; Ger. Sahel-Rennmäuschen Taterillus arenarius Robbins, 1974. Proc. Biol. Soc. Wash., 87: 399. Tiguent, Trarza Region, Mauritania.

Taxonomy Chromosome number distinguishes this species from its sibling species T. gracilis, T. petteri, T. pygargus and T. tranieri (Matthey 1969, Petter 1970,Volobouev & Granjon 1996, Dobigny et al. 2003). Synonyms: none. Chromosome number: 2n = 30 (!!), 2n = 31 (""). Description Medium-sized gerbil. Dorsal pelage pale sandyyellow; hairs grey at base. Ventral pelage white, clearly delineated from dorsal pelage on flanks. Cheeks white, with white supraorbital and postorbital patches. Muzzle pointed, often with dark markings on upper part of nasal region. Large elongated ears. Large eyes. Foreand hindfeet white. Hindfeet relatively long; soles dark and naked or nearly so; three median toes of similar length, Digit 1 short; rather long claws. Tail long (ca. 130–140% of HB) covered by short hairs, long terminal pencil of dark blackish-brown hairs. Nipples: 2 + 2 = 8. Geographic Variation None recorded. Similar Species T. gracilis. Similar in morphology and size; chromosome number: 2n = 36/37; widely distributed in West Africa. T. petteri. Similar in morphology and size; chromosome number: 2n = 18/19; Mali, Niger and Burkina mostly south of Niger R. T. pygargus. Similar in morphology and size; chromosome number: 2n = 22/23; Senegal and Niger. T. tranieri. Similar in morphology and size; chromosome number: 2n = 14/15; only in S Mauritania and W Mali. Distribution Endemic to Africa. Sahel Savanna BZ. Presence confirmed (by chromosome number) only in Mauritania (Matthey 1969, Petter 1970,Volobouev & Granjon 1996). Recorded eastwards to Mali and Niger by Robbins (1974) on the basis of morphological

Taterillus arenarius

characters, but not yet confirmed by karyology (L. Granjon unpubl., Dobigny et al. 2002a). Habitat Sandy-clay plains and inland dunes (Granjon et al. 1997b). Trapped in dry areas where average annual rainfall does not exceed 400 mm. Abundance

Densities probably low (Granjon et al. 1997b).

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Taterillus congicus

Remarks Found in pellets of Barn Owls Tyto alba; 10% of HF) on Digit 3 of hindfoot; claws of hindfeet >10% of hindfoot. Tail short (ca. 30% of HB), mostly naked except for a few short bristles, tapering towards tip. Skull: incisors pro-odont, forming semicircle on cutting edge, single faint groove on anterior face of each incisor; interorbital constriction very conspicuous; slight supraorbital ridges; palate extends well posterior to the posterior end of cheekteeth (more so than in other murid rodents); M1 and M2 with lamina-like transverse rows; t1 on M1 absent (Figure 58). The skull is similar in form to that of Lophuromys (due to convergence), although detailed structure suggests affinities to the Dendromurinae. (Description based on original description and on Rosevear 1969.) Geographic Variation None recorded. Similar Species Steatomys spp. Similarly very short tail (ca. 50% of HB); four digits on forefoot; incisor teeth orthodont/opisthodont; zygomatic plate very small; zygomatic arches flared; palate short, not extending far behind the cheekteeth; tendency to become fat at certain times of year; widespread geographic distribution. Distribution Endemic to Africa. Northern Rainforest–Savanna Mosaic. Recorded only from the type locality in Togo (see also below). Habitat The type locality was described as in the ‘high forest belt’ (Rosevear 1969). However, the only known locality for this species is 08° 11´ N, 00° 41´ E, which is in the Rainforest–Savanna Mosaic where patches of riverine and relict forest alternate with grassland and woodland savanna. Rosevear also mentions that the generic name comes from the Greek Lima (garden or meadow) and mys (mouse), which suggests that the habitat may be dense grasslands close to riverine forest where the soil is moist and suitable for digging (see Remarks) – a habitat similar to that of Lophuromys spp. Abundance Known only by the two type specimens. Remarks Virtually nothing is known about this species. However, the morphology of the feet suggest that it is terrestrial and burrowing, and the very short tail suggests that it is not a climbing mouse like Dendromus spp. The single stomach was full of termites (Dieterlen 1976c). These limited observations suggest that it is an insectivore that probably gathers its prey by scratching in the soil and litter with its long claws – in much the same way as do Lophuromys spp.

Leimacomys buettneri

Conservation IUCN Category: Data Deficient. Likely to be ‘Critically Endangered’ because of very limited range. No individuals of this species have been encountered since the type specimens were obtained more than 100 years ago. Two extensive surveys in Togo during the 1960s failed to find any evidence of this species, and none has been found in recent years. On this evidence, Schlitter (1989) considers it may be extinct. Measurements Leimacomys buettneri HB: 118 mm T: 37 mm HF: 23 mm E: 14 mm WT: ca. 30 g GLS: 30.3 mm GWS: 15.6 mm M1–M3: 4.9 mm Bismarkburg, Togo Body measurements: holotype (Matschie 1893) Skull measurements: Dieterlen 1976c Key References Denys 1993; Dieterlen 1976c; Rosevear 1969. Fritz Dieterlen

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Subfamily MURINAE – Rats and Mice Murinae Illiger, 1811. Abhandl. K. Akad. Wiss. Berlin for 1804–11, p. 46, 129. Aethomys (11 species) Apodemus (1 species) Arvicanthis (7 species) Colomys (1 species) Dasymys (5 species) Dephomys (1 species) Desmomys (2 species) Grammomys (11 species) Heimyscus (1 species) Hybomys (6 species) Hylomyscus (8 species) Lamottemys (1 species) Lemniscomys (11 species) Malacomys (3 species) Mastomys (8 species) Muriculus (1 species) Mus (20 species) Mylomys (2 species) Myomyscus (3 species) * Nesokia (1 species) Nilopegamys (1 species) Oenomys (2 species) Pelomys (5 species) Praomys (16 species) Rattus (2 species) Rhabdomys (1 species) Stenocephalemys (4 species) Stochomys (1 species) Thallomys (4 species) Thamnomys (3 species) Zelotomys (2 species)

Veld Rats Long-tailed Field Mouse Grass Rats Water Rat Shaggy Rats Defua Rat Scrub Rat Thicket Rat Smoky Mouse Forest Mice Wood Mice Mount Oku Rat Grass Mice Swamp Rats Multimammate Mice Ethiopian Striped Mouse Mice, Pygmy Mice Mill Rats (Three-toed Grass Rats) Meadow Mice Short-tailed Bandicoot Rat Ethiopian Water Rat Rufous-nosed Rats Creek Rats Soft-furred Mice Rats Four-striped Grass Mouse Ethiopian Rats Target Rat Acacia Rats (Tree Rats) Thicket Rats Broad-headed Mice

p. 362 p. 377 p. 379 p. 389 p. 392 p. 400 p. 402 p. 404 p. 418 p. 420 p. 429 p. 439 p. 441 p. 455 p. 460 p. 472 p. 473 p. 499 p. 502 p. 506 p. 508 p. 509 p. 513 p. 519 p. 539 p. 544 p. 547 p. 554 p. 556 p. 563 p. 567

*Formerly Myomys.

The Murinae forms the largest assemblage of species within either Muridae or the larger Muroidea and exhibits the most expansive geographic range. Totalling about 124 extant genera and 543 living species, the subfamily is indigenous to Eurasia, the Middle East, Arabian Peninsula, Indomalayan region, Japan, Ryukyu Islands, Philippine Islands, the numerous archipelagos stretching from the Sunda Shelf to the New Guinea and Australian region, and Africa (Musser & Carleton 2005). The subfamily is represented in Africa by 31 genera and 145 species. African endemics (27 genera, 139 species) comprise about 20% of all murine taxa. Most species of Arvicanthis and Myomyscus are also restricted to Africa, but a single species of each (A. niloticus and M. yemeni) is also found on the Arabian Peninsula. Of the 39 species of Mus, 17 (44%) are African endemics; some researchers view these sub-Saharan endemics, currently arranged as the subgenus Nannomys, as a separate evolutionary radiation that should be recognized as a genus. Mus spretus (subgenus Mus), native to the western Mediterranean region of south-western Europe, also occurs in the Maghreb of North Africa where, judged by late Pleistocene samples, it may have originated (Dobson 1998, 2000). Two genera, Apodemus and Nesokia, have large geographic

ranges outside of Africa and only marginal distributions on the continent. Apodemus sylvaticus, one of the 20 Palaearctic species of Apodemus, reaches the Mediterranean fringe of North Africa (Kock & Felten 1980), which may represent an accidental introduction by humans followed by subsequent spread throughout Mediterranean coastal habitats (Dobson 1998, 2000, Michaux et al. 2002). Nesokia indica, indigenous to western Asia, the Middle East and the Arabian Peninsula, also occurs in north-eastern Egypt, and Pleistocene fossils from Egypt and northern Sudan point to a formerly broader African distribution (Osborn & Helmy 1980). Three non-native murines – Mus musculus, Rattus norvegicus and R. rattus – have invaded parts of the continent, but have not been able to invade natural communities. They are mostly restricted to humanized environments such as cities, towns, villages and food stores. Although their initial evolution and original distribution are rooted in Eurasia, they are now found in many parts of the world and are members of a small cluster of murines (four genera and 14 species, possibly including Apodemus sylvaticus; see above) that have expanded their distributions far from their natural ranges through intentional or accidental processes associated with human migration and settlement (Musser & Carleton 2005). Also, the Indomalayan Bandicota bengalensis was introduced to Patta Island, Kenya, but whether a population became established is unknown (Corbet & Hill 1992). No other subfamily of Muroidea, or even Rodentia, has such a relatively large contingent of species whose present geographic distributions have been mediated by anthropogenic activities. Living native African murines range in body size from small (e.g. most species of Mus) to large (e.g. some species of Arvicanthis and Aethomys). No endemic very large-bodied or giant murines are present. Variation in body form reflects terrestrial (e.g. Arvicanthis), scansorial (e.g. Hylomyscus), arboreal (e.g. Thallomys) and limnetic (e.g. Colomys) adaptations. Although some species may excavate burrows or shallow places to nest, none exhibits the morphological and physiological adaptations associated with muroids that are extremely fossorial (e.g. Tachyoryctes and Spalax in Spalacidae). Most African murines are nocturnal, some are diurnal; and they fill a variety of trophic niches, from omnivorous, herbivorous and granivorous to insectivorous and carnivorous. The bulk of endemic murine species occurs in the sub-Saharan biotic zones, inhabiting deserts, grasslands, savannas, rainforests and afromontane forests. Only three genera of typically subSaharan murine rodents are represented in Africa north of the Sahara: Lemniscomys barbarus (a Maghreb endemic), Mastomys erythroleucus (primarily sub-Saharan with an isolated population in west-central Morocco) and Arvicanthis niloticus (also primarily sub-Saharan with populations in Egypt and Sudan; Musser & Carleton 2005). The Murinae is characterized by a cohesive suite of morphological traits (Carleton & Musser 1984), but derived molar conditions form the cardinal basis for defining the subfamily. Two neomorphic cusps, the anterostyle (t1) and enterostyle (t4), are present on the lingual border of the upper first molar and form two chevron-shaped, transverse lamina; both upper and lower molars lack longitudinal enamel crests between lamina; and cusps on the lower molars are positioned opposite one another (Flynn et al. 1985, Jacobs et al. 1989, 361

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Freudenthal & Martin Suárez 1999). Other derived cranial features include their modified carotid circulatory pattern (sphenofrontal foramen and squamosal-alisphenoid groove absent; stapedial foramen present) and a reduced tegmen tympani that does not contact the posterior squamosal (Bugge 1970, Carleton & Musser 1984 and unpubl.). The monophyly of Murinae is additionally supported by phylogenetic analyses of mitochondrial and nuclear genes (e.g. Verneau et al. 1997, 1998, Michaux et al. 2001, Debry 2003, Jansa & Weksler 2004), although the number of genera sampled to date is relatively few and in some studies includes only Mus and Rattus. Such studies identify Gerbillinae and Deomyinae as the closest relatives of Murinae (Martin et al. 2000, Michaux et al. 2001, Adkins et al. 2003, Jansa & Weksler 2004).The three subfamilies may have diverged from an ancestral muroid stock about 20.8–17.9 mya (early Miocene), an estimate derived from molecular-clock assumptions (Michaux et al. 2001). By late Miocene, but not earlier, representatives of each subfamily are recorded from African sediments (Jaeger 1977b, Mein et al. 1993, Geraads 2001, Winkler 2001). While recognition of Murinae as a natural group is strongly supported, uncertainty over relationships among endemic African murines has produced two viewpoints to explain their diversity. Employing microcomplement fixation of albumin,Watts & Baverstock (1995a) identified an African murine clade separate from New Guinean, Australasian and South-East Asian evolutionary lineages, and suggested that living African murines represent ‘a period of rapid radiation from a single ancestor, beginning 8–10 mya and still continuing’. Mitochondrial and nuclear gene sequences have disclosed a large African murine clade (Aethomys, Arvicanthis, Dasymys, Desmomys, Grammomys, Hybomys, Lemniscomys, Mylomys, Pelomys and Rhabdomys) that partly accords with the phylogenetic grouping identified by Watts and Baverstock (Ducroz et al. 2001). The alternate view, that living African

murines represent a paraphyletic assemblage, draws support from DNA/DNA hybridization (Chevret 1994) and from other genesequence analyses (Lecompte 2003, Jansa & Weksler 2004). In addition to an Arvicanthis clade (containing the same genera as given by Ducroz et al. 2001), Lecompte identified Praomys (Colomys, Heimyscus, Hylomyscus, Mastomys, Myomyscus, Praomys, Stenocephalemys, and Zelotomys), Mus (species in Nannomys) and Malacomys (containing only species of Malacomys) lineages, all three distantly isolated from the Arvicanthis clade and from each other. The meagre number of Miocene taxa recovered from African sediments offer no resolution to whether the derivation of modern African murines is from a single ancestral group or from several ancestral groups, or to their origin as a result of one or more immigrations. Southern Asia is vaguely mentioned as the source area, with arrivals in Africa occurring in middle to late Miocene–Pliocene (Jacobs 1985, Winkler 1994, 2002). Late Miocene is the earliest documentation for Progonomys and Paraethomys in North Africa (Jaeger 1977b, Mein et al. 1993) and for Karnimata and Saidomys in Kenya (Winkler 2001). Murines from earlier Miocene strata are known only from northern Pakistan, including the middle Miocene Antemus, generally considered the earliest murine (Jacobs & Downs 1994, Freudenthal & Martin Suarez 1999). Fossils of living endemic African murines first appear in the Pliocene (Denys 1999, Winkler 2002). Although now largely confined to Africa, Mastomys and Arvicanthis are each extralimitally represented by a single species, now extinct, that lived in Israel during the Pleistocene (Tchernov 1968, 1996), and by an extinct Pliocene species on the Mediterranean island of Rhodes (De Bruijn et al. 1996). The subfamily is currently represented in Africa by 31 genera. Guy G. Musser & Michael D. Carleton

GENUS Aethomys Veld Rats Aethomys Thomas, 1915. Ann. Mag. Nat. Hist., ser. 8, 16: 477. Type species: Epimys hindei Thomas, 1902.

The genus Aethomys currently includes 11 species (Table 29), of which ten are endemic to East, central and southern Africa, and one species is endemic to West Africa (Musser & Carleton 1993, Chimimba 1998, Chimimba et al. 1999). An additional proposed species (A.

Aethomys kaiseri.

halleri) (see Denys & Tranier 1992) requires assessment. Species in the genus occur in a variety of savanna woodland and grassland habitats, preferring those that include shrubs or thick grass, hollow tree trunks and logs, creviced rocky areas, and/or termitaria. Some species may occur commensally with humans in agricultural areas. Although some are widely distributed (e.g. A. chrysophilus), others are restricted (e.g. A. stannarius, A. silindensis). Veld Rats are generalized medium- to large-sized murid rodents. Species in the genus vary in body size, body proportions and pelage colour. The genus is most similar to Rattus, from which it differs in having molars that are clearly cuspidate and anterior palatal foramina that extend posteriorly to between the molars (Figure 59). Although generally considered ‘long-tailed’, tail length varies among species (ca. 70–150% of head and body length). Pelage is short, sleek or rough, and the dorsal pelage is often ‘sunburnt’ (hence the Greekderived generic name: eithos = sunburnt; mys = mouse) in various shades of brown, grey, red and yellow. The hairs of the ventral pelage may be completely white, or grey at base with a white tip. Limbs are equal in size, the feet usually white above. Forefoot has four digits

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Table 29. Species in the genus Aethomys. Arranged in order of increasing mean head and body length. (n. d. = no data.) Species

HB mean (mm)

A. granti

111

A. namaquensis

113

A. chrysophilus

138

A. nyikae

T mean (mm) [% of HB]

Dorsal pelage

GLS mean (mm)

Chromosome number

Width M1

Nipples

Notes

30.0

2n = 32

n. d.

3 + 2 = 10

South Africa

31.3

2n = 24

n. d.

1+2=6

117 [97–113%] 154 [140%] 156 [120%]

Dull yellowish-brown to dark brown Yellowish-brown; black tips Reddish-brown

36.1

2n = 50

3.2

0+2=4

A. ineptus

147

162 [110%]

See A. chrysophilus

35

2n = 44

n. d.

n. d.

A. stannarius

148

Medium brown

36.4

n. d.

n. d.

1+2=6

Widespread in southern Africa Widespread. Kenya to South Africa NE Angola, S DR Congo, N Zambia, Malawi East Africa to Zambia and Angola South Africa. Morphologically indistinguishable from A. chrysophilus N Nigeria and N Cameroon

A. thomasi

150

Grizzled rufousbrown

26.0

n. d.

≥2.2

n. d.

W and C Angola

A. hindei

158

Medium brown

37.4

2n = 50

600 mm annual rainfall), while M. coucha has a preference for higher altitude/relatively drier parts of South Africa (Dippenaar et al. 1993, Venturi et al. 2003). The species is absent from natural habitats in the Rainforest BZ in central Africa, but has been recorded in human settlements in that zone. Many records of M. natalensis throughout Africa, and the associated biological information, are uncertain because of lack of karyological identification and the large geographic overlap with most other Mastomys species. Molecular identification of tissue samples should resolve the distribution of this species in the near future. Habitat Grassland, with or without bushes, thickets or trees, and disturbed patches. Very common in fields and fallow land as well as in and around buildings in human settlements. Does not occur in the rainforest but may be found in human settlements and agricultural fields within the Rainforest BZ. At the extremes of its geographic range may be limited to specific habitats, e.g. only inside villages in Senegal (Duplantier et al. 1990). It is a common pioneer species after fire (Meester et al. 1979). Abundance One of the commonest species in savanna habitats, with densities of up to 1000/ha in disturbed areas. In farmland and fallow areas in Tanzania, 85% of the captured small mammals were multimammate mice while in natural miombo woodland and moist grassland 50 km away, they comprised only 27% of the trapped small mammals in a community of 17 different species (Leirs 1995). The species is well known for its seasonal fluctuations in numbers and irregular population explosions: maximum population numbers may be up to 40 times minimum numbers (Leirs et al. 1996). In a natural thicket-grassland savanna in Malawi, multimammate mice comprised 47% of small rodents in a community of ten species on an annual basis; densities fluctuated from 1/ha (dry season) to 40/ha (wet season) (Happold & Happold 1991). Abundance may be patchy, large numbers occurring in (preferred) habitats while other nearby habitats have no (or very few) individuals. Often the commonest of all rodents in food stores and houses where they may be very abundant and are regarded as a serious pest. Adaptations Terrestrial and nocturnal. Activity is highest during the first half of the night, starting immediately after sunset. Multimammate mice can make their own burrows, but often use existing burrows or cracks in the soil. They can easily dig to a depth of more than 500 mm and can jump over 600 mm high (S. VibePetersen unpubl.). The nest is constructed underground, often not very deep, and is a simple cavity lined with dried plant parts torn to pieces. The digestive system is adapted to a generalist diet (Perrin & Curtis 1980). Subadult animals become reproductively active very quickly after a period of heavy rainfall (Leirs et al. 1993, 1994). Survival of subadult and adult animals is affected in a complex way by a combination of density-dependent and density-independent (rainfall) factors (Leirs et al. 1997a).

Foraging and Food Opportunistic omnivores. The diet includes seeds of cereals, leaves and stems of grasses and dicotyledonous plants, insects and sometimes carrion (Leirs et al. 1994). Contents of stomachs reported in the literature often relate to the kind of habitat where the studies were carried out, illustrating the opportunistic nature of foraging. Seeds often make up a large proportion of the stomach contents (Field 1975,Taylor & Green 1976). Multimammate mice dig up newly-planted maize seeds, causing huge loss of potential harvest. They also climb maize stalks and damage the cobs. They are generally considered a pest in agriculture throughout their geographic range. Social and Reproductive Behaviour Very lively, but generally not aggressive (although reported more aggressive than other Mastomys species). Home-ranges show a wide degree of overlap and there is no evidence of territoriality. For resident animals, home-ranges are rather small, ca. 1000 m2, larger for males than for females (Leirs et al. 1997a). At times of dispersal, animals can move several hundred metres in only a few hours. Reproduction and Population Structure Reproduction very seasonal, starting soon after the onset of the wet season and lasting well into the dry season. Germinating seed and young grass seedlings in the diet stimulate reproductive maturation (Leirs et al. 1994, Firquet et al. 1996). Mean litter-size varies widely between populations, e.g. 10–12 in Tanzania (with a reported maximum litter-size of 23) (Leirs 1995), 6.5 in Senegal (Duplantier et al. 1996) and 4.5 in E DR Congo (Rahm 1970). Gestation: 21–22 days. Postpartum oestrus a few days after parturition; mean litter interval 28 days during the breeding season. Copulatory plug only rarely formed (Johnston & Oliff 1954). Young altricial at birth; eyes open Day 15; weaned Day 21 (Baker & Meester 1977).Young animals rarely mature during the season of birth; they become reproductively active after abundant rainfall and vegetation growth in the next breeding season.When such rains come early, subadults may start reproducing at the age of three months. Most animals die towards the end of their first breeding season when not more than about 12 months of age. Sex ratio of subadults near 1:1, but strongly biased towards "" during the breeding season. Population structure is strongly influenced by the large litters, high reproductive rate and early mortality. In Malawi, the population in thicket–grassland savanna was composed primarily of subadults at end of dry season (when the population is at its lowest – see above); most of these subadults matured during the early wet season. Young began to enter the population towards the end of the wet season and early dry season, and soon formed the majority of the population as the adults died. By mid-dry season all young had become subadults. Population turnover was rapid due to dispersal of young, immigration of new animals from elsewhere and mortality of animals of all agegroups; individuals rarely live for more than 12 months (Happold & Happold 1991). Predators, Parasites and Diseases Multimammate mice are a common prey of many owls, raptors, snakes and small grounddwelling carnivores. Predation is an important source of mortality. The mice adapt their foraging behaviour and become more cautious in open spaces when the presence of raptors in the environment increases (Mohr et al. 2003). Attracting or excluding avian predators 469

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has complex and sometimes compensatory effects on survival, reproduction and dispersal of the mice so there is no clear effect on the resulting population densities (Vibe-Petersen et al. 2006). Multimammate mice are the main reservoir for Lassa fever virus in West Africa, and also implicated in the epidemiology of bubonic plague and leptospirosis. Very little is known about diseases affecting the mice themselves. Conservation IUCN Category: Least Concern. One of the commonest small mammals in Africa. Considered to be a serious agricultural pest in many regions. Measurements Mastomys natalensis HB (!!): 108 (74–145) mm, n = 801 HB (""): 106 (66–171) mm, n = 751 T (!!): 108 (63–147) mm, n = 786 T (""): 106 (70–175) mm, n = 736

HF (!!): 22.3 (17–25) mm, n = 797 HF (""): 21.7 (16–25) mm, n = 747 E (!!): 17.2 (12–24) mm, n = 790 E (""): 16.9 (11–25) mm, n = 741 WT (!!): 37.5 (9–84) g, n = 799 WT (""): 35.3 (10–88) g, n = 752 GLS (!!): 28.5 (22.9–34.8) mm, n = 754 GLS (""): 28.1 (22.3–32.3) mm, n = 699 GWS (!!): 13.6 (10.9–16.9) mm, n = 747 GWS (""): 13.5 (10.9–16.4) mm, n = 702 M1–M3 (!!): 5.0 (4.4–5.6) mm, n = 803 M1–M3 (""): 5.0 (4.5–5.7) mm, n = 754 Morogoro, Tanzania (H. Leirs unpubl., RMCA) Key References Dippenaar et al. 1993; Duplantier et al. 1990; Granjon et al. 1996; Leirs 1995; Leirs et al. 1997a. Herwig Leirs

Mastomys pernanus DWARF MULTIMAMMATE MOUSE (DWARF MASTOMYS) Fr. Souris à mammelles multiples naine; Ger. Zwerg-Vielzitzenmaus Mastomys pernanus (Kershaw, 1921). Ann. Mag. Nat. Hist., ser. 9, 8: 568. Amala River, Kenya (= upper course of the Mara River, situated at 00° 58´ S, 25° 24´ E according to Misonne & Verschuren 1964).

Taxonomy Originally described in the genus Rattus. The generic position of this species has been questioned, and recent molecular evidence indeed suggests that it should no longer be considered a member of Mastomys but is more closely related to Hylomyscus (Lecompte et al. 2002b). Synonyms: none. Chromosome number: not known. Description Very small, grey mouse, smaller than all other Mastomys. Dorsal pelage darkish-grey with, in most specimens, an ochraceous stripe between flanks and ventral pelage. Ventral pelage grey, belly slightly washed with buff; hairs grey with white tip; small white chest patch in some individuals. Prominent spot of white hairs behind each ear. Tail long (85% of HB), well covered with short hairs (up to 1.5 mm). Typical Mastomys skull; mesopterygoid fossa triangular, very narrow on the posterior margin of the palatine and relatively wide towards the end (Van der Straeten 1999). Nipple number: not known. Geographic Variation None recorded. Similar Species M. natalensis. Much larger mean HB and T; tail longer as % of HB; ventral pelage dark grey; widespread distribution and common; nipples 12 × 2 = 24; 2n = 32. Distribution Endemic to Africa. Southern part of Somali– Masai Bushland BZ. Recorded from N Tanzania and S Kenya, with one morphologically somewhat aberrant specimen from Dakawa in C Tanzania. Unconfirmed records from Rwanda (Van der Straeten 1999).

Mastomys pernanus

Habitat The one specimen in Dakawa, Tanzania, was trapped in Brachystegia woodland (W.Verheyen, R.Verhagen & H. Leirs, unpubl.). Abundance The very low number of specimens in collections indicates that the species is rare or not easily captured with usual trapping techniques. Only a single specimen was trapped at Dakawa Ranch,Tanzania, although 1333 other small mammals were collected at the same locality over a period of two years (W. Verheyen, R.

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Mastomys shortridgei

Verhagen & H. Leirs, unpubl.). In Rwanda, however, 14 out of 201 rodent specimens in owl pellets were identified as M. pernanus (Misonne & Verschuren 1964). Remarks Apparently no other information available. Conservation

IUCN Category: Data Deficient.

Measurements Mastomys pernanus HB: 78 (73–88) mm, n = 7 T: 66 (60–78) mm, n = 6

HF: 16.1 (15–18) mm, n = 7 E: 14.9 (14–16), n = 7 WT: 18, 20 g, n = 2 GLS: 24.1 (23.2–25.1) mm, n = 3 GWS: 12.1 (11.8–12.5) mm, n = 3 M1–M3: 4.0 (3.8–4.4) mm, n = 6 S Kenya, N Tanzania (Van der Straeten 1999; all known specimens) Weight: E. Van der Straeten and H. Leirs unpubl. Key Reference

Van der Straeten 1999. Herwig Leirs

Mastomys shortridgei SHORTRIDGE’S MULTIMAMMATE MOUSE (SHORTRIDGE’S MASTOMYS) Fr. Souris à mammelles multiples de Shortridge; Ger. Shortridges Vielzitzenmaus Mastomys shortridgei (St Leger, 1933). Proc. Zool. Soc. Lond. 1933: 411. Okavango–Omatako junction, Grootfontein District, Namibia.

Taxonomy Originally described in the genus Myomys. Van der Straeten & Robbins (1997) confirmed the classification of this species in the genus Mastomys. It has been suggested that M. shortridgei was conspecific with M. angolensis, but the nipple arrangement is different in the two species, and angolensis is now placed in the genus Myomyscus. Referred to as Myomys shortridgei by Shortridge (1934) and as Praomys shortridgei by Smithers (1971). Synonyms: legerae. Subspecies: none. Chromosome number: 2n = 36, FN = 50. Description Medium-sized dark grey mouse. Dorsal pelage dark grey, sometimes nearly black; hairs grey at base with buffy tip. Ventral pelage grey; hairs grey at base with white tip. Upper surface of fore- and hindfeet white.Tail long (ca. 88% of HB), dark above and below. Skull typical of Mastomys, but pterygoid fossa wider than in other Mastomys and the anterior palatal foramina do not reach the inner root of M1 (Meester et al. 1986). There is confusion in the literature about the number of nipples: Gordon (1985) mentioned two rows of eight nipples, but Van der Straeten (2001 in litt.) confirmed that the holotype has two rows of five nipples each (total ten). Geographic Variation None recorded. Mastomys shortridgei

Similar Species M. natalensis. Smaller mean HB; tail longer as % of HB; ventral pelage dark grey; widespread distribution and common; nipples 12 × 2 = 24; 2n = 32. Distribution Endemic to Africa. ZambezianWoodland BZ. Usually thought to be endemic to extreme NW Botswana and NE Namibia in the region of the confluence of Okavango and Kwito rivers. However, Crawford-Cabral (1998) reported a few specimens from scattered localities in E Angola. Habitat Marshes and the banks and terraces of rivers. Sometimes commensal. Abundance Uncertain. Smithers (1971) recorded that it occurs with M. natalensis in Botswana but was the least common of the two

species. Rare in collections from Angola (Crawford-Cabral 1998), although Shortridge (1934) mentioned that it was plentiful locally. Remarks Terrestrial and nocturnal. Granivorous; probably omnivorous (Smithers 1971). None of 18 "" collected in Apr and May in NW Botswana showed signs of breeding, but a tiny juvenile was trapped in Feb (Smithers 1971). Conservation

IUCN Category: Least Concern.

Measurements Mastomys shortridgei HB: 120 (103–137) mm, n = 25 T: 105 (86–118) mm, n = 25 HF: 25 (23–27) mm, n = 25 471

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E: 18 (17–20) mm, n = 25 WT: 46 (35–74) g, n = 25 GLS: n. d. GWS: n. d. M1–M3: n. d.

Okavango Region and Western Caprivi, NE Namibia (Shortridge 1934) Weight: Smithers 1971 Key References

Shortridge 1934; Skinner and Smithers 1990. Herwig Leirs

GENUS Muriculus Ethiopian Striped Mouse Muriculus Thomas, 1903. Proc. Zool. Soc. Lond. 1902 (2): 314. (publ. 1903). Type species: Mus imberbis Rüppell, 1842.

Muriculus imberbis.

A monotypic genus confined to higher altitudes in Ethiopia. The genus has been allied to Mus and Zelotomys (Osgood 1936) and especially to Mus (Ellerman 1941).The genus is characterized by pro-odont incisors (but without notch on the posterior side as in Mus), the cutting edges of the incisors form a semi-circular shape (similar to those of Ammodillus), the rostrum is thin and narrow (especially when viewed laterally), and the coronoid process on the mandible is low without a point and is approximately the same height as the articular condyle (Figure 75). There is a dark middle-dorsal stripe. The single species is Muriculus imberbis. Figure 75. Skull and mandible of Muriculus imberbis (BMNH 28.1.1.153).

D. W. Yalden

Muriculus imberbis ETHIOPIAN STRIPED MOUSE Fr. Souris à crinière; Ger. Simien-Maus Muriculus imberbis (Rüppell, 1842). Mus. Senckenberg. 3: 110, pl. 6 (Fig. 4). Enschetgab, Ethiopia. 2800 m.

Taxonomy Originally placed in the genus Mus. Synonyms: chilaloensis. Subspecies: two. Chromosome number: not known. Description Distinctive very small murine, with pro-odont upper incisors that distinguish the skull from related species. Pelage soft and dense. Dorsal pelage brown to olive-grey, with slight speckling; hairs dark grey at base, brownish at tip; some all-black hairs especially along mid-dorsal line. Faint mid-dorsal stripe along the middle of the back, which does not extend as far forward as the head nor over the rump. (Pelage colour very similar to Mus mahomet except for mid-dorsal

stripe.) Flanks paler, less black, often with yellowish line separating dorsal and ventral colours. Ventral pelage greyish to buff-white, with tinge of orange-buff in some individuals. Ears grey, with sparse short buffy hairs. Fore- and hindfeet whitish-grey dorsally. Tail (ca. 70% of HB) very clearly bicoloured with small hairs; dark above, pale below. Skull: see genus profile. Nipples: not known. Geographic Variation M. i. imberbis: Ethiopian plateau west of the Rift Valley. Ventral pelage greyish-white or buff.

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Abundance Rare. Only 12 specimens known (Yalden et al. 1976; Yalden & Largen 1992). Collecting in Ethiopia (1968–98, n = ca. 6300 rodents) yielded no individuals of this species. Some individuals were found in houses in the Simien Mts in the 1970s (Müller 1977), but none were recorded there in the 1990s (Nievergelt et al. 1998). Despite extensive recent trapping, not known to occur in the Bale Mts. Remarks Probably terrestrial and nocturnal.The pro-odont upper incisors suggest that these mice may dig their own burrows in the otherwise rather open habitat. Presumably granivorous. One juvenile found in Sep (label, BMNH). Conservation IUCN Category: Endangered. The grassland habitat is threatened by continuing modification and destruction by humans and their livestock; Yalden et al. (1976) speculated that the species was less common than 50 years previously. Schlitter (1989) recommended that the species should be classified as Vulnerable.

Muriculus imberbis

M. i. chilaloensis: south-eastern plateau of Ethiopia, east of the Rift Valley. Ventral pelage white with a yellow tinge. Similar Species Mus mahomet. Very similar, but on average slightly smaller, lacks the mid-dorsal stripe; incisors orthodont (not pro-odont). Distribution Endemic to Africa. Afromontane–Afroalpine BZ of Ethiopia. Confined to the grasslands (woina dega) of the high plateaux of Ethiopia, from 1900 to 3400 m. Known from only about nine localities (Yalden et al. 1976,Yalden & Largen 1992).

Measurements Muriculus imberbis HB: 70, 78 mm, n = 2 T: 50, 52 mm, n = 2 HF: 16.5 (16–17) mm, n = 3 E: 11, 12 mm, n = 2 WT: n. d. GLS: 21.5 (20.6–22.5) mm, n = 3 GWS: 11.0 mm, n = 1 M1–M3: 4.0 (3.8–4.1) mm, n = 3 Ethiopia (D.W.Yalden unpubl.) Key References Müller 1977; Nievergelt et al. 1998;Yalden et al. 1976;Yalden & Largen 1992. D. W. Yalden

Habitat Typically open upland grasslands, but also reported as commensal in Simien Mts (Müller 1977).

GENUS Mus Old World Mice and Pygmy Mice Mus Linnaeus, 1758. Syst. Nat., 10th edn., 1: 59. Type species: Mus musculus Linnaeus, 1758.

Mus minutoides.

In Africa, the genus Mus is represented by about 20 species (Table 37). Species within the genus occur throughout most of the continent and are recorded from forest, savanna, highland grassland and semi-

arid habitats, and from sea level to ca. at least 2000 m. The only habitats where Mus does not occur are rainforests and arid deserts. Extralimitally, the genus is represented throughout the Old World. One species, Mus musculus, has been introduced into all continents (including Africa) and many islands, and has, in general, proved to be a very successful colonizer. The genus, as exemplified by African species, is characterized by small size (HB: ca. 43–90 mm, GLS: 16–25 mm, WT: 3–20 g), delicate build, shortish pelage and shortish tail without a terminal pencil. Upper incisors opisthodont or slightly pro-odont; uniquely there is a notch on the posterior face of the upper incisors. Other diagnostic skull characters include pronounced masseteric knob on the zygomatic plate near the lower edge, anterior palatal foramina, 473

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Table 37. Species in the genus Mus. Arranged in order of increasing mean head and body length. (n. d. = no data.)

a b

Species

Subgenusa

HB mean (range) (mm)

M. tenellus

N

48.4 (43–53)

M. haussa

N

49.7 (44–52)

M. mattheyi

N

52 (46–60)

M. indutus

N

53.2 (45–65)

M. minutoides

N

54.8 (45–68)

M. sorella

N

59.9 (51–73)

M. orangiae

N

59.6 (52–63)

M. musculoides

N

60.8 (55–70)

M. goundae

N

61 (60–62)

M. setzeri

N

61 (ca.)

M. oubanguii

N

62.7 (50–75)

M. baoulei

N

65.9 (59–73)

M. mahomet

N

67 (63–73)

M. bufo

N

68.4 (63–78)

M. spretus

M

77 (ca.)

M. triton

N

75.9 (69–80)

M. setulosus

N

81.8 (77–86)

M. musculus

M

83.7 (78–91)

M. neavei

N

88.8 (58–106)

M. callewaerti

N

88.8 (84–97)

Tail mean (range) (mm) [% of HB] 34 (30–36) [70%] 38 (35–42) [75%] 38.4 (33–44) [70%] 42 (30–52) [80%] 41 (38–49) [75%] 38.9 (34–46) [65%] 37.5 (36–39) [63%] 43.6 (32–55) [72%] 34 (31–37) [55%] 36 (31–48) [59%] 38 (26–44) [61%] 36.9 (32–45) [50–65%] 53.5 (46–60) [74%] 66.6 (61–74) [97%] 62 (55–71) [ca. 80%] 54 (49–63) [71%] 55.5 (52–59) [68%] 77.2 (66–85) [92%] 38.4 (33–48) [43%] 44.8 (43–46) [50%]

GLS mean (range) (mm)

White spots on head

Choanaeb

16.8 (16–17.5)

Subauricular and postauricular

U

16.6 (15–17)

None

U or V

16.6 (15–17)

None

U or V

17.7 (16.5–19.5)

Subauricular

U

18.8 (17.5–20.4)

Subauricular, small

U

19.7 (17.8–21.0)

Subauricular, small

V

18.6 (18.5–18.7)

n. d.

U

19.3 (18.3–20.2)

Subauricular, small

U

19 (18–20)

Postauricular, large

V

18.0 (17.5–18.3)

None

U

20 (18.6–21.5)

Postauricular, large

V

18.8 (17.7–20.4)

Suborbital, and subauricular (ochre)

U

19.9 (18.4–21.0)

None

U

20.9 (20.1–21.6)

None

U

19.8 (18.3–21.8)

None

U

21.8 (20.5–22.7)

None

U

21.1 (19.3–21.9)

None

U

21.6 (21–22.3)

None

U

18.5 (18–18.9)

?

V

25.4 (23.4–25.1)

None

?U

N = subgenus Nannomys; M = subgenus Mus. See genus profile for details. U = U-shaped choanae; V = V-shaped choanae.

Figure 76. Skull and mandible of Mus setulosus (MNHN 1998/887).

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Colour of ventral pelage

Chromosome number

Distribution and Notes

Pure white

n. d.

Sudan, S Ethiopia, S Somalia, S Kenya and C Tanzania; postauricular patch is obvious tuft

Pure white

2n = 28–34, FN = 38

Sudan and Sahel Savanna BZs; Senegal to Sudan

Pure white

2n = 36, FN = 36

Sudan and Guinea Savanna BZs; West Africa

Pure white

2n = 36, FN = 36–37

Southern Africa

Pure white

2n = 18–34, FN = 36

Southern and eastern Africa; widespread

Pure white

n. d.

Central Africa

Pure white

n. d.

C South Africa

Pure white

2n = 38 (polymorphic)

Guinea and Sudan Savanna BZs. W Africa (and towards central Africa)

Pure white

2n = 16–19, FN = 30

Central African Republic. Very rare

Pure white

n. d.

Namibia, Botswana, Zambia

Pure white

2n = 28, FN = 30–34

Central African Republic

Pure white

2n = 20, FN = 32

Guinea to Togo

Greyish-white (sometimes with orange)

2n = 36, aFN = 34

Ethiopia, SW Kenya, SW Uganda

Buff to greyish-buff

2n = 36, FN = 36

Mountains of Albertine Rift in E DR Congo, W Uganda and Burundi

Whitish

n. d.

N Morocco to N Libya

Medium-dark grey, white tips

2n = 34, FN = 34 (variable)

Central Africa; widespread

Pure white

2n = 36, FN = 36

Sierra Leone to S Sudan and Ethiopia

Usually grey, sometimes white

2n = 40

Mostly ports on coastline

Pure white

n. d.

Tanzania to South Africa; ears large and pale

Medium-dark grey, white tips

n. d.

NE and C Angola, S and W DR Congo

which extend well posterior to the anterior end of M1, laminae of the molar teeth, which tilt posteriorly, t1 of M1 distorted posteriorly and hence almost in line with t4 and t5, and very small M3 (Figure 76). There is great variation in the distribution and abundance of species of Mus in Africa. Some are widespread and numerous (M. minutoides, M. musculoides, M. setulosus), some have very limited distributions (e.g. M. bufo, M. goundae, M. oubanguii), and others are known from only few widely scattered localities and their full geographic range is uncertain (e.g. M. callewaerti, M. sorella, M. tenellus).Typically, individuals of many species of Mus are numerous in favoured environments and contribute a relatively large percentage to the total rodent community; they have short life expectancies and high fecundity, and the population

turnover is rapid. They form an important source of food for smaller predators, and their remains often contribute a high percentage of total prey in the pellets of owls. Individuals of Mus spp. are mostly gregarious and non-territorial. They are granivorous or omnivorous, their small size necessitating that they eat only high quality foods. The genus is divided into four subgenera, two of which are represented in Africa: subgenus Mus (typical Old World Mice) distinguished by larger size, flat anterior face to the zygomatic plate, masseteric knob at the lower anterior corner of the plate, and M3 with two (often inconspicuous) laminae (2 spp. – M. musculus, M. spretus); and the subgenus Nannomys (African Pygmy Mice), distinguished by smaller size, convex anterior face to the zygomatic plate, masseteric 475

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Family MURIDAE

knob at the lower centre of the plate, and M3 usually without laminae (18 spp.). In the past, Nannomys was referred to as Leggada (e.g. De Graaff 1981, Smithers 1983, Meester et al. 1986) and has been raised to the rank of genus by some authorities (e.g. Allen 1939, Roberts 1951, Bonhomme 1992). The distinction between the two subgenera is not always clearcut (Petter 1963b) although, in West Africa, for example, the two can be easily separated (Rosevear 1969). For Africa, the recognition of Nannomys as a subgenus appears to be warranted (Musser & Carleton 2005). Here, the African species are allocated to the genus Mus (rather than Leggada or Nannomys) following Petter & Matthey (1975) and Musser & Carleton (2005). Because all species of Mus (especially Nannomys) are small and morphologically similar, distinctions between species are often blurred. Geographical variation within widespread species is common and has resulted in the naming of many forms (species or subspecies), many of which are now regarded as synonyms. Within Nannomys, some species have been classified into groups implying at least some degree of relationship between the group-members. Based on some external and craniodental morphological similarities, Petter (1981) defined a ‘sorella group’, which included M. sorella, M. goundae, M. neavei, M. oubanguii, M. kasaicus (considered here to be conspecific with M. minutoides [but see Musser & Carleton 2005]) and two other species (acholi, wamae) now considered to be conspecific with M. sorella. Musser & Carleton (1993, 2005) indicate that the diagnostic characters of M. baoulei are closely similar to those of the sorella group, but suggest that the relationship of M. baoulei to other members of the sorella group requires fresh assessment. Musser & Carleton (2005) do not mention either M. minutoides or M. musculoides as belonging to the sorella group. Based on morphology of the sex chromosomes, two

groups within Nannomys can be distinguished. The first group (with acrocentric X and Y chromosomes) includes M. bufo, M. indutus, M. mattheyi, M. mahomet and M. tenellus, which all have 36 acrocentric chromosomes (diploid number 2n = 36, and fundamental number FN = 36). Two additional species belong to this group: M. setulosus (2n = 36, FN = 36), which is distinguished from the others by large heterochromatin additions on several pairs of autosomes, and M. haussa, which is distinguished by a pericentric inversion and varying numbers of autosomal centric fusions (2n = 28–34, FN = 38) (Veyrunes et al. 2004).The second group (with metacentric X and/or Y chromosomes formed by sex-autosome translocations) includes M. triton, M. oubanguii, M. musculoides, M. minutoides and M. goundae (Matthey 1966a, b, Jotterand 1972, Veyrunes et al. 2004). This group is of special interest because of the large variation in the number and morphology of chromosomes between populations and even within populations (2n = 18–34, FN = 30–36). This is particularly evident for M. minutoides and probably also for M. triton. This diversity most likely indicates the occurrence of cryptic species (Veyrunes et al. 2004). In the absence of chromosomal data, the other species have not yet been assigned to either group (i.e. M. baoulei, M. callewaerti, M. neavei, M. orangiae, M. setzeri and M. sorella). Consequently, it is not yet possible to determine if the sorella group of Petter (1981) corresponds to the second of the groups based on chromosome morphology (although there is at least some overlap in species composition); pending resolution of this problem, the affinities of species to these groups are not given in the species profiles. Species in the genus may be distinguished by size, pelage colour, skull characters and karyology. D. C. D. Happold & F. Veyrunes

Mus baoulei BAOULE PYGMY MOUSE Fr. Souris naine de Baoulé; Ger. Baouli-Zwergmaus Mus baoulei (Vermeiren & Verheyen, 1980). Rev. Zool. Afr. 94: 573. Lamto, Côte d’Ivoire.

Taxonomy Originally described in the genus Leggada. Subgenus Nannomys. Synonyms: none. Chromosome number: 2n = 20, FN = 32 (M. Tranier unpubl.). Description Very small mouse. Dorsal pelage grey-brown to dark-brown; hairs grey at base, brown at tip. Flanks paler than dorsal pelage, becoming yellowish- to reddish-brown towards ventral surface. Ventral pelage pure white, clearly delineated from dorsal pelage on flanks and cheeks. One ochre spot below each eye and one below each ear. Tail short (50–65% of HB), brown above, paler below, covered with fine and short hairs. Skull: anterior palatal foramina very long (3.65 mm, range 3.65–4.70), and longer than for sympatric species of Mus. Nipples: not known.

M. kasaicus, M. neavei and M. sorella. All belong to the ‘sorella’ group of Petter (1981), are of similar size, and have a more easterly distribution. M. setulosus. A little larger; chromosome number: 2n = 36, FN = 36; no auricular spots; relatively longer tail; West to East Africa. M. musculoides. Brighter pelage; chromosome number: 2n = 18–34, FN = 36; relatively longer tail; West and East Africa. M. haussa. Paler colour; chromosome number: 2n = 28–34, FN = 38; West Africa. M. mattheyi. Chromosome number: 2n = 36, FN = 36; West Africa. Cranial and dental characters also enable discrimination between these species (Vermeiren & Verheyen 1980, Musser & Carleton 1993).

Geographic Variation None recorded. Similar Species M. goundae. Chromosome number: 2n = 16–19, FN = 30; Central African Republic. M. oubanguii. Chromosome number: 2n = 28, FN = 30–34; Central African Republic.

Distribution Endemic to Africa. Guinea Savanna BZ and Northern Rainforest–Savanna Mosaic. Known only from E Guinea, Côte d’Ivoire and Togo (Vermeiren & Verheyen 1980, Musser & Carleton 1993, Robbins & Van der Straeten 1996). Habitat Savanna.

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Mus bufo

Abundance Uncertain; may be locally abundant (e. g. Lamto, Côte d’Ivoire, cf M. musculoides, L. setulosus) (Vermeiren & Verheyen 1980). Remarks Apparently no other information available. Conservation

IUCN Category: Least Concern.

Measurements Mus baoulei HB: 65.9 (59–73) mm, n = 47 T: 36.9 (32–45) mm, n = 35 HF (c.u.): 13.3 (12–15) mm, n = 40 E: 10 (8–12) mm, n = 35 WT: n. d. GLS: 18.8 (17.7–20.4) mm, n = 6* GWS: 9.1 (8.3–9.8) mm, n = 40 M1–M3: 3.1 (2.8–3.4) mm, n = 66 Côte d’Ivoire (Vermeiren & Verheyen 1980, MNHN) and Guinea (MNHN) *MNHN only Key Reference

Vermeiren & Verheyen 1980.

Mus baoulei

Laurent Granjon

Mus bufo TOAD PYGMY MOUSE (RWENZORI MOUSE) Fr. Souris naine crapaud; Ger. Kröten-Zwergmaus Mus bufo Thomas, 1906. Ann. Mag. Nat. Hist., ser. 7, 18: 145. Rwenzori East, Uganda, 6000 ft (1830 m).

Taxonomy Subgenus Nannomys. Morphologically similar to sympatric Mus triton but different in external, skull, dental, karyological and ecological characters (F. Dieterlen unpubl.). The form ablutus from Idjwi I. in L. Kivu could be a valid subspecies (Allen & Loveridge 1942). Electrophoretic analysis of protein enzymes at 24 loci indicates that M. bufo is more closely related to M. gratus (= M. minutoides) than to M. triton (Van Rompaey et al. 1984). Synonyms: ablutus, wambutti. Subspecies: none currently recognized. Chromosome number: 2n = 36, FN = 36 (Jotterand-Bellomo 1988, Maddalena et al. 1989). Description Small mouse, smaller than sympatric M. triton. Dorsal pelage stiff and thick, hairs long (8–9 mm). Dorsal pelage dark coppery-brown, rather variable; hairs slate-grey to black at base, dull reddish-brown at tip; softer underfur black, grey or buff. Rump darkish. Ventral pelage buff to greyish-buff, shorter than dorsal pelage. Yellowish line between dorsal and ventral pelage in some individuals. Ears dark with scattered short buffy hairs. No postauricular or subauricular spots. Upper surface of forefeet and hindfeet brown with a tinge of buff. Tail long (ca. 97% of HB), finely scaled, with dense short blackish bristles above, whitish below. Skull stoutly built but less so than in M. triton; anterior palatal foramina long, extending nearly to the middle of M1; mesopterygoid fossa broad compared to M. triton (Thomas 1906a). Nipples: 3 + 1 = 8.

Mus bufo

Geographic Variation None currently recognized (seeTaxonomy). 477

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Family MURIDAE

Similar Species M. triton. Larger; tail shorter (71% of HB); ventral pelage paler (usually greyish-white); different chromosome number (see species profile).

normally the second or third commonest species of the three species of Mus, and much less abundant than M. minutoides (Rahm 1967). More common in selected habitats in Kahuzi-Biega N. P., E DR Congo.

Distribution Endemic to Africa. Afromontane–Afroalpine BZ in montane regions of the Albertine Rift Valley, normally 1500–3000 m (Misonne 1963, Delany 1975, F. Dieterlen unpubl.). Recorded in Upper Ituri region west of L. Albert and on both sides of the Albertine Rift Valley from ca. 01° N to ca. 03° S. Specific locations include Rwenzori Mts (DR Congo, Uganda); Virunga Mts and mountains west of L. Edward (DR Congo); and on both sides of L. Kivu (E DR Congo, W Rwanda, NW Burundi) (Elbl et al. 1966, Dieterlen 1967a, Rahm 1967, Maddalena et al. 1989, J. Kerbis Peterhans, in litt.). One record from Itombwe Mts, west of L. Tanganyika (von Wettstein-Westersheim 1923). Records from lowland forests below 1000 m need verification, especially Ituri Forest (the type locality of M. b. wambutti; Lönnberg & Gyldenstolpe 1925) and Irangi, E DR Congo (Rahm 1967). An extralimital record from the Aberdare Mts, Kenya (Misonne 1963) also needs verification (not shown on map).

Adaptations et al. 1998).

Habitat In Kahuzi-Biega N. P., E DR Congo (at 2300–2400 m), preferred habitats are bamboo forest (Arundinaria alpina) and stands of Hagenia abyssinica, both with grass cover (Panicum massaiense) (F. Dieterlen unpubl.). Other habitats include secondary growth on the edge of dense montane forest (2000–2200 m), low and dense afroalpine vegetation (e.g. with Erica, Senecio, Lobelia, Helichrysum) on the peak of Mt Kahuzi, E DR Congo (3260–3308 m), and dense stands of Pennisetum purpureum in the cultivated zone west of L. Kivu, DR Congo (1500–1900 m). On Rwenzori Mts, Uganda, recorded in grassland (1500 m), low montane forest (1900 m), high montane forest (2600 m) and bamboo (2960 m) (Kerbis Peterhans et al. 1998).

Predators, Parasites and Diseases No information.

Abundance Generally rare in the cultivated zone west of L. Kivu (1500–1900 m). Less common than M. minutoides on Idjwi I. in L. Kivu (Rahm & Christiaensen 1966). Often occurs sympatrically with M. triton and M. minutoides in E DR Congo; e.g. at five locations, M. bufo comprised 4.38% of all small mammals (M. minutoides 8.24%, M. triton 3.91%, n = 2340) although the percentage values varied according to locality from 0.7% to 6.5% (Rahm 1967). Numbers varied from month to month over four years (at a single locality); M. bufo was

Mainly nocturnal (Delany 1975, Kerbis Peterhans

Foraging and Food Herbivorous, occasionally omnivorous. Most stomach contents (eight of 13) contained exclusively starchlike vegetable material, a whitish-brownish pulp of seeds and tubers; five stomachs contained animal food (chitinous parts, fibres, parts of intestines) comprising only up to ca. 20% of stomach contents (F. Dieterlen, unpubl.). Reproduction and Population Structure Seasonal trends in reproduction not known. A few pregnant and/or lactating "" recorded in different seasons. Embryo number: 3 and 4 (n = 2). As for other species of the genus Mus, captures of !! outnumber "" (36 : 25; F. Dieterlen unpubl.).

Conservation

IUCN Category: Least Concern.

Measurements Mus bufo HB: 68.4 (63–78) mm, n = 30 T: 66.6 (61–74) mm, n = 27 HF: 15.3 (13–18) mm, n = 30 E: 11.8 (10–13) mm, n = 30 WT: 10.4 (6–16) g, n = 30 GLS: 21.0 (20.1–21.6) mm, n = 11 GWS: 10.1 (9.9–10.3) mm, n = 9 M1–M3: 3.4 (3.1–3.5) mm, n = 10 Parc National de Kahuzi-Biega, DR Congo (F. Dieterlen unpubl., SMNS) Key References

Delany 1975; Misonne 1963. Fritz Dieterlen

Mus callewaerti CALLEWAERT’S PYGMY MOUSE Fr. Souris naine de Callewaert; Ger. Callewaerts Zwergmaus Mus callewaerti (Thomas, 1925). Ann. Mag. Nat. Hist., ser. 9, 15: 668. Lualaba, Luluabourg, DR Congo. 610 m.

Taxonomy Subgenus Nannomys. Described in the genus Hylenomys (Thomas 1925), a genus distinguished from Leggada (= Nannomys) by the presence of pro-odont incisors (a character now known to be present in several species of Leggada). Hylenomys now synonymized with Mus (Musser & Carleton 1993, 2005). Synonyms: none. Chromosome number: not known. Description The largest species of the genus, similar externally to M. triton. Pelage stiff, dense and coarse. Dorsal pelage mediumbrown, hairs dark grey at base, with subterminal medium brown

band and black tip. Ventral pelage whitish-grey; hairs dark grey on basal half, off-white on terminal half. (Hill & Carter [1941] state that ventral pelage may have a pale pinkish-brown tinge especially on chest.) Ears short, brown. Lips and chin white. No postauricular or subauricular spots. Fore- and hindfeet dirty-white. Tail short (ca. 50% of HB), indistinctly bicoloured, brown above, whitish below. Skull: large for the genus; supraorbital crests well developed; auditory bullae large; incisor teeth white on anterior surface, slightly pro-odont. Nipples: not known.

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Mus goundae

Geographic Variation None recorded. Similar Species M. triton. Smaller; ventral pelage grey; supraorbital crests on skull absent; auditory bullae small; incisor teeth yellowish-orange on anterior face; sympatric at some localities (e.g. Chitau, Angola). M. sorella. Much smaller; ventral pelage pure white. M. minutoides. Much smaller; dorsal pelage brownish-buff; ventral pelage pure white; incisor teeth yellowish on anterior surface. Distribution Endemic to Africa. Zambezian Woodland BZ and Southern Rainforest–Savanna Mosaic. Recorded from C and NE Angola and S and SE DR Congo. Altitudes from 610 m (holotype) to 1810 m (SE DR Congo) (Crawford-Cabral 1998). Sympatric with M. triton. Habitat ‘Forest’ (holotype in SE DR Congo; Cabrera & Ruxton 1926) and savanna (Sanborn 1952). Abundance Uncertain; known by only a few specimens. Recorded as ‘peu commun’ (Sanborn 1952). Remarks One specimen had eaten fruits of prickly pear Opuntia sp. In Angola, Hill & Carter (1941) remarked ‘Apparently in the debris they live on the many insects frequenting such places.’ Females collected in Jul and Aug were ‘apparently lactating’ (Hill & Carter 1941). Conservation IUCN Category: Least Concern. The limited number of records and rarity of specimens are causes for concern. In Angola, there are many seemingly suitable localities where the species has not been encountered (Crawford-Cabral 1998). Eaten by local people (Sanborn 1952).

Mus callewaerti

T: 44.8 (43–46) mm, n = 4 HF: 15.3 (12–17) mm, n = 4 E: 11.3 (10–14) mm, n = 4 WT: n. d. GLS: 24.5 (23.4–25.1) mm, n = 4 GWS: 11.3 (10.7–11.9) mm, n = 3 M1–M3: 3.8, 3.8 mm, n = 2 Auditory bulla: 4.9, 5.1 mm, n = 2 Angola and DR Congo (Hill & Carter 1941, Misonne 1965b) Key Reference 1952.

Measurements Mus callewaerti HB: 88.8 (84–97) mm, n = 4

Hill & Carter 1941; Misonne 1965b; Sanborn D. C. D. Happold

Mus goundae GOUNDA RIVER PYGMY MOUSE Fr. Souris naine de la Gounda; Ger. Gounda-Fluss Zwergmaus Mus goundae Petter and Genest, 1970. Mammalia 34: 455. Gounda River, N Central African Republic.

Taxonomy Subgenus Nannomys. Related to M. oubanguii. Synonyms: none. Chromosome number: 2n = 16–19, FN = 30; chromosomal polymorphism caused by Robertsonian translocations (Jotterand 1970, 1972).

with prelobe and without accessory cusp; M3 small; M1 with anterior lobe quadricuspidate. Nipples: 2 + 2 = 8.

Description Very small brownish-orange mouse, with pure white ventral pelage. Dorsal pelage ochraceous-brown, with orangerufous on flanks. Ventral pelage pure white. Colour of dorsal pelage and ventral pelage clearly delineated on lower flanks. Ears large, blackish, slightly pointed at tip. White postauricular patch present. Fore- and hindfeet comparatively large, white. Tail short (ca. 55% of HB). Skull: rostrum elongated; incisors slightly pro-odont, choanae V-shaped; anterior palatal foramina short (one tenth of M1–M3), M1

Similar Species M. oubanguii. Similar size; dorsal pelage reddish-brown; large white postauricular patch; nipples 3 + 2 = 10; chromosome number: 2n = 28. M. setulosus. Larger (HB: 81.8 [77–86] mm,T: 55.5 [52–59] mm, GLS: 21.1 [19.3–21.9]); dorsal pelage blackish-brown; no auricular patches; nipples not known; chromosome number: 2n = 36.

Geographic Variation

None recorded.

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Family MURIDAE

M. musculoides. Similar size; dorsal pelage golden-brown; no postauricular patch; nipples 2 + 2 = 8; chromosome number: 2n = 25–32 (polymorphic); common and widespread. Distribution Endemic to Africa. Guinea Savanna BZ. Only known from near the Gounda R., north-east of N’Délé, N Central African Republic. Habitat

Wooded savanna.

Abundance No information; known only from the type locality. Remarks

Apparently no other information available.

Conservation IUCN Category: Data Deficient. The single known population is very small, and the number of individuals is probably declining. Measurements Mus goundae HB: 60, 62 mm, n = 2 T: 31, 37 mm, n = 2 HF: 12, 14 mm, n = 2 E: 12, 13 mm, n = 2 WT: n. d. GLS: 18, 20 mm, n = 2 GWS: n. d.

Mus goundae

M1–M3: 3.5, 3.7 mm, n = 2 Central African Republic (MNHN) Key Reference

Petter & Genest 1970. F. Petter

Mus haussa HAUSA PYGMY MOUSE Fr. Souris naine Haussa; Ger. Haussa Zwergmaus Mus haussa (Thomas and Hinton, 1920). Novit. Zool. 27: 319. Farniso, near Kano, N Nigeria.

Taxonomy Originally described in the genus Leggada. Subgenus Nannomys. Morphologically and ecologically similar to M. tenellus (Rosevear 1969, Musser & Carleton 2005). Synonyms: none. Chromosome number: 2n = 28–34, FN =38 (Jotterand 1972, Veyrunes 2002, Granjon & Dobigny 2003). Description Very small pale-coloured mouse; the smallest species of Mus in sub-Saharan Africa. Dorsal pelage pale sandy. Ventral pelage pure white. Colour of dorsal pelage and ventral pelage clearly delineated on flanks. Ears sandy-grey, with small whitish or sandy hairs. No postauricular patch of white hairs. Cheeks, lips and throat white. Fore- and hindfeet white. Tail short (ca. 75% of HB), scaly, pale or white, more or less naked. Skull: GLS 17 mm or less (cf. M. minutoides); upper incisors opisthodont, choanae U-V shaped, anterior palatal foramina elongated; M1 elongated and 65–70% of M1–M3; M1 with anterior lobe tricuspidate. Nipples: 3 + 2 = 10. Geographic Variation None recorded.

pelage ochre-tawny to chestnut; parapatric. M. musculoides. Larger (HB: 61 [55–70] mm, GLS: 18.4 [17.8– 18.9] mm); dorsal pelage darker, golden-brown flecked with dark brown; distribution in less arid habitats. Distribution Endemic to Africa. Sahel and Sudan Savanna BZs from Senegal to Chad and Sudan. Habitat

Semi-arid grassland savanna of the sub-Sahara region.

Abundance Evidence from remains in owl pellets (from Senegal, Nigeria and Chad) suggest this species may be locally common (see below). Remarks Nocturnal and terrestrial. Tolerates the hottest and driest climates of all species of pygmy mice. Diet is seeds and insects (no detailed studies). Remains found in owl pellets in W Senegal (F. Petter unpubl.), Mali (Meinig 2000) and N Nigeria where they formed 13% of the rodent prey (Demeter 1978).

Similar Species M. mattheyi. On average slightly larger (HB: 52 [46–60] mm); dorsal 480

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Mus indutus

Conservation

IUCN Category: Least Concern.

Measurements Mus haussa HB: 49.7 (44–52) mm, n = 7 T: 38 (35–42) mm, n = 7 HF: 12 (12–13) mm, n = 7 E: 8.8 (8–10) mm, n = 7 WT: 3 g, n = 1 GLS: 16.6 (15–17) mm, n = 6 GWS: 8.4 (8.2–8.6) mm, n = 4 M1–M3: 2.9 (2.9–3.1) mm, n = 7 Senegal (St Louis), Niger (Niamey), Nigeria (Farniso, Kano) and Chad (Ndjamena); BMNH, MNHN, SMF Key References

Happold 1987; Rosevear 1969. F. Petter

Mus haussa

Mus indutus DESERT PYGMY MOUSE Fr. Souris naine du désert; Ger. Wüstenzwergmaus Mus indutus (Thomas, 1910). Ann. Mag. Nat. Hist., ser. 8, 5: 89. Molopo River, South Africa.

Taxonomy Originally described as Leggada bella induta. Subgenus Nannomys. Formerly considered as a subspecies of M. minutoides. Currently, indutus refers to Mus from the western parts of southern Africa, but the limits of its distribution are not known. Synonyms: deserti, pretoriae, valschensis. Subspecies: two or three. Chromosome number: 2n = 36, FN = 36–37 (Matthey 1966a,Veyrunes et al. 2004). Description Very small mouse with soft pelage. Dorsal pelage variable shades of pale buff or pale buffy-orange; hairs slate-grey at base, some with black tip giving a grizzled appearance. Flanks buffyorange without black-tipped hairs. Ventral pelage (including chin) pure white. Clear delineation between colour of flanks and ventral pelage. Head with pointed nose and long vibrissae. Ears moderately sized and rounded, brownish; small white subauricular patch (usually absent in eastern part of distribution). Limbs short with whitish feet; four digits on forefeet; five digits on hindfeet. Digits 3 to 4 elongated on both forefeet and hindfeet. All digits with well-developed claws. Tail long (ca. 80% of HB), pale buff above, white below. Nipples: 2 + 2 = 8. Geographic Variation M. i. indutus: Botswana and Namibia. Dorsal pelage comparatively pale buff. M. i. pretoriae/valschensis: Gautung Province, South Africa. Dorsal pelage tawny ochraceous-buff, reminiscent of Mus minutoides.

M. setzeri. Ears longer; white ventral pelage extends onto upper rump and muzzle; sympatric.

Similar Species M. minutoides. Dorsal pelage generally darker, tail dark above; allopatric.

Distribution Endemic to Africa. Zambezian Woodland and South-West Arid (Kalahari Desert) BZs. Recorded from C and

Mus indutus

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Family MURIDAE

N Namibia (Matson & Blood 1994), through Botswana to NW South Africa and W Zimbabwe. Extends northwards into S Angola (Crawford-Cabral 1998) and SW Zambia (Ansell 1978). Habitat Tolerates a wide range of habitats in semi-arid savannas (Nel 1978); avoids open micro-habitats (Kerley et al. 1990). Abundance Abundant to very abundant. May undergo population explosions during periods of high food supply (Smithers 1971). In the Kalahari Desert of South Africa, M. indutus is the third most abundant species of small mammal after Rhabdomys pumilio and Gerbillurus paeba (Nel 1978). Adaptations Nocturnal and terrestrial. Excavates its own burrows in sandy soil, but also uses burrows excavated by other species; may hide under rocks, fallen logs or beneath the bark of trees. Spherical nests are constructed underground from grass or other soft fibres, in which young are born. Foraging and Food Diet is predominantly grass seeds and pods of Acacia trees, but includes insects (Smithers 1971, Nel 1978). Green plant material may also be important (Kerley et al. 1990). Social and Reproductive Behaviour Highly aggressive towards conspecifics, with cases of cannibalism reported in captivity (Skinner & Smithers 1990). Such aggressive behaviour suggests territoriality, but critical studies are lacking. Offspring remain with both parents until they are weaned.

Reproduction and Population Structure May breed throughout the year, but peaks in pregnancy are recorded in Botswana during the wet season (Oct–Apr; Smithers 1971). Embryo number: 4.9 (2–8, n = 17; Botswana; Smithers 1971). Population explosions in M. indutus occur at the same time as those of Mastomys natalensis. High numbers of M. natalensis may possibly provide predator cover for M. indutus. Predators, Parasites and Diseases No information. Conservation IUCN Category: Least Concern. Widespread and abundant throughout its range. Measurements Mus indutus HB: 53.2 (45–65) mm, n = 12* T: 42 (30–52) mm, n = 90 HF: 14 (13–16) mm, n = 90 E: 11 (8–12) mm, n = 90 WT: 5.4 (3–8) g, n = 85 GLS: 17.7 (16.5–19.5) mm, n = 11 GWS: 9.3 (8.5–10.0) mm, n = 10 M1–M3: 3.0 (2.8–3.1) mm, n = 11 Body measurements and weight: Botswana, unsexed individuals (Smithers 1971) Skull measurements: Botswana, South Africa (Roberts 1951) *Specimen labels (TM) Key References

Nel 1978; Smithers 1971. A. Monadjem

Mus mahomet MAHOMET PYGMY MOUSE Fr. Souris naine de Mahomet; Ger. Mohammed-Zwergmaus Mus mahomet Rhoads, 1896. Proc. Acad. Nat. Sci., Philadelphia 1896: 532. Sheik Mahomet, Ethiopia.

Taxonomy Subgenus Nannomys. Within Ethiopia, M. mahomet is a distinctive species. The name kerensis Heuglin 1877 might be a prior name (Yalden et al. 1976). The status of this taxon is uncertain, and it may belong to another species from further south in Africa, such as M. minutoides, M. bufo, M. sorella or bella (synonym of M. minutoides) (Yalden & Largen 1992). Synonyms: emesi. Subspecies: none. Chromosome number: 2n = 36, aFN = 34 (Aniskin et al. 1998). Description Very small mouse, similar to M. minutoides. Pelage dense, short and slightly coarse. Dorsal pelage dark greyish-brown, slightly speckled with buff; hairs grey at base, brown or buff at tip. Ventral pelage greyish-white, sometimes with pale orange tinge; hairs grey at base, off-white at tip. Ventral pelage clearly delineated from flanks usually by thin orange-buff or sandy-yellow line. Ears darkly pigmented, prominent, covered with sparse short buffy hairs. No postauricular or subauricular white patches. Fore- and hindfeet white. Tail short (ca. 74% of HB), scaly, with many very short hairs; dark above, slightly paler below. Nipples: 3 + 2 = 10.

Geographic Variation

None recorded.

Similar Species M. tenellus. Mostly smaller; dorsal pelage pale sandy-brown, ventral pelage pure white; white postauricular and/or subauricular patches; in Ethiopia, < ca. 2000 m). M. triton. On average larger, dorsal pelage olive-brown; ventral pelage grey or whitish-grey; in Ethiopia, SE forests only. M. musculoides. Similar size; sympatric in Uganda and Kenya. M. setulosus. Slightly larger; dorsal pelage grey, ventral pelage pure white; sympatric or parapatric in Ethiopia; in Ethiopia, 1000– 1500 m. Distribution Endemic to Africa. Afromontane–Afroalpine BZ of Ethiopia and Eritrea; perhaps confined to the plateaux at 1500–3200 m (Yalden & Largen 1992). Considered by Musser & Carleton (1993, 2005) to occur also in SW Kenya and SW Uganda (not mapped). Limits unknown.

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Mus mattheyi

Habitat Upland forest-edges, scrub and grasslands, habitats partly shared with Stenocephalemys albipes, but unlike S. albipes, M. mahomet is not a true forest species (Yalden 1988,Yalden & Largen 1992). Abundance Moderately common and widespread at middle altitudes in Ethiopia. Comprised 11% (n = 285) of rodents caught in Menagesha State Forest (Afework Bekele 1996a), and 6% (n = 905) in a wide range of sites in S Ethiopia (Rupp 1980). Remarks Little is known about the biology of this species. Nocturnal and presumed to feed on seeds and insects. Parous "", and !! with enlarged testes, were trapped in Aug in Bale, Ethiopia, and a lactating " was recorded in Jan at Illubabor, Ethiopia (1500 m; BMNH). Conservation

IUCN Category: Least Concern.

Measurements Mus mahomet HB: 67 (63–73) mm, n = 21 T: 53.5 (46–60) mm, n = 21 HF: 15 (14–15) mm, n = 21 E: 11.8 (9–13) mm, n = 21 WT: 10.3 (6–13) g, n = 17 GLS (CbL): 19.9 (18.4–21.0) mm, n = 21 GWS: n. d. M1–M3: 3.2 (3.0–3.6) mm, n = 21 Ethiopia (Rupp 1980,Yalden 1988)

Mus mahomet

Key References Largen 1992.

Afework Bekele 1996a; Rupp 1980; Yalden & D. W. Yalden

Mus mattheyi MATTHEY’S PYGMY MOUSE Fr. Souris naine de Matthey; Ger. Mattheys Zwergmaus Mus mattheyi Petter, 1969. Mammalia 33: 118. Accra, Ghana.

Taxonomy Subgenus Nannomys. Closely related to M. haussa. Musser & Carlton (1993, 2005) comment that they were ‘unable to assign anything to mattheyi [and] the morphological traits used to distinguish mattheyi from haussa by Petter (1969) and by Petter & Matthey (1975) vary in a continuous fashion from typical haussa morphology to that considered diagnostic for mattheyi’. However, Veyrunes (2002) showed that M. mattheyi must be considered as a valid species because of its very high molecular divergence and chromosomal differences compared with M. haussa. Synonyms: none. Chromosome number: 2n = 36, FN = 36. The karyotype is considered to be ancestral for African Mus (Jotterand-Bellomo 1986). Description Very small mouse, on average larger than M. haussa but on average smaller than M. musculoides. Dorsal pelage ochretawny to chestnut, usually darker along mid-dorsal line. Flanks fawn. Ventral pelage white. Colour of dorsal pelage and ventral pelage clearly delineated on flanks. Ears grey; no postauricular white patch. Fore- and hindfeet white. Tail short (ca. 70% of HB). Skull: upper incisors opisthodont; choanae V-shaped, sometimes tending to U-shaped (as in M. haussa); anterior palatal foramina elongated (but less than in M. haussa) reaching to level of posterior end of M1; M1 with prelobe; M1 with anterior lobe tricuspidate. Nipples 2 + 2 = 8.

Mus mattheyi

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Geographic Variation None recorded.

Remarks

Similar Species M. haussa. On average slightly smaller (HB: 49.7 [44–52] mm, HF: 12 [11.5–12.5] mm, GLS: 16.6 [15–17] mm); dorsal pelage pale sandy; distribution in more arid habitats. M. musculoides. On average larger (HB: 61 [55–70] mm, HF: 13.6 [12–14] mm, GLS: 18.4 [17.8–18.9] mm); dorsal pelage darker, golden-brown flecked with dark brown; sympatric.

Conservation

Apparently no other information available. IUCN Category: Least Concern.

Measurements Mus mattheyi HB: 52 (46–60) mm, n = 27 T: 38.4 (33–44) mm, n = 22 HF: 12.1 (11–13) mm, n = 27 E: 8.8 (7–10) mm, n = 27 Distribution Endemic to Africa. Sudan Savanna and Guinea WT: n. d. Savanna BZs. Recorded from Senegal to Côte d’Ivoire (and perhaps GLS: 16.6 (15–17) mm, n = 6 Togo), and Burkina (Veyrunes 2002), but distribution may be more GWS: 8.4 (8.2–8.6) mm, n = 4 extensive. Musser & Carleton (2005) record only from the type M1–M3: 2.9 (2.8–3.0) mm, n = 4 locality (see above). Geographic range is further south than for M. Throughout geographic range (MNHN) haussa. Key Reference Petter 1969. Habitat Moist woodland savanna and grass savanna. The holotype was taken on the Accra plain.

F. Petter

Mus minutoides TINY PYGMY MOUSE Fr. Souris naine d’Afrique australe; Ger. Kleine Zwergmaus Mus minutoides Smith, 1834. S. Afr. Quart. J., ser. 2, 2: 157. Cape Town, South Africa.

Taxonomy Subgenus Nannomys. Taxonomic status of this species is uncertain. Formerly, M. indutus and M. orangiae were considered subspecies of M. minutoides, but are now regarded as valid species (Musser & Carleton 1993, 2005). Currently, minutoides refers to Mus from the eastern side of Africa, from East Africa to South Africa, but the limits of the geographic range are uncertain. If M. musculoides is included in M. minutoides (as in Musser & Carleton 2005), then M. minutoides has a distribution throughout much of the savanna regions of Africa. The minutoides–musculoides complex and the relationship between M. minutoides and M musculoides is not yet resolved. Diploid chromosome number apparently variable, suggesting minutoides may include sibling species. Synonyms: kasaicus, marica, minimus, umbratus. Subspecies: three, validity uncertain. Chromosome numbers: 2n = 18, FN = 36 (Western Cape Province, South Africa); 2n = 34 (KwaZulu– Natal, South Africa) (Matthey 1966a, Skinner & Smithers 1990, Veyrunes et al. 2004). Description Very small mouse with soft, shiny pelage. Dorsal pelage variable shades of brownish-buff to brownish-orange; hairs slate-grey at base with black tip. Flanks buffy-orange. Ventral pelage (including chin) pure white. Clear delineation between colour of flanks and ventral pelage. Head with pointed nose and long vibrissae. Ears brownish, moderately sized and rounded. Limbs short. Feet white with well-developed digits; four digits on forefoot; five digits on hindfoot. Digits 2–4 elongated. All digits with well-developed claws. Tail short (ca. 75% of HB), brownish above, paler below. Nipples: 3 + 2 = 10. Geographic Variation Meester et al. (1986) list three subspecies for southern Africa:

M. m. minutoides: Western Cape, Eastern Cape and KwaZulu–Natal Provinces, South Africa. M. m. umbratus: Swaziland and NE South Africa. M. m. marica: E Mpumalanga Province, South Africa; S Mozambique. Specimens from Western and Eastern Cape Provinces (South Africa) are slightly darker than specimens from KwaZulu–Natal and E Mpumalanga Provinces (South Africa), Swaziland and S Mozambique. Differences, however, not very noticeable due to great variation within each region. Specimens from S Mozambique have fewer black-tipped hairs in dorsal pelage, resulting in brighter brownish-orange colour (but highly variable). Similar Species M. indutus. Dorsal pelage generally paler; tail pale above. M. setzeri. Ears longer; white ventral pelage extends on to upper rump and muzzle. Distribution Endemic to Africa. Zambezian Woodland BZ (eastern part), South-West Cape BZ and southern Somalia–Masai Bushland BZ. Also occurs at higher altitudes in Swaziland. Recorded in S and E South Africa, northwards through Swaziland and S Mozambique to Zimbabwe and Malawi.Two isolated records from NE Namibia (Matson & Blood 1994). Northern limit of the geographic range where it adjoins that of M. musculoides in eastern Africa is uncertain (shown by striped lines on map). Replaced by M. indutus in western part of Zambezian Woodland BZ. Habitat Tolerates a wide range of habitats. In Swaziland, occurs in afromontane and riparian forests, short to tall grasslands, rocky outcrops, all forms of Acacia and broad-leaved woodland, cultivated

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Mus minutoides

bark of trees. During the non-breeding season, shelters may be changed regularly. Construct spherical nests, under cover, from grass or other soft fibres in which young are born. In captivity, does not enter spontaneous torpor at low temperatures or during periods of reduced food availability (Webb & Skinner 1995, Downs & Perrin 1996). Foraging and Food Omnivorous. Diet consists mostly of grass seeds and insects (Rowe-Rowe 1986), but foliage predominates in the Karoo of SW South Africa (Kerley 1992). Proportional contribution of food types in stomach contents: 44% vegetable material, 40% seeds, 16% arthropods (n = 17, Swaziland; Monadjem 1997b).

Mus minutoides (southern Africa) and Mus musculoides (western Africa). See also text.

fields and recently burnt areas (Monadjem 1998a). May also occur in suburban gardens, young pine plantations and on the fringe of wetlands. Altitude range: near sea level to 2700 m (Rowe-Rowe & Meester 1982a). Abundance Abundant to very abundant. Densities of up to 28/ ha have been recorded (Monadjem 1998b), and presumably may reach much higher densities during favourable conditions. Adaptations Nocturnal and terrestrial.Tiny Pygmy Mice excavate their own burrows in soft soil, and will also use holes excavated by other species, or will rest under rocks, fallen logs or beneath the

Social and Reproductive Behaviour Reported to forage independently, but frequently pairs are captured in the same livetrap, suggesting that they may forage together (Monadjem 1998a). In Swaziland, mean distance between successive monthly captures (on a 1 ha grid) was 18.6 m, indicating that movements are limited to a small area. In captivity, copulation is preceded by grooming of " by her mate (Willan & Meester 1978). Females aggressively defend their nests when young are present. Self-grooming commences at 10–11 days of age and continues through adulthood. Reproduction and Population Structure Poorly known for such an abundant species. Reproduction peaks in the wet season (Nov–Feb) but may continue throughout the year. Mean litter-size 4.5 (n = 4, KwaZulu–Natal, South Africa; Taylor 1998) and 4.0 (range 1–7, n = 27, in captivity; Willan & Meester 1978). Gestation (in captivity): 18–19 days. At birth, young hairless, weight 0.8 g. Incisors erupt Day 7–9. Young weaned Day 18. Sexual maturity attained Day 42 (Willan & Meester 1978). Mean interval between litters 22 days. Population numbers fluctuate widely at some localities (Monadjem 1999b) but not at others (Monadjem 1998b). Survival rates are low and few individuals live to one year of age. Sex ratio does not deviate from parity (Monadjem 1999b).

Mus minutoides.

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Predators, Parasites and Diseases Preyed upon by several species of owls (Vernon 1972, Perrin 1982). Ectoparasites include 12 species of fleas, and three species of ticks (details in De Graaff 1981). Susceptible to plague in laboratory conditions. Conservation IUCN Category: Least Concern. Widespread and abundant throughout its range.

E: 8.5 (6–11) mm, n = 11 WT: 6.2 (4–12) g, n = 16 GLS: 18.8 (17.5–20.4) mm, n = 4 GWS: 9.6 (8.8–10.3) mm, n = 4 M1–M3: 3.0 (2.8–3.2) mm, n = 4 Body measurements: Swaziland (Monadjem 1998a) Skull measurements: South Africa (Roberts 1951) Key References De Graaff 1981; Downs & Perrin 1996; Monadjem 1999b; Smithers 1983; Willan & Meester 1978.

Measurements Mus minutoides HB: 54.8 (45–68) mm, n = 16 T: 41.0 (38–49) mm, n = 16 HF: 12.5 (11–14) mm, n = 15

A. Monadjem

Mus musculoides WEST AFRICAN PYGMY MOUSE Fr. Souris naine d’Afrique de l’Ouest; Ger. Westafrikanische Zwergmaus Mus musculoides Temminck, 1853. Esquisses Zool. sur la Côte de Guiné, p. 161. ‘Côte de Guine’, West Africa. Exact locality uncertain.

Taxonomy Subgenus Nannomys. The taxonomic relationship of this species with M. minutoides of eastern and southern Africa is uncertain. Within West Africa, this species has been to referred to both M. musculoides and M. minutoides. The taxon musculoides has been placed as a subspecies of M. minutoides (Petter & Matthey 1975), as a valid species (Rosevear 1969, Happold 1987, Musser & Carleton 1993, Grubb et al. 1998), or placed in synonymy with L. minutoides (Musser & Carleton 2005). The species occurs throughout West Africa; however, its eastern geographic border is uncertain (see Distribution); Musser & Carleton (1993) consider this to be Ethiopia and Somalia, but R. Hutterer (pers. comm.) places the eastern limit asW Cameroon. Hence the identity of C African populations is questionable. Rosevear (1969) placed Mus musculoides in the subgenus Leggada, now a synonym of Nannomys (Musser & Carleton 1993). See also Mus minutoides for

further comment. Synonyms: bella (or bellus), enclavae, gallarum, gondokorae, grata, marica, paulina, petila, soricoides, sungarae, sybilla, vicini (Musser & Carleton 1993). Subspecies: none. Chromosome number shows Robertsonian polymorphism: at Ippy, Central African Republic, 2n = 25 to 2n = 32, with 2n = 28 and 2n = 29 forming about 60% of population (n = 31); another study at Ippy, 2n = 38, FN = 33 and 34 (n = 4) (Jotterand-Bellomo 1984); in Côte d’Ivoire, 2n = 33 (n = 6) and 2n = 33 (n = 2) (Jotterand-Bellomo 1984); in S Nigeria, 2n = 32 (n = 1), 2n = 33 (n = 4) and 2n = 34 (n = 7) (R. Matthey pers. comm.); in Mali, 2n = 18 (""), 2n =19 (!!) (n = 8) (Veyrunes et al. 2004). Description Very small delicate mouse with short tail and white ventral pelage. Dorsal pelage golden-brown, flecked with dark brown; hairs pale grey at base, golden-brown at tip. Flanks similar in colour to dorsal pelage, becoming paler on lower flanks. Head similar in colour to dorsal pelage; small white subauricular patch.Ventral pelage pure white. Colour of flanks and ventral pelage clearly delineated. Chin, chest, fore- and hindlimbs white. Tail short (ca. 70% of HB) and slender, pale brown above, whitish below. Total length of skull more than 17 mm (cf. M. haussa). Nipples 2 + 2 = 8. Geographic Variation None except in karyotype (seeTaxonomy). Similar Species M. setulosus. Larger (HB: 81.8 [77–86] mm, HF: 14.1 [14–16] mm, M1–M3: 3.58 [3.5–3.6] mm); dorsal pelage darker. M. haussa. On average smaller (HB: 49.7 [44–52] mm, HF: 12 [11.5– 12.5] mm, M1–M3: 2.9 [2.9–3.1] mm); dorsal pelage paler; distribution further north, mostly in Sudan and Sahel Savanna BZs. M. mattheyi. On average smaller (HB: 52 [46–60] mm, HF: 12 [11.5– 12.5] mm, M1–M3: 2.9 [2.9–3.1] mm); Ghana, distribution limits and status uncertain.

Mus musculoides (western Africa) and Mus minutoides (southern Africa). See also text.

Distribution Endemic to Africa. Widely distributed in Guinea Savanna BZ, Northern Rainforest–Savanna Mosaic, and grassy patches in Rainforest BZ. May extend into southern parts of Sahel Savanna

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Mus musculus

BZ in west of range. Recorded from Gambia and Senegal to Cameroon, and may extend further eastwards through N DR Congo, Central African Republic and Uganda to Ethiopia and Somalia (see above). Western extension of range to East Africa, and where M. minutoides replaces M. musculoides, also uncertain (shown by striped lines on map). Geographic range has extended southwards into ‘savanna-like’ habitats within the Rainforest BZ in recent years as a result of humaninduced activities (clearing of rainforest, farming, urbanization). Habitat Savanna woodlands and grasslands, as well as new and old farmlands, secondary bush and immature cocoa plantations. Grassy areas within Rainforest BZ, such as road verges, farmlands and clearings. Does not occur in rainforest except, very occasionally, in natural grassy clearings where trees have fallen. Abundance At 14 widely separated savanna habitats in W Nigeria, comprised on average 14% (0–53%) of captured small rodents (Happold 1975b), and at Olokomeji F. R. (in Rainforest– Savanna Mosaic) they formed 56% of all small rodents in grasslands at certain times of year (Anadu 1973). Relative abundance (and absolute numbers) declines from south to north in W Nigeria from 90% in Rainforest–Savanna Mosaic to 11–41% in southern Guinea Savanna BZ and (usually) 0% in northern Guinea Savanna BZ (where Tatera kempi [now Gerbilliscus kempi], Myomys daltoni [now Praomys daltoni] and Mastomys spp. are common species) (Happold 1975b). In natural clearings in rainforest in Ghana they comprise ca. 2% of small rodents, the fifth most numerous species after Lophuromys sikapusi, Praomys tullbergi, Dephomys defua and Hybomys trivirgatus (Jeffrey 1977). Population numbers vary seasonally and annually in W Nigeria (Anadu 1973). Numbers tended to be lowest at the end of the wet season (Oct/Nov), increasing during the dry season and early wet season, and declining gradually during the late wet season. Density: 2–4/ha (end of wet season) to 7–10/ha (end of dry season). In some years, highest density is 23–35/ha (early dry season). Adaptations Terrestrial and nocturnal. Construct spherical or cup-shaped nests of shredded grass under logs and in shallow burrows. Locomotion is a rapid scuttle; when disturbed, individuals hide under logs, dry grass and any other available cover. Numbers decline markedly where savanna is burned during dry season (probably mostly due to emigration), but immigrants from unburnt areas return as soon as grasses sprout again (Anadu 1973). Foraging and Food Herbivorous. In captivity, feeds on small seeds, grass stems and fruits. Social and Reproductive Behaviour

many individuals huddle together in a nest; huddling is probably an important aspect of thermoregulation in this small species, especially during cooler days of the dry season. Reproduction and Population Structure Reproduction is seasonal in S Nigeria.Young born Aug–May; most litters at end of dry season and beginning of wet season (Mar–Apr), and at end of wet season (Sep–Dec). Embryo number (wild-caught ""): 3.38 (2–6, mode 2, n = 13); embryo number (captive ""): 3.00 (1–5, mode 3, n = 18 litters from seven ""; Anadu 1976). Females may have several litters in close succession. Gestation: 22–24 days. Litter interval (in captivity): 41–58 days. At birth, young are naked, eyes are closed, and mean WT = 0.8 g. External ear open Day 11. Eyes open Day 14. Weaned at Day 24. Almost (92%) adult HB length by Day 30; adult size by Day 60. Sexual maturity 10–12 weeks. Longevity: probably not longer than ca. one year. Mice born at beginning of wet season (Apr) attain sexual maturity during the wet season and begin to breed in Aug; mice born at end of wet season (Nov–Dec) are not mature until the beginning of next breeding season in Mar–Apr. Hence continual recruitment of young throughout most of year, and high annual turnover (Anadu 1973). No details from other parts of range. Predators, Parasites and Diseases Preyed upon by owls. Second most numerous species in pellets of Barn Owls Tyto alba on Mt Nimba, Liberia (Heim de Balsac & Lamotte 1958). In N Nigeria, comprised 4–8% of prey numbers (total n = 64) in pellets of Barn Owls and 3% of prey numbers (total n = 83) in pellets of Spotted Eagle-owls Bubo africanus (Demeter 1981). Conservation IUCN Category: Least Concern (as M. minutoides). Widespread and not threatened. Measurements Mus musculoides HB: 60.8 (55–70) mm, n = 8 T: 43.6 (32–55) mm, n = 8 HF: 13.6 (12–14) mm, n = 8 E: 9.6 (9–10) mm, n = 8 WT: 8.5 (6.5–10.3) g, n = 8 GLS: 19.3 (18.3–20.2) mm, n = 8 GWS: 9.5 (8.9–10.0) mm, n = 8 M1–M3: 3.1 (2.6–3.2) mm, n = 8 W Nigeria (Happold 1987) Key References

Anadu 1976; Happold 1975b, 1987. D. C. D. Happold

Gregarious. In captivity,

Mus musculus HOUSE MOUSE Fr. Souris domestique; Ger. Haus-Maus Mus musculus Linnaeus, 1758. Syst. Nat., 10th edn., 1: 62. Uppsala, Sweden.

Taxonomy Subgenus Mus. Many forms or subspecies of Mus musculus have been described, most of which are now considered as synonyms (see Musser & Carleton 1993, 2005) although others

may be species (Marshall & Sage 1981). The taxonomy of the ‘house mouse’ has been less well studied in Africa than elsewhere (but see Granjon et al. 1992, Prager et al. 1998). Recent research suggests 487

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that the ‘correct’ name of the domestic mouse is likely to be Mus domesticus (Boursot et al. 1993, 1996, Din et al. 1996, Prager et al. 1998). One taxon (M. spretus of North Africa and S Europe), originally included in Mus musculus, is now considered to be a valid species. Traditionally, there appear to be two forms of Mus musculus: a domestic form (which lives only in houses and has a relatively longer tail) and a feral form (which lives in human-modified habitats and has a relatively shorter tail). Synonyms that refer to Africa include brevirostris, domesticus, gentilis, praetextus, vignaudi (all pale-bellied forms from North Africa – see Osborn & Helmy 1980, Aulagnier & Thévenot 1986) and modestus (dark-bellied form from southern Africa (see Meester et al. 1986). Musser & Carleton (2005) provide a full list of synonyms. Chromosome number: 2n = 40, FN = 38 (Niger: Dobigny et al. 2002b; Senegal: Granjon et al. 1992); 2n = 22 (Robertsonian populations in Tunisia; Said & Britton-Davidian 1991). Description Very small greyish mouse, normally associated with houses. Dorsal pelage greyish or brownish-grey; hairs dark grey at base, pale grey or brown at terminal end, sometimes with black tip. Flanks paler. Ventral pelage buffy-brown, pale grey, or white, merging into colouration of flanks. Head rather pointed, dark ‘beady’ eyes; ears large, mostly naked. Fore- and hindlimbs short, greyish, with small naked unpigmented digits. Tail long (ca. 90–100% of HB), thin, almost naked, slightly darker above than below. Nipples: 3 + 2 = 10. Geographic Variation Throughout its range, M. musculus shows large differences in colour and pattern. For Africa, the following forms may be recognized: gentilis (including praetextus): North Africa, Sudan, Ethiopia. Pale (usually white) ventral pelage. modestus (? domesticus): North Africa; South Africa (and perhaps Namibia and Zimbabwe). Buffy-brown or grey ventral pelage. In North Africa, both dark and pale-bellied forms are usually distributed in a geographical mosaic in separate farms and oases (Marshall & Sage 1981). Similar Species Mus (Nannomys) spp. On average smaller body size; tail relatively shorter (i.e usually less than 75% of HB); natural habitats (Table 37). Distribution Not indigenous to Africa. Occurs in most countries of the world, but not widespread in Africa. Recorded in temperate Africa, especially in the northern parts of Morocco, Algeria, Libya and Tunisia, and in the Nile Valley. In Algeria and Libya, known from some oases in the desert. Occurs up to 2000 m in the High Atlas of Morocco. Introduced to sub-Saharan Africa by Arab and European shipping, but has formed permanent populations only in selected localities in South Africa (including Marion I.), Namibia, Zimbabwe and Senegal (north of Dakar). Mostly rare in tropical Africa, and recorded from only a few localities near the coastline where populations may not be permanent. Of the three species of small murid rodents introduced into Africa, M. musculus has not colonized the continent to the same extent as Rattus rattus but has been more successful than R. norvegicus.

Mus musculus

Habitat In Africa, mostly commensal and confined to houses, food stores and other urban buildings, although a few populations are feral (e.g. in Morocco [Aulagnier & Thévenot 1986], Algeria [Kowalski & Rzebik-Kowalska 1991] and southern Africa [Smithers 1983]). In Libya, recorded as commensal and quite widely distributed in the wild where the habitat is mesic and has a dense cover of plants (Ranck 1968). In Egypt, also found in a variety of natural habitats including gardens, barley fields, sand flats, salty waste land (with Suaeda and other halophytic plants), canal banks and palm groves (Osborn & Helmy 1980). In N Sudan, recorded only from Khartoum during winter months, but not in villages away from the Nile R. (Happold 1967c). In Senegal, House Mice are strictly commensal (J.-M. Duplantier, unpubl.). Abundance In North Africa, may be fairly abundant in suitable habitats when conditions are optimal. Little information for Africa south of the Sahara; Smithers (1983) records that, in southern Africa, House Mice never occur in sufficient numbers to become a major problem, a contrasting situation to temperate regions of the world, and to some countries where they have been introduced, where they may be major pests, e.g. Australia. Adaptations Nocturnal and diurnal; terrestrial and scansorial. Extreme adaptability with respect to food and climate has enabled House Mice to colonize many habitats in most parts of the world. Most populations are commensal, some are feral. House Mice require adequate water (as free water or in food) and hence do not survive in arid habitats (unless close to a river, a man-made water supply, or in oases). Nevertheless, in Senegal, they live in drier areas than Rattus rattus. Populations are able to breed (and remain viable) in a wide range of climates where the ambient temperatures are very high (as in hot deserts) or very low (sub-Antarctic islands). Rapid reproduction (see below) enables populations to increase rapidly when environmental conditions are good.

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Mus neavei

Foraging and Food Omnivorous. Primarily granivorous and herbivorous, but will feed on anything edible. Occasionally feed on insects and earthworms.

& Traub 1965b). Several human diseases can be carried by House Mice including tularaemia, leptospirosis, borreliosis and several Salmonella-like bacteria that cause food poisoning.

Social and Reproductive Behaviour Communal and sociable. In established groups, !! defend an area of variable size (which is related to food availability). When populations are low, any increase in resources promotes rapid reproduction; when populations are high, reproductive rate declines, and at very high densities and when food is limiting, reproduction may cease completely. House Mice are good colonizers, and in a very short period of time can become very numerous (Corbet & Southern 1977).

Conservation IUCN Category: Least Concern. House Mice are major pests in some parts of the world, and control of their populations is essential. They eat and contaminate stored foods, and are a threat to public health in many parts of the world. The domestic white mouse, used extensively for medical research, is derived from this species.

Reproduction and Population Structure Little information for Africa. Very prolific. In Egypt, pregnant "" and nestling young present throughout year (Osborn & Helmy 1980). Typically, in other parts of the world, "" may produce 5–10 litters/year. Gestation: 19–20 days; fertilization may occur during postpartum oestrus. Littersize: ca. 5.5 (4–8). Young weaned by Day 20. Sexual maturity when 7.5 g ("") to 10 g (!!) (Corbet & Southern 1977). Rapid reproduction maintains high population numbers even when predation rates are high. Predators, Parasites and Diseases Many small mammalian carnivores, birds of prey and snakes feed on House Mice. Ectoparasites include several species of fleas and a sucking louse.The fleas Xenopsylla cheopis, X. ramesis, Pulex irritans, Nosopsyllus londiniensis and Synosternus cleopatrae are commonly found on House Mice in Egypt (Hoogstraal

Measurements Mus musculus HB: 83.7 (78–91) mm, n = 10 T: 77.2 (66–85) mm, n = 10 HF: 18.0 (16–20) mm, n = 10 E: 13.7 (13–15) mm, n = 9 WT: 13.4 (9–20) g, n = 9 GLS: 21.6 (21.0–22.3) mm, n = 11 GWS: 11.2 (9.9–12.1) mm, n = 9 M1–M3 (alveolar): 3.6 (3.3–3.8) mm, n = 9 Egypt (Osborn and Helmy 1980) Key References (Africa only) Corbet & Southern 1977; Kowalski & Rzebik-Kowalska 1991; Osborn & Helmy 1980; Ranck 1968; Smithers 1983. D. C. D. Happold

Mus neavei NEAVE’S PYGMY MOUSE Fr. Souris naine de Neave; Ger. Neaves Zwergmaus Mus neavei (Thomas, 1910). Ann. Mag. Nat. Hist., ser. 8, 5: 90. Petauke, Loangwe district, E Zambia. 2500 ft (762 m).

Taxonomy Originally described in the genus Leggada. Subgenus Nannomys. Described as Leggada neavei (Thomas 1910a). Belongs to the ‘sorella group’ (see Mus sorella). Considered as a subspecies of M. sorella (Verheyen 1965b, Smithers 1983, Meester et al. 1986), although Petter (1981) suggested that it should be treated as a valid species. Relationship to M. oubanguii and M. baoulei is uncertain (Musser & Carleton 2005). Synonyms: none. Chromosome number: not known. Description Small mouse with rich tawny pelage. Dorsal pelage ochraceous-brown to rich tawny-brown, tending to blackish on middorsal region.Ventral pelage pure white. Colour of dorsal pelage and ventral pelage clearly delineated on lower flanks. Head with pointed muzzle. Ears large, pale grey. Fore- and hindfeet white. Tail short (ca. 43% of HB). Skull: rostrum elongated, incisors slightly pro-odont, choanae V-shaped; M1 with small prelobe. Nipples: not known. Geographic Variation None recorded. Similar Species M. sorella. Mean HB smaller; mean GLS larger; mean M1–M3 longer; dorsal pelage dark greyish-brown. Mus neavei

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M. minutoides. Mean HB smaller (51.3 [45–63] mm); mean M1–M3 slightly smaller (3.0 (2.8–3.2) mm); dorsal pelage brownish-buff (variable).

Habitat Woodland savanna.

Measurements Mus neavei HB: 88.8 (58–106) mm, n = 5 T: 38.4 (33–48) mm, n = 5 HF: 13 (12–14) mm, n = 6 E: 11 (10–12) mm, n = 6 WT: n. d. GLS: 18.5 (18–18.9) mm, n = 6 GWS: 9.4, 9.7 mm, n = 2 M1–M3: 3, 3.5 mm, n = 2 E Zambia (MNHN)

Remarks Apparently no other information on this species.

Key Reference Thomas 1910a.

Distribution Endemic to Africa. Zambezian Woodland BZ. Recorded from S Tanzania, S DR Congo, SE Zambia, S Zimbabwe, W Mozambique and South Africa (former Transvaal). Limits of geographic range unknown. Records from Malawi represent other species (Ansell & Dowsett 1988).

Conservation

F. Petter

IUCN Category: Data Deficient.

Mus orangiae ORANGE PYGMY MOUSE Fr. Souris naine d’Orange; Ger. Orange Zwergmaus Mus orangiae (Roberts, 1926). Ann. Transvaal Mus. 11: 251. Viljoensdrift, near Vereeniging, Kruisementfontein, South Africa.

Taxonomy Originally described in the genus Leggada. Subgenus Nannomys. Formerly considered a subspecies of Mus minutoides (e.g. De Graaff 1981; Meester et al. 1986) but now considered as a valid species. Considered to be allied, on craniological grounds, to Mus setzeri (Vermeiren & Verheyen 1983). Synonyms: none. Chromosome number: not known.

relatively long and rounded; white subauricular patch (probably) absent. Limbs short with white feet and well-developed digits; four digits on forefeet; five digits on hindfeet.All digits with well-developed claws.Tail short (ca. 60% of HB), brownish above, paler below. Nipples: 2 + 2 = 8. Geographic Variation

Description Very small mouse with soft pelage. Dorsal pelage bright orange-buff; hairs with pale grey base; orange-buff at tip; black tips of some hairs result in a slightly grizzled appearance. Flanks orange-buff, without any black-tipped hairs. Ventral pelage pure white. Head with pointed nose and long vibrissae. Ears brownish,

None recorded.

Similar Species M. minutoides. Similar size and shape; dorsal pelage reddish-brown with some black-tipped hairs; may differ craniologically; marginally parapatric. M. indutus. On average slightly smaller; HF usually larger; dorsal pelage generally paler; tail pale above; marginally sympatric. M. setzeri. Similar size; rump white; allopatric. Distribution Endemic to Africa. Highveld BZ. Recorded from C South Africa and Lesotho. Taxonomic status of Mus occurring in the southern part of the Free State, South Africa, uncertain, but may represent M. orangiae. Limits of geographic range not known. Habitat Lynch (1983, as M. minutoides but presumably referring to M. orangiae) records that in the Free State, specimens were mostly collected in short, open grassland. Abundance ‘Mus minutoides is not abundant in the Orange Free State’ (Lynch 1983). No other information. Remarks Nocturnal and terrestrial; constructs a grass nest amongst piles of rocks or in disused termitaria (Lynch 1983). Conservation IUCN Category: Least Concern. However, too little is known about this species to be able to assess its conservation status adequately.

Mus orangiae

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Mus oubanguii

Measurements Mus orangiae HB (!!): 59.6 (52–63) mm, n = 8 HB (""): 60.7 (56–69) mm, n = 6 T (!!): 37.5 (36–39) mm, n = 8 T (""): 37.8 (36–40) mm, n = 6 HF (!!): 12.9 (12–14) mm, n = 8 HF (""): 13.5 (12–14) mm, n = 6 E (!!): 11.9 (9–14) mm, n = 8 E (""): 12.7 (12–13) mm, n = 6 WT: n. d.

GLS: 18.5, 18.7 mm, n = 2 GWS: 9.0, 9.7 mm, n = 2 M1–M3: 3.0, 3.3 mm, n = 2 South Africa Body measurements: TM Skull measurements: Roberts 1951 Key Reference

Lynch 1983. A. Monadjem

Mus oubanguii OUBANGUI PYGMY MOUSE Fr. Souris naine de l’Oubangui; Ger. Oubangui-Zwergmaus Mus oubanguii Petter and Genest, 1970. Mammalia 34: 454. Bangassou near La Maboké (near Mongoumba), Central African Republic.

Taxonomy Subgenus Nannomys. Synonyms: none. Chromosome number: 2n = 28; FN variable; polymorphism is non-Robertsonian (Matthey & Jotterand 1970, Jotterand 1972, Jotterand-Bellomo 1984). Description Very small reddish-brown mouse with pure white ventral pelage. Dorsal pelage reddish-brown.Ventral pelage pure white. Colour of dorsal pelage and ventral pelage clearly delineated on flanks. Head with pointed muzzle. Ears large, blackish, slightly pointed at tip. Large white postauricular patch. Fore- and hindfeet white. Hindfeet comparatively large (cf. M. mattheyi). Tail short (ca. 60% of HB). Skull: rostrum elongated; incisors orthodont or slightly pro-odont; choanae V-shaped; anterior palatal foramina short; M1 and M2 well developed, M3 small; M1 with well-developed prelobe and without accessory cusp; M1 with anterior lobe quadricuspidate. Nipples: 3 +2 = 10.

Geographic Variation

None recorded.

Similar Species M. goundae. Similar size; dorsal pelage ochraceous-brown; nipples 2 + 2 = 8; chromosome number: 2n = 16–19; N Central African Republic. M.musculoides. Similar size; dorsal pelage golden-brown; no postauricular patch; nipples 2 + 2 = 8; chromosome number: 2n = 25–32 (polymorphic); common and widespread. M. setulosus. Much larger (HB: 81.8 [77–86] mm, T: 55.5 [52– 59] mm); dorsal pelage blackish-brown; no auricular patches; chromosome number: 2n = 36; widespread. Distribution Endemic to Africa. Northern Rainforest–Savanna Mosaic. Recorded only from the type locality at Bangassou near La Maboké, and at Ippy on the right bank of Oubangui R., SW Central African Republic. Habitat Savanna patches on sandy slightly lateritic soil, close to rainforest, with grasses (mostly Loudetia arundinacea) and trees (mostly Hymenocardia acida, Annona senegalensis, Lophira alata and introduced Borassus aethiopium).Type locality (and study site – see Genest-Villard 1973) was an unburnt reserve. Sympatric with M. setulosus and M. minutoides (Petter & Genest 1970). Abundance Uncertain. In suitable habitats, 10–25 burrows/ 100 m2 (Genest-Villard 1973). Distribution is very localized. Adaptations Nocturnal and terrestrial. During the day, rests in simple underground burrows. Burrows are up to ca. 60 cm in length, with a spherical nest chamber (ca. 20 cm diameter) and an escape entrance. The nest is lined with leaves of grass and trees. The main entrance is blocked with fragments of grass and dead leaves. Burrows may be constructed close of those of Tatera (now Gerbilliscus) spp. and occasionally the burrows of the two species join up (Genest-Villard 1973).

Mus oubanguii

Foraging and Food Mainly granivorous. In captivity, eats seeds and small insects. 491

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Social and Reproductive Behaviour Normally nest alone, except when " has her young. Adult !! nest alone. Reproduction and Population Structure Litter-size: 4–5. Births recorded in May and Jun (no data for other months) (GenestVillard 1973). Predators, Parasites and Diseases No information. Conservation IUCN Category: Data Deficient. The very limited geographic range, isolation of populations and decline in habitat are cause for concern.

Measurements Mus oubanguii HB: 62.7 (50–75) mm, n = 50 T: 38 (26–44) mm, n = 29 HF: 13.9 (13–15) mm, n = 50 E: 11.5 (9–14) mm, n = 50 WT: n. d. GLS: 20 (18.6–21.5) mm, n = 26 GWS: 10 (9.5–10.7) mm, n = 21 M1–M3: 3.5 (3.2–3.9) mm, n = 26 Central African Republic (MNHN) Key References

Genest-Villard 1973; Petter & Genest 1970. F. Petter

Mus setulosus PETERS’S PYGMY MOUSE Fr. Souris naine de Peters; Ger. Peters Zwergmaus Mus setulosus Peters, 1876. Monatsb K. Preuss. Akad. Wiss. Berlin: 480. Victoria, Cameroon.

Taxonomy Subgenus Nannomys. One form (proconodon) from Ethiopia was considered to be a valid endemic species by Yalden et al. (1976). Synonyms: pasha, proconodon. Subspecies: none. Chromosome number: 2n = 36, FN = 36 (Matthey 1966a, Jotterand-Bellomo 1986). Description Small dark-coloured mouse; the largest of pygmy mice in West Africa. Pelage short, slightly coarse (cf. soft in other pygmy mice); comparatively long (7 mm). Dorsal pelage blackishbrown, usually ‘dull’, very finely speckled with buff (without the ‘bright’ russet colouration of M. musculoides); hairs grey at base, buff or black at tip. Lower flanks slightly paler, with fewer black-tipped hairs. Ventral pelage pure white or off-white. Head similar in colour to dorsal pelage. Ears darkly pigmented, with sparse short grey or buff hairs. Upper lips with narrow white fringe (not wide as in M. haussa or M. musculoides). Auricular patches absent. Fore- and hindfeet white. Tail short (ca. 67% of HB), dark above and below, with small scaly rings and scattered short dark bristles. Skull: larger than in other species of pygmy mice in West Africa (see also below); total length of skull ca. 21 mm; anterior palatal foramina 4.8 (4.4– 5.1) mm; M2 with small anterio-external cusp (Rosevear 1969).

recorded from E Nigeria (Gotel Mts,Yelwa), S Cameroon (Adamaoua Mts, Victoria) and Gabon (Hutterer & Joger 1982, Hutterer et al. 1992, Grubb et al. 1998). Scattered localities in Central African Republic, NE DR Congo and S Sudan. Isolated populations in Ethiopia (as proconodon). Not recorded from Benin and S Nigeria, where perhaps it occurs. One locality in SE DR Congo (not mapped) needs verification. Habitat Most records are from relict forests in the Rainforest BZ, and from gallery forests and grasslands in montane habitats, e.g. gallery forest (E Nigeria, Cameroon), montane grassland and gallery forest (Liberia), tropical deciduous forest and cultivated areas of cleared tropical forest (Ethiopia), and grassfields at edge of forest on

Geographic Variation Individuals from Cameroon (see Measurements) are larger than proconodon (see Taxonomy) from Ethiopia. Similar Species M. haussa. Smaller in most measurements; dorsal pelage sandy; auricular patches present; geographic range further north. M. musculoides. Smaller; dorsal pelage buff or russet, usually rather bright; auricular patches present; sympatric. Distribution Endemic to Africa. Rainforest BZ (Western and West Central Regions), Northern Rainforest–Savanna Mosaic and Afromontane–Afroalpine BZ of Ethiopia. Most locality records are from E Sierra Leone, Liberia, Côte d’Ivoire, Ghana and Togo; also

Mus setulosus

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Mus setzeri

Mt Nimba (Guinea). Recorded from 1000 to 1750 m in Ethiopia (where it is considered a ‘lowland species’ – Yalden et al. 1976), at 1550–2300 m in highlands of Cameroon and Nigeria, and 500– 1300 m on Mt Nimba, Liberia. Abundance Generally uncommon, and not captured as often as sympatric M. musculoides. May be locally common, as in parts of S Cameroon (Eisentraut 1973). Remarks There is no detailed biological information on this species. Rosevear (1969) suggested that it is probably very similar to M. musculoides.

Measurements Mus setulosus HB: 81.8 (77–86) mm, n = 6 T: 55.5 (52–59) mm, n = 6 HF: 14.1 (14–16) mm, n = 6 E: 11.8 (11–12) mm, n = 6 WT: n. d. GLS: 21.1 (19.3–21.9) mm, n = 4 GWS: 9.9 (9.6–10.2) mm, n = 4 M1–M3: 3.6 (3.5–3.6) mm, n = 4 Cameroon (BMNH) Key Reference

Conservation

Rosevear 1969.

IUCN Category: Least Concern. D. C. D. Happold

Mus setzeri SETZER’S PYGMY MOUSE Fr. Souris naine de Setzer; Ger. Setzers Zwergmaus Mus setzeri Petter, 1978. Mammalia 42: 377. 82 km west of Mohembo, Botswana.

Taxonomy Subgenus Nannomys. Craniologically most similar to Mus orangiae. Synonyms: none. Chromosome number: not known. Description Very small mouse with soft pelage. Dorsal pelage variable shades of pale buff or pale buffy-orange; hairs slate-grey at base; some have black tip giving a slightly grizzled appearance (though less so than in M. indutus). Flanks lack black-tipped hairs and are buffy-orange. Ventral pelage pure white; white colour extends high up on flanks, and dorsally across rump to form white band anterior to tail – a unique character of this species. Head with pointed nose and long vibrissae. Chin, cheeks and muzzle white. Ears comparatively

long (cf. M. minutoides) and rounded, pale brown; small white patch at base of each ear. Limbs short with white feet; four digits on forefeet; five digits on hindfeet. All digits with well-developed claws. Tail short (ca. 60% of HB), whitish. Nipples: not known. Geographic Variation

None recorded.

Similar Species M. minutoides. Dorsal pelage generally darker, tail dark above; white ventral pelage does not extend on to upper rump and muzzle; allopatric. M. indutus. Ears shorter; white ventral pelage does not extend on to upper rump and muzzle; sympatric. Distribution Endemic to Africa. Parts of South-West Arid Zone (Kalahari Desert) and Zambezian Woodland BZ. Recorded only from NE Namibia, NW Botswana and W Zambia, with a single record from S Botswana (Vermeiren & Verheyen 1983). Perhaps present in SE Angola. The record from S Botswana suggests a wider geographic range than currently known. Habitat Poorly known. In Botswana, recorded from the fringes of pans and wetlands in relatively arid habitats (mean annual rainfall 400–450 mm). Abundance No information. The species is poorly represented in museum collections, suggesting that it is not abundant. Remarks

Apparently no other information available.

Conservation IUCN Category: Least Concern. The restricted geographic range, apparent rarity and lack of records from protected areas may be cause for concern. Mus setzeri

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M1–M3: 3.3 (3.3–3.5) mm, n = 18 Body measurements and weight: Botswana, unsexed individuals (as Leggada sp.; Smithers 1971) Skull measurements: Zambia (Vermeiren & Verheyen 1983) *From specimen label (TM)

Measurements Mus setzeri HB: 61 mm, n = 1* T: 36 (31–48) mm, n = 9 HF: 14 (13–15) mm, n =11 E: 13.6 (13–14) mm, n = 11 WT: 6.8 (5–9) g, n = 11 GLS: 18.0 (17.5–18.3) mm, n = 11 GWS: 10.1 (9.6–10.5) mm, n = 17

Key References

Petter 1978; Vermeiren & Verheyen 1983. A. Monadjem

Mus sorella SORELLA PYGMY MOUSE (THOMAS’S PYGMY MOUSE) Fr. Souris naine de Thomas; Ger. Zentralafrikanische Zwergmaus Mus sorella (Thomas, 1909). Ann. Mag. Nat. Hist., ser. 8, 4: 548. Kirui, Mt Elgon, W Kenya. 6000 ft (1830 m).

Taxonomy Originally described in the genus Leggada. Subgenus Nannomys. A species within the ‘sorella group’, characterized by the presence of V-shaped choanae, which also includes the closely related M. goundae, M. neavei and M. oubanguii. Phylogenetic relationships within the ‘sorella group’ need systematic revision (Musser & Carleton 1993, 2005). Synonyms: acholi, kasaicus, wamae. Subspecies: none. Chromosome number: not known. Description Very small greyish mouse with pure white ventral pelage. Dorsal pelage dark greyish-brown, with pale-coloured speckles; hairs grey at base, beige or pale brown at tip; some hairs on back longer with black tip. Flanks paler, due to presence of rufous or pale brown hairs and absence of black-tipped hairs. Ventral pelage pure white. Colour of dorsal pelage and ventral pelage very clearly delineated high up on flanks (more so than in other species of Mus) without a brownish-orange line of delineation. Head long. Ear comparatively long, dark brown, sparsely covered with short fine hairs. Small white subauricular spot. Upper lips, lower cheeks, chin, throat and chest pure white. Fore- and hindfeet white. Tail short (ca. 65% of HB). Skull: rostrum slender; choanae V-shaped; incisors proodont, anterior palatal foramina long, reaching to the anteriolateral cusp of M1. Nipples: 2 + 2 = 8. Geographic Variation None recorded. Similar Species M. musculoides/minutoides. Similar in total length; HB on average smaller (M. minutoides); tail longer (mean 50 mm, ca. 100% of HB); ear on average shorter (mean 8.9 mm); common and widespread.

Mus sorella

Remarks In Garamba N. P., E DR Congo, sympatric with M. minutoides (Verheyen 1965b). Seeds and fragments of burnt material were the principal items in stomach contents (Verheyen & Verschuren 1966). Preyed on by owls in Kagera N. P., Rwanda (X. Misonne in Verheyen 1965b). Conservation

Distribution Endemic to Africa. Northern and Eastern Rainforest– Savanna Mosaics close to the Rainforest BZ (East Central and West Central Regions). Recorded from NE and SE DR Congo, Uganda, Kenya and N Tanzania (Musser & Carleton 1993). Presence in E Cameroon and EC Angola (Musser & Carleton 2005) uncertain (not mapped). Limits of geographic range unknown. Habitat Savanna grasslands and woodlands, especially where grasses are tall; grassland areas close to gallery forests along rivers (NE DR Congo; Verheyen 1965b).

IUCN Category: Least Concern.

Measurements Mus sorella HB: 59.9 (51–73) mm, n = 22 T: 39.8 (34–46) mm, n = 21 HF: 13.1 (12–14) mm, n = 22 E: 10.8 (10–12) mm, n = 22 WT: n. d. GLS: 19.7 (17.8–21.0) mm, n = 26 GWS: 10.1 (9.3–10.7) mm, n = 20

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Mus spretus

M1–M3: 3.4 (3.1–3.7) mm, n = 29 Garamba N. P., NE DR Congo (Verheyen 1965) Key References

Petter 1981; Verheyen 1965b. F. Petter

Mus spretus ALGERIAN MOUSE (WESTERN MEDITERRANEAN MOUSE) Fr. Souris sauvage; Ger. Algerische Maus Mus spretus Lataste, 1883. Actes Soc. Linn. de Bordeaux, ser. 7, 4: 27. Oued Magra, between M’sila and Barika, N of Hodna, Algeria.

Taxonomy Subgenus Mus. Variously considered as a valid species, or as a subspecies of M. musculus. Karyology, electrophoresis, DNA hybridization and mitochondrial RNA sequences suggest M. spretus is linked with M. spicilegus and M. musculus, and is a member of the clade that includes Mus macedonicus, M. spicilegus (both non-African) and M. musculus (details in Musser & Carleton 2005). Synonyms: caoccii, hispanicus, lusitanicus, lynsei, mogrebinus, parvus, rifensis. Subspecies: none. Chromosome number: 2n = 40, FN = 40. Description Small mouse similar to Mus musculus, but smaller and often with shorter tail. Pelage short and dense. Dorsal pelage brownish, slightly flecked with pale brown; hairs dark grey with pale brown or black tip. Ventral pelage greyish-white, clearly delineated from colour of flanks; hairs grey on basal half, white on terminal half. Head similar in colour to dorsal pelage; face rather pointed. Ears darkly pigmented. Limbs short. Fore- and hindfeet with small sparse white hairs. Tail long (ca. 80% of HB), bicoloured, ringed with very small scales; many extremely small bristles. Notch on inner surface of upper incisors not well developed (cf. M. musculus); sometimes barely visible. Nipples: not known.

Geographic Variation

None recorded.

Similar Species M. musculus. Dorsal pelage dark grey; ventral pelage grey; tail on average longer (66–85 mm, 90–100% of HB); notch on inner surface of upper incisors well developed; commensal. Apodemus sylvaticus. Larger in all dimensions; tail ca. 107% of HB; often syntopic. Distribution Mediterranean Coastal BZ and coastal regions of Sahara Arid BZ. In Africa, recorded from Morocco, Algeria, Tunisia and Libya. In Algeria, extends from sea level to the northern parts of Haut Plateaux (Kowalski & Rzebik-Kowalska 1991), and in Morocco from coastal regions near Tangiers to limits of cultivation in the subSahara as well as some oases (Aulagnier &Thévenot 1986). Distribution in Libya uncertain; reference to the ‘wild form’ of M. musculus (Ranck 1968) probably refers to M. spretus, which is commoner in the interior than near the coast. Extralimital in S France, Spain, Portugal and Balearic Islands. Probably indigenous to the Mahgreb and subsequently expanded northwards into southern Europe (Dobson 1998, Dobson & Wright 2000). Habitat In Algeria, recorded from the seashore to alpine meadows at 1600 m (Kowalski & Rzebik-Kowalska 1991). Tends to prefer habitats with sparse woody vegetation interspersed with open ground, as well as agricultural fields (Khidas et al. 2002). In Morocco, lives on plains and hills, as well as cultivated areas, grasslands and forests (Aulagnier & Thévenot 1986). Although largely sympatric with M. musculus, it is usually not syntopic and is not commensal (cf. M. musculus). Abundance Varies greatly according to habitat. Commonest species of small mammal in mixed woodlands (dominated by Phylliria latifolia, Pinus halepensis and Arenia maritima) near sea level in Algeria (M. spretus 46%, Gerbillus campestris 32%, Apodemus sylvaticus 12%, Lemniscomys barbarus 9%; n = 43). In an oak–olive forest, comprised 80% of two spp. of small mammals (n = 10) (Khidas 1993). Rare or absent in five other sampled habitats from 250 to 1800 m (Khidas 1993). Highest population numbers are often in agricultural fields.

Mus spretus

Adaptations Terrestrial. Individuals appear to be less capable of coping with cold conditions than M. musculus. Experimentally, in both species, when Ta = 5–35 °C, Tb = 35–37 °C. When Ta is reduced to 0 °C,Tb decreased to ca. 33 °C (whereas for M. musculus,Tb remained 495

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normal). At Ta = 20–25 °C, oxygen consumption (as a measure of metabolic rate) was 3–7 cm3/g/h for both species; when Ta = 0 °C, metabolic rate remained at 3–7 cm3/g/h in M. spretus, but increased to 11–20 cm3/g/h in M. musculus (S.D. ±0.63–±1.07). Although results are variable (and for M.musculus depend on where the individuals originated), M. spretus seems less capable of altering metabolic rate (and therefore maintaining Tb) when Ta is low than M. musculus (Gorecki et al. 1990).

Predators, Parasites and Diseases Probably common in owl pellets in Algeria, although difficult to distinguish from M. musculus (Kowalski & Rzebik-Kowalska 1991). In Morocco, comprised 65% of small prey in pellets of Barn Owls Tyto alba (Aulagnier & Thévenot 1986). Ectoparasites include a flea, Nosopsyllus barbarus (Beaucornu & Kowalski 1985).

Foraging and Food Mainly fruits and seeds (Aulagnier &Thévenot 1986, Khidas et al. 2002).

Measurements Mus spretus TL: 139.7 (125–155) mm, n = 41 T: 62.1 (55–71) mm, n = 44 HF: 16.0 (14–18) mm, n = 44 E: 12.8 (12–15) mm, n = 41 WT: 13.9 (12–16) g, n = 7 GLS (CbL): 19.8 (18.3–21.8) mm, n = 40 GWS: 11.0 (10.0–12.1) mm, n = 41 M1–M3: 3.6 (3.3–3.9) mm, n = 44 Algeria (Kowalski & Rzebik-Kowalska 1991)

Social and Reproductive Behaviour

No information.

Reproduction and Population Structure In Algeria, !! with enlarged testes from May–Nov, and lactating "" from Jun–Nov (Kowalski & Rzebik-Kowalska 1991). Embryo number: 4 (n = 1). In S Spain (a climate similar to that of the Mahgreb), reproductive season is from Mar–Nov, with peaks in reproductive activity in Apr– May and Aug–Sep (Antunez et al. 1990). Reproductive activity is probably associated with food abundance, and therefore varies from year to year (Duran et al. 1987, in Fons & Saint Girons 1993).

Conservation

Key References 1991.

IUCN Category: Least Concern.

Khidas et al. 2002; Kowalski & Rzebik-Kowalska D. C. D. Happold

Mus tenellus DELICATE PYGMY MOUSE Fr. Souris naine délicate; Ger. Zarte Zwergmaus Mus tenellus (Thomas, 1903). Proc. Zool. Soc. Lond. 1903: 298. Roseires, Blue Nile, Sudan.

Taxonomy Originally described in the genus Leggada. Subgenus Nannomys. Morphologically and ecologically similar to M. haussa. Setzer

(1956) placed aequatorius and delamensis (in the Sudan) as subspecies of bellus (now a synonym of M. musculoides), but Musser & Carleton (1993, 2005) allocated these forms to M.tenellus. Synonyms: aequatorius, delamensis, gerbillus, suahelicus (Petter 1972b, Musser & Carleton 1993). Subspecies: none. Chromosome number: not known. Description Very small pale-coloured mouse, similar in colour to M. haussa. Dorsal pelage pale sandy, sometimes darker on middorsal line. Ventral pelage pure white. Colour of dorsal pelage and ventral pelage clearly delineated on flanks. Chin and throat pure white. Ears short, grey. White sub- and postauricular patches (which may form a white ring around the base of each ear); the only species of Mus with such extensive patches. Fore- and hindfeet white. Hindfeet narrow.Tail of moderate length (ca. 70% of HB), scaly, with very small bristles, dark above, pale below. Skull: choanae U-shaped, anterior palatal foramina small, rounded; incisors opisthodont; M1 56–61% of M1–M3, M3 15–20% of M1–M3; M1 with anterior lobe tricuspidate (cf. M. haussa). Nipples: 2 + 2 = 8. Geographic Variation

None recorded.

Similar Species No other species of Mus has such an extensive postauricular patch (or tuft) of white hairs (see Description). Mus tenellus

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Mus triton

M. haussa. Similar size; no postauricular patch of white hairs; allopatric. M. minutoides/musculoides. On average larger; dorsal pelage darker (golden-brown flecked with dark brown). Distribution Endemic to Africa. Guinea Savanna and Somalia– Masai Bushland BZs. Recorded from C Sudan, Ethiopia (below 2000 m), S Somalia (not mapped) and southwards through Kenya to C Tanzania (Musser & Carleton 1993, 2005). Limits of geographic range unknown. Habitat

Grass steppe with thicket clumps.

Abundance Uncertain. Rarely collected. In S Ethiopia comprised 1500 m) of Angola. Abundance No information. Remarks Very little is known about this species. Apparently makes a screeching sound ‘like a squirrel’. Lactating !! collected in Aug (Hill & Carter 1941). Conservation

Otomys anchietae

Measurements Otomys anchietae HB (""): 213 (209–217) mm, n = 5 HB (!!): 197 (187–209) mm, n = 6 T (""): 119 (115–127) mm, n = 5 T (!!): 111 (87–121) mm, n = 6 HF (""): 41 (40–41) mm, n = 5 HF (!!): 37 (36–38) mm, n = 6 E: n. d. WT: n. d. GLS: 49.4 (46.2–52.8) mm, n = 5 GWS: 26.5 (25.0–28.0) mm, n = 5 M1–M3: 12.0 (11.3–12.5) mm, n = 5 Body measurements: Chitau, Angola (Hill & Carter 1941) Skull measurements: Angola (Taylor & Kumirai 2001) Key References

Crawford-Cabral 1998; Hill & Carter 1941. P. J. Taylor

IUCN Category: Least Concern.

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Otomys angoniensis

Otomys angoniensis ANGONI VLEI RAT Fr. Rat du vlei d’Angoni; Ger. Angonis-Lamellenzahnratte Otomys angoniensis Wroughton, 1906. Ann. Mag. Nat. Hist., ser. 7, 18: 274. M’Kombhuie, Malawi (= Matipa Forest, Misuku Range, Malawi; fide Ansell & Dowsett 1991). 7000 ft (2120 m).

Taxonomy Otomys angoniensis maximus may be a distinct species (Musser & Carleton 1993, 2005, Crawford-Cabral 1998), but morphometric overlap (Bronner & Meester 1988, P. J.Taylor unpubl.) and genetic similarity (Maree 2002) suggest maximus should be retained as a subspecies of O. angoniensis. The Angolan cuanzensis, previously allocated to angoniensis, is recognized as a distinct species following Musser & Carleton (2005). Synonyms: canescens, divinorum, elassodon, mashona, maximus, nyikae, pretoriae, rowleyi, sabiensis, tugelensis. Subspecies: three. Chromosome number: 2n = 56, aFN = 54. All chromosomes are acrocentric. X-chromosome is second largest member of the karyotype and Y-chromosome is smallest. Limited G-band homology detected between this species and O. irroratus (Contrafatto et al. 1992c). Description Large stocky rat with dense shaggy pelage. Dorsal pelage pale to dark greyish-buffy. Colour varies geographically from darker to paler, with the palest individuals on Mt Kilimanjaro. Ventral pelage dark grey. Head large. Small, well-haired ears held close to head. Tail short (ca. 55% of HB). Each upper incisor with single groove. Each lower incisor with one deep and one shallow groove (except for O. a. maximus where the shallow groove is practically invisible). M3 with seven laminae (occasionally six). M1 with four laminae. Nasal bones moderately expanded. Petrotympanic foramen small, slit-like (as in O. burtoni and O. sloggetti). Baculum spatulate basally with central raised portion in ventral view; length of proximal portion ca. 4 mm, maximum width ca. 1 mm (Davis 1973). Body size varies geographically (see Measurements). Nipples: 0 + 2 = 4.

O. tropicalis. Round petrotympanic foramen. O. denti, O. sloggetti and O. unisulcatus. Single groove in each lower incisor. O. typus. Two deep grooves in each lower incisor. O. anchietae, O. barbouri, O. lacustris, O. occidentalis. Five laminae in M1. Distribution Endemic to Africa. Zambezian Woodland BZ and southern part of the Somalia–Masai Bushland BZ; also present in highland habitats throughout the eastern side of the continent. Recorded from South Africa, Swaziland, Zimbabwe, Mozambique, Malawi, Angola, Zambia, DR Congo, Rwanda, Tanzania and Kenya (Ansell 1960, Misonne 1974, Ansell & Dowsett 1988, Bronner & Meester 1988, Crawford Cabral 1998). Records from outlying Eastern Cape Province, South Africa (De Graaff 1981, Bronner & Meester 1988) could not be corroborated on the basis of known museum specimens (Lynch 1994). Specimens having slit-like petrotympanic foramina from Uvira (SMNS) and Albert N. P. (MNHN) extend the known distribution into E DR Congo and Rwanda, respectively. Distribution in East Africa is patchy. See also Geographic Variation above. Habitat Mesic grassland and savanna woodland habitats (from coastal to high montane habitats, mostly at 4.5 mm; P. J. Taylor, unpubl.). Each lower incisor with deep outer and shallow inner groove. O. a. maximus: N Botswana, NW Zambia, SW & SE Angola, SE & E DR Congo. Largest subspecies; HB: 154–207 mm, GLS: 40–49 mm (Roberts 1951, P. J. Taylor unpubl.). Each lower incisor with single groove. O. a. tugelensis: South Africa, SE Botswana. Similar size to O. a. angoniensis, comparatively narrow interorbital distance (2 m3) are necessary for main shelters (Gouat & Gouat 1983). Natural caves and human constructions (retaining walls along roads, piles of stones along fields, small rocky dams) may also be used as shelters. Atlas Gundis avoid forests and dense vegetation cover (e.g. esparto grass Stipa tenacisissima steppe),

Ctenodactylus gundi

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The social group is stable and long lasting; one adult reproductive ", for example, was observed in the same colony in the Aures Mts for more than three years (Gouat 1988a). Members of the colony occupy a common home-range, which remains stable throughout the year, and from one year to the next. The size of home-ranges depends on food abundance, stability of resources and on local social constraints. In the Aures Mts, the home-range of three colonies was 1031 m2 and 5468 m2 in semi-desert habitats, and 685 m2 in a high elevation habitat (Gouat 1991b). The whole of the home-range is equivalent to a territory. It is actively marked and defended against conspecific intruders (Gouat 1991a). Animals deposit scent marks all around their territory, and the entrances to shelters are covered with piles of faecal pellets, which are a good indication that gundis are present (George 1974). Animals mark and dust-bathe at least once each day in a specific place near the main shelter (P. Gouat & J. Gouat 1987). Gundis may chase conspecific intruders. In captivity, even in large cages, intruders are chased and bitten to death (Eisentraut 1977, P. Gouat & J. Gouat 1987). Animals in a colony share a common shelter during the night. In addition to this night shelter, numerous secondary shelters used during the day are scattered throughout the home-range. Activity starts at sunrise. Animals come out of the night shelter, and after a period of amicable interaction around the entrances, disperse throughout their home-range to feed individually or in small groups (Gouat 1988a, 1991a).The territory appears as a mosaic of foraging patches adjacent to their rocky shelters. Atlas Gundis spend a large amount of time sun-bathing on the ground or on rocks. Adults and juveniles cooperate to detect predator and conspecific intruders. Animals take turns to watch the surroundings from the top of rocks and to give alarm calls when a potential predator is detected (Gouat & Gouat 1989). Alarm calls are typically simple, monosyllabic chirps which are emitted in quick succession (up to 20 at a time) (George 1981a), but they vary according to the age of the emitter and the circumstances (Séguignes 1979, Gouat et al. 1985). On hearing alarm chirps, members of the colony stop their activity and stay vigilant.The sudden appearance of a predator elicits one or two very loud chirps, and all the members of the colony run into shelters. Sound emissions are numerous and varied. The repertoire includes audible range communication (vocalizations, tooth chattering and foot drumming) and some ultrasonic vocalizations (Gouat et al. 1985). These sounds are mainly displayed during interactions between conspecifics. At mid-day, animals rest in the shade or inside the shelters, individually or in small groups. The length of the resting period depends on the ambient temperature. In the hot period (Apr–Sep) in the southern part of the Aures Mts, this resting period lasts from 10:00h until 16:00h. During the cold period (Dec–Feb), or in high elevation sites, resting periods may be absent or may last only one hour (Gouat 1991a). In the late afternoon, animals forage again. Around sunset, they return individually to the main shelter. Interactions are rare at this time except when there is a change of night shelter. One animal may stay at the entrance of the old night shelter and drive the incoming animals to the new night shelter (Gouat 1988a). In the semi-desert sites of Aures Mts, reproduction starts in Dec. It begins with the exclusion of supernumerary "" by both the !! and the reproducing " (Gouat 1988a). This time of the year is one of the rare occasions when gundis are aggressive (P. Gouat & J. Gouat 1987). Copulation may occur outside the shelter (Gouat 1985). Prior

to mating, " approaches ! by walking with measured tread with the head bent down slightly, and emits a sexual trill of low intensity (Gouat et al. 1985). He sniffs the genital area of !. Mounting is followed by some long pelvis thrusts, which precede ejaculation. After ejaculation, " remains on ! for ca. 30 sec. In natural conditions, "" have been observed ejaculating five times during a 45 min period with the same or different !! (Gouat 1985). Reproduction and Population Structure Polyoestrous. Two litters are produced each year. In the semi-desert sites of Aures Mts, reproduction begins in late Dec. First litters born in late Feb to early Mar. The birth of young is preceded by a prepartum oestrus (Gouat 1985; see also C. vali). Second births occur between late Apr and mid-May. In high altitude sites of the Aures Mts, this reproductive schedule is delayed by two months. Females have two litters per year only under favourable conditions, and scarcity of food may preclude any reproduction and promote the exclusion of supernumerary "" (Nutt, 2005). Gestation: 73 days. Litter-size: 2 (1–3, n = 17; Gouat 1985). At birth, young are precocial, fully furred, with the eyes and ears open and functional, and able to walk and to chew solid food (Gouat 1985). Birth-weight: 29.9 ± 11.9 g (range 18–40 g, n = 15; Gouat 1985). Weaning: ca. Week 6. Males and !! do not breed until the reproductive season following their birth, when aged 7–9 months of age (Gouat 1985). Detailed physiological studies on reproduction are required. Communal nursing is the rule, but young have seldom been observed sucking any !! other than their mother (J. Gouat & P. Gouat 1987). Young gundis emit an extension trill when approaching their mother, or when the mother leaves them after feeding (Gouat et al. 1985). This vocalization accompanies lordosis and anogenital eversion by the young. The extension trill is attractive to adults and may elicit similar trills in other young animals (J. Gouat & P. Gouat 1987).Young animals emit a modulate trill during sucking (Gouat et al. 1985). During their first week, young animals remain near the main shelter watched over by an adult ! or a juvenile. Adults may carry the young animals in their mouth, one by one, from one shelter to another (J. Gouat & P. Gouat 1987). At the end of the second week, play behaviour (running and head-shaking) and young vocalizations tend to disappear, and agonistic behaviour develops. At three weeks of age, young animals display most behavioural patterns of the adult but sexual behaviour remains absent until the next reproductive season (J. Gouat & P. Gouat 1987). Young animals from about two weeks of age feed mainly on vegetation, but weaning seldom occurs before six weeks of age (Gouat & Gouat unpubl.). Females may supplement young with water through long-lasting mouth to mouth contact (Gouat 1988a). Three types of population structure have been observed dependent on the bioclimatic conditions. (1) In high altitude habitats, and in habitats with a Mediterranean climate, where mean annual rainfall is ca. 400 mm and food resources are abundant from spring to autumn, high levels of reproduction and a high turnover of the population are observed. Colonies in these habitats, however, may disappear suddenly if devastated by summer storms or by a long period of snow cover during winter (Séguignes 1979, Gouat & Gouat 1983). (2) In semi-desert habitats, populations are more stable, and the level of reproduction is correlated with the abundance of resources. Summer is the most difficult period of the year for gundis because the vegetation becomes dry and sparse. When the conditions are good enough, and 631

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the resources remain abundant, mortality may be low during summer. In this case, in summer or autumn, a group composed of adults and juveniles may migrate to establish themselves in a vacant site (Gouat 1988b). (3) In desert habitats on the edge of the Sahara Desert, where mean annual rainfall is less than 100 mm, social units are generally composed of only one breeding pair. Generally, animals are unable to survive in the desert habitats when there is more than one year without any rain. The maintenance of populations under these conditions depends primarily on recolonization from more favourable sites (Gouat & Gouat 1982, Séguignes & Vernet 1998). Predators, Parasites and Diseases Gundis are hunted and eaten by humans in the Maghreb. The most dangerous predators are snakes because they are able to enter the rocky shelters of gundis, even during the night. The presence of a large adder (Daudin’s Viper Vipera lebetina), 1.20 m long, caused a colony of gundis to change their main shelter and to modify the way they used their home-range (Gouat 1991a). Other predators are dogs, foxes, birds of prey and ravens (Séguignes 1979, Gouat 1988a).

Measurements Ctenodactylus gundi HB: 193 (150–228) mm, n = 25 T: 31 (20–45) mm, n = 20 HF: 38 (33–44) mm, n = 113* E: 18 (14–22) mm, n = 113* WT: 268 (185–396) g, n = 22 GLS: 48.6 (42.9–53.5) mm, n = 44 GWS: 32.1 (27.8–35.9) mm, n = 43 M1–M3: 9.3 (7.5–10.8) mm, n = 48** Algeria and Tunisia (MNHN); Aures Mts (Algeria) and from their descendants born in captivity (P. Gouat & J. Gouat unpubl.) *Beni Kheddache, Tunisia (K. J. Nutt unpubl.) **P4 may occur in some adults; if so cheekteeth measurement is P4–M3 Key References Gouat 1985; Gouat, P. 1991b; Gouat & Gouat, 1983; Gouat & Gouat 1989; Gouat et al. 1985. Patrick Gouat

Conservation

IUCN Category: Least Concern.

Ctenodactylus vali THOMAS’S GUNDI Fr. Goundi de Thomas (Goundi du Sahara); Ger. Thomas Gundi Ctenodactylus vali Thomas, 1902. Proc. Zool. Soc. London 1902: 11. ‘Wadi Bey’ (NW of Bonjem, Tripoli), Libya.

Taxonomy Corbet (1978) included vali as a synonym of gundi, but George (1982) listed vali and gundi as separate valid species (see also Dieterlen 2005c). The form joleaudi (originally described as a species, C. joleaudi) is considered to be a subspecies of vali by Petter (1961) and subsequent authors. Synonyms: joleaudi. Subspecies: none. Chromosome number: 2n = 40 (George 1979b).

Description General appearance of a small guinea-pig with a short hairy tail and small ears. Pelage very dense and soft. Dorsal pelage buffy-brown with dull grey underfur. Ventral pelage yellowish-grey. Head flat and broad. Forehead narrow and slightly hooked. Muzzle short with long black vibrissae. Nostrils naked and black. Eyes large and round surmounted by long vibrissae. Ears flattened on the head, oval, black inside bordered by a dense fringe of short stiff whitishgrey hairs. Fore- and hindlimbs short; four digits on each foot. Digits of the hindfeet are surmounted by comb-like bristles. Claws sharp, not reaching the ground in normal position. Tail relatively short (ca. 102% of HF), usually hidden by pelage of rump and not visible. Skull: see family and genus profiles; each upper incisor with single groove; angular process of mandible elongated posteriorly forming the most posterior part of the mandible (as in Massoutiera, cf. Felovia, Pectinator); hugely inflated auditory bullae (ca. 17.9 mm, 39% of GLS), distance between bullae ca. 7 mm. Emits chirp alarm call when disturbed. Mean length of faecal pellets: 7.9 (7.3–8.2) mm. Nipples: 2 + 0 = 4. Geographic Variation

None recorded.

Similar Species C. gundi. HB on average larger; forehead large and straight; whistle alarm call; less inflated auditory bullae; mean length of faecal pellets >9.5 mm; partially parapatric. Massoutiera mzabi. HB similar; tail relatively longer; present in northern Sahara; allopatric. Ctenodactylus vali

Distribution Endemic to Africa. Northern edge of Sahara Arid BZ. Distributed in two discrete and widely separated areas of NW

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Africa: (1) Algeria and Morocco in the west, and (2) Libya in the east. Morocco: hamada of Wadi Ziz. Algeria: Saharan Atlas from Beni Ounif to Bechar, basin of Wadi Saoura, and hamada of Wadi Guir, Ougarta Mts. Libya: Tripolitania, transitional desert between Soda Mts and the Gulf of Sirte (Ranck 1968). Often parapatric with C. gundi. See also below. Habitat Wide range of rocky desert habitats such as mountain slopes, edges of hamadas, wadis, rift and small mountainous massifs. Also found in semi-desert areas, on the southern slopes of the Saharan Atlas in Algeria and SE Morocco. Abundance Generally sparsely distributed, but high density may be observed under favourable climatic conditions during the breeding period (e.g. 18 animals/ha; George 1981a). Adaptations Thomas’s Gundis are diurnal and have similar activity patterns to Atlas Gundis: they are active at sunrise and in the early morning, rest at the hottest time of the day, and are active again in the afternoon before sunset. Thomas’s Gundis are parapatric with Atlas Gundis on the western edge of the Saharan Atlas and in the southeastern part of Morocco. In semi-desert habitats, the highly structured occupation of space displayed by Atlas Gundis prevents any extension of the geographic range of Thomas’s Gundis further to the west. However, Thomas’s Gundis are capable of rapid colonization of sites left vacant by Atlas Gundis following a period of severe drought (Gouat 1988b). The present distribution of Thomas’s Gundis in two isolated areas of distribution is best explained by competition with Atlas Gundis and the climatic fluctuations of the Sahara Desert. The relationships between the two species may be explained as follows (Gouat 1988a): (1) Ctenodactylus gundi, thought to belong to the ancestral species, originally occupied their present range. Following an extension of the Sahara Desert to the north, populations of Ctenodactylus gundi gradually moved away and settled in the northern mountains (e.g. Tel Atlas Mts in Algeria). Several rocky corridors may have allowed this resettlement from southern sites to northern sites (e.g. Hodna Mts in Algeria). (2) Ctenodactylus vali appears to have evolved in one of the populations of Ctenodactylus gundi remaining in the Saharan Atlas, in response to the drought caused by the extension of the Sahara Desert to the north. This new species extended its range throughout the range occupied previously by Ctenodactylus gundi where desert conditions prevail. (3) Regression of the Sahara Desert to the south enabled Ctenodactylus gundi to return to its previous range, and as a consequence, the range of Ctenodactylus vali became smaller and restricted to more arid desert habitats. As a result, the range of Ctenodactylus vali was split into two parts and the species is present only in habitats where Ctenodactylus gundi is unable to survive. Foraging and Food Herbivorous, feeding mainly on grasses and herbaceous plants.The diet encompasses food plants such as Eremophyton chevallieri, Amberboa leucantha, Cymbopogon sp. and Aristida sp. (George 1974). Animals forage mainly in early morning and late in the afternoon, alternating foraging with resting. Animals pick up food

items while they travel through their home-range, without having any clear foraging places.The area of foraging each day varies from 20 m2 to 275 m2 (Gouat 1988a). Social and Reproduction Behaviour Thomas’s Gundis, in contrast to Atlas Gundis, are mainly solitary; social bonds are weak, and animals behave as ‘floaters’ for a large part of the year. At the end of autumn, ! settles at a place where she will produce her young. Several "" may try to join !, but only one " will succeed. A familiarization period is necessary for " to become accepted by !. In captivity, reproduction was successful in groups composed of one ! and 2–3 "" (Grenot 1973); "" develop a hierarchy (George 1978b). In captivity, confrontation between unfamiliar animals may cause the death of the intruder (Grenot 1973, Gouat 1988a). During these agonistic encounters, animals emit different vocalizations, including the ‘trilled whistle’ and the ‘repeated whistle’, and non-vocal sounds such as foot-drumming and tooth-chattering (J. Gouat 1991). The repeated whistle is composed of a repetition of short whistles (mean frequency of the plateau: 4.5 KHz; each unit lasts between 0.12 and 0.5 sec). This vocalization is also displayed in the case of a sudden alert and is named the ‘alarm call’ by George (1981a); on hearing this call, conspecifics respond by adopting an alert posture, or by disappearing inside a shelter. The trilled whistle is emitted only in an agonistic context. It consists of a modulate whistle followed by a trill. The trilled whistle is emitted alone or coupled with the alarm call. Thomas’s Gundis have two litters each year (see also below). In Dec in Djeniene Bou Rezg (Beni Ounif region, Algeria), individuals settle in a suitable habitat after several months of nomadic existence and become sedentary. A ! and several "" establish a territory where ! will later give birth to her young. By Jan, ! and one of the "" have developed a social bond; and the supernumerary "" have disappeared. The pair may spend the night in a common shelter but each forages independently. First copulation occurs in the days following pair formation. Male stays with ! during gestation but leaves her soon after the prepartum oestrus and no later than the birth of the first litter in Jan–Mar. Young animals are precocial but have difficulty in moving around and remain close to the shelter. Even before they are weaned, the mother leaves her young at night. She joins them in the morning, and may spend the afternoon rest with them.Young animals emit a calling trill quite similar to the extension trill of the Atlas Gundi (J. Gouat 1991), but without lordosis or anogenital eversion by the young. This vocalization attracts the mother and the littermates. Littermates continue to use a common shelter during the night.Young animals of the first litter disperse before the birth of the second litter in Apr–May. In captivity, mothers may become aggressive towards their young of the first litter in the days preceding birth of the second litter. In natural conditions, however, dispersion does not seem to be caused by the mother.Young animals disperse simply by extending their home-range. The mother uses a similar strategy with her second litter. Even while she is still lactating, the mother spends the night alone. She forages on her own, and shortly before her young are weaned, she increases her home-range significantly. In Djeniene Bou Rezg (Beni Ounif region, Algeria), the size of the daily home-range varies from 100 to 500 m2 in Dec–Mar, and increases to 1125 m2 in May and Jun. At this time of the year, the mother is still lactating. There is no significant correlation between the size of the daily home-range, either with food abundance or with the presence 633

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of young. From day to day, animals use different parts of their habitat and the percentage of overlap between the daily home-range on two consecutive days varies from 12% to 33%, with a minimum in May and Jun. In the early summer, mothers leave their reproductive homerange and become nomadic. The young animals may stay for a while on their natal home-range, but soon become nomadic and solitary. Animals become sedentary again in Dec, with the beginning of the next reproductive season.

to survive. After four years without significant rainfall, Thomas’s Gundis were still present in the Beni Abbes region but at a very low density (Gouat & Gouat 1984).When the climatic conditions became favourable again, reproduction recommenced (Gouat 1988a).

Reproduction and Population Structure Two litters per year under favourable conditions; first litter in late Jan–Mar; second litter Apr–May. Gestation: 2 months. Litter-size: 2 (1–3), n = 21 litters (combined data from Grenot 1973, George 1978b, Gouat 1988a). Females are lactating and pregnant with the second litter at the same time.Young precocial, fully furred, with eyes and ears open and functional at birth. Weaned: 1–2 months. Young reach sexual maturity and adult size in Dec when aged 7–9 months. In times of severe drought (more than one year without rain), there is no sedentary period and no reproduction. First copulations occur at the end of the autumn (late Nov–Jan). The period of mating may vary between animals of a given region. In the Taghit region, two !! were captured 20 km apart during the same week of Mar, and one gave birth four weeks before the other (Gouat 1988a). As in Atlas Gundis, a prepartum oestrus is suspected for the second copulation. Female is able to store spermatozoa in her genital tract for at least two months, and to use these spermatozoa without further copulation in order to produce a replacement litter (Gouat 1986). Young animals disperse soon after weaning and recolonization of vacant sites occurs rapidly (Gouat 1988b). In times of severe drought, reproduction stops and the density of local populations decreases slowly. At such times, Thomas’s Gundis may extend their home-range in order to find sufficient resources

Conservation IUCN Category: Data Deficient. Probably not threatened. Human population density is very low in the geographic range of the species; any decreases in population density are probably due to climatic changes.

Predators, Parasites and Diseases Preyed upon by humans, shepherd’s dogs, foxes, birds of prey, snakes (Gouat 1988a) and jackals (George 1974).

Measurements Ctenodactylus vali HB: 160 (124–185) mm, n = 13 T: 36 (27–43) mm, n = 13 HF: 35 (30–39) mm, n = 9 E: 17 (15–20) mm, n = 9 WT: 129.6 (87–180) g, n = 7 GLS: 47.8 (45.1–50.9) mm, n = 19 GWS: 32.0 (28.4–34.4) mm, n = 20 M1–M3: 8.4 (7.4–9.7) mm, n = 20 Algeria (MNHN); and individuals from the Taghit region (Algeria) and their descendants born in captivity (J. Gouat & P. Gouat unpubl.) Key References George 1974; J. Gouat, 1986, 1991; P. Gouat, 1988b; Grenot 1973. Patrick Gouat

GENUS Felovia Felou Gundi Felovia Lataste, 1886. Le Naturaliste 7 (36): 287. Type species: Felovia vae Lataste, 1886.

Felovia is a monotypic genus, restricted to the semi-desert rocky habitat of the Felou Hills of the upper Senegal R. in Mali and Mauritania. Originally proposed as a subgenus of Massoutiera but recognized as a valid genus by Thomas (1913) and St Leger (1931). Characters of the genus include: palate extends posteriorly to the cheekteeth, toothrows converge anteriorly, upper molar teeth simple with large infoldings on both sides (Figure 100), and tail about double the length (ca. 221%) of the hindfoot (Misonne 1974); further details are given in the species profile. The single species is Felovia vae. Patrick Gouat Felovia vae.

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Figure 100. Skull and mandible of Felovia vae (BM 19.7.7.3698).

Felovia vae FELOU GUNDI Fr. Goundi du Félou; Ger. Felou-Gundi Felovia vae (Lataste, 1886). Le Naturaliste, 7 (36): 287. Upper Senegal River, Felou Hills, Medine, south of Kayes, Mali.

Taxonomy Originally described in the genus Massoutiera. Phylogenetic evidence suggests this species is closely related to Massoutiera mzabi (George 1979a, 1985a). Synonyms: none. Chromosome number: 2n = 36 (George 1979a).

genus profiles; each upper incisor with single groove; angular process of mandible not elongated posteriorly; auditory bullae not greatly inflated (ca. 13.8 mm, 29% of GLS, least inflated of all species of gundis). Mean length of faecal pellets: 10.6 (10.0–10.9) mm. Nipples: 2 + 0 = 4.

Description General appearance of a small guinea-pig with small ears and a short movable hairy tail. Pelage very dense and soft. Dorsal pelage reddish-brown. Ventral pelage russet. Head flat and broad. Ears small, with a whitish tuft at the base but without white on the back of the ear; flattened on the head. Fore- and hindlimbs short; four digits on each foot. Digits of the hindfeet surmounted by comb-like bristles. Tail relatively long (ca. 221% of HF), longer than in other gundis (except P. spekei). Tail is folded back on the rump when the gundi is at rest, but begins to flick as soon as the gundi begins to move. Skull: see family and

Geographic Variation

None recorded.

Similar Species Massoutiera mzabi. Upper incisors not grooved or slightly grooved; auditory bullae greatly inflated. All other species of gundis are allopatric. Distribution Endemic to Africa. Sahel Savanna BZ. Known only from Felou Hills, upper Senegal R. in Mali, and the Tagant and Adrar regions in Mauritania. Suspected to occur in Senegal. Habitat Long deep fissures of ancient sandstone hills, in semi-desert habitats with some trees (e.g. Adenium obesum, 2 spp. of fig trees) and shrubs (Tephrosia mossiensi) (George 1974). In Mauritania, animals have been found in an oasis on the rocky banks of wadis, on rocky mountain slopes, on the edges of hamada (F. Colas pers. comm.) and on gueltas (i.e. water ponds) where they drink free water (Vale et al. 2012). Abundance Very limited geographic distribution, but in selected localities up to 25 animals/ha (George 1981a).

Felovia vae

Remarks Herbivorous. During the dry season (Mar) feeds on leaves of a leguminous shrub Tephrosia mossiensis, dropped petioles of fig trees, dry grass and seeds (George 1974). Food resources fluctuate seasonally. Felou Gundis lives in family groups, not far from other family groups. They are noisy animals and in alert situations they emit repeated ‘chee-chee-chee’ calls (George 1981a). They occupy the same shelters for a long time; in 1972, they were still present at the same site where Lataste found them in 1885 (George 1974). Births recorded between mid-Dec and Jan. One young per litter (George 1978b). African Wild Cats Felis sylvestris are likely predators (George 1974). See Rosevear (1969) for further information. 635

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Conservation IUCN Category: Data Deficient, previously assessed as Vulnerable. Felou Gundis are the most threatened of all species of gundis (but see Abundance above). They are not preyed on by humans in Mauritania. Measurements Felovia vae HB: 179 (169–190) mm, n = 5 T: 73 (67–80) mm, n = 4 HF: 33.6 (31–37) mm, n = 9 E: 15.7 (14–17) mm, n = 7

WT: 185.8 ± 8 g, n = 10 GLS: 45.1 (42.8–47.3) mm, n = 8 GWS: 29.7 (27.4–31.8) mm, n = 8 M1–M3: 8.5 (7.5–8.9) mm, n = 8** Mauritania (MNHN) Weight: George (1978b) **P4 may occur in some adults; if so, cheekteeth measurement is P4–M3 Key References

George 1974, 1978a, 1981a; Rosevear 1969. Patrick Gouat

GENUS Massoutiera Mzab Gundi Massoutiera Lataste, 1885. Le Naturaliste 7 (3): 21. Type species: Ctenodactylus mzabi Lataste, 1881.

A monotypic genus distributed in desert and semi-desert rocky habitats in the Sahel Savanna BZ and the Sahara Arid BZ. The type species, although originally named Ctenodactylus mzabi by Lataste (1881), was transfered by him to his new genus, Massoutiera, on the basis of the bilobate pattern of the upper molar teeth. Characters of the genus include: palate extends posteriorly to the cheekteeth, toothrows converge anteriorly, upper molar teeth simple with narrow infoldings on both sides, and tail about one and a half times the length (ca. 148%) of the hindfoot (Misonne 1974); further details are given in the species profile.The single species is Massoutiera mzabi. Patrick Gouat

Figure 101. Skull and mandible of Massoutiera mzabi (BMNH 19.7.7.2923).

Massoutiera mzabi.

Massoutiera mzabi MZAB GUNDI Fr. Goundi du Mzab; Ger. Sahara-Gundi Massoutiera mzabi (Lataste, 1881). Bull. Soc. Zool. de France 6: 214. Ghardaia, Mzab, Algeria.

Taxonomy Two other species have been described in this genus (Massoutiera haterti and Massoutiera rothschildi), but although there are slight differences in the two forms, similar differences may be observed within a single population (Petter & Roche 1958). Synonyms: harterti, rothschildi. Subspecies: none. Chromosome number: 2n = 36 (George 1979b).

Description General appearance of a small guinea-pig with small ears and a short, movable hairy tail. Pelage very dense and soft. Dorsal pelage cream to reddish-brown. Ventral pelage paler. Young have a pinker colouration than their parents. Head flat and broad. Forehead large and straight. Muzzle short with long black vibrissae. Nostrils naked and black. Eyes large and round surmounted by long vibrissae.

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Ears flattened on head, oval, black inside bordered by a dense fringe of short stiff whitish-grey hairs. Fore- and hindlimbs short; four digits on each foot. Digits of the hindfeet are surmounted by comb-like bristles. Claws are sharp, not reaching to the ground in normal position. Tail of medium relative length (ca. 148% of HF), longer than in Ctenodactylus spp. but shorter than in Felovia vae; tail conspicuous (with flicking movements) when gundi is moving. Skull: see family and genus profiles; each upper incisor with single faint groove or without groove; angular process of mandible elongated posteriorly, but not to the extent as in Ctenodactylus; auditory bullae hugely inflated (ca. 18.2 mm, 40% of GLS) (Figure 101). Mean length of faecal pellets: 7.3 (6.6–7.9) mm. Nipples: 2 + 0 = 4. Geographic Variation None recorded. Similar Species Ctenodactylus gundi and Ctenodactylus vali. Tail smaller, less visible; unilobate pattern of the upper molar teeth. Felovia vae. Tail longer; upper incisors each with single groove; less inflated auditory bullae. Pectinator spekei. Tail similar size; ears only partially flattened on the head; occurs only on the Horn of Africa. Pectinator spekei and Felovia vae are allopatric, and there is no risk of confusion with M. mzabi in the field. Distribution Endemic to Africa. Sahara Arid BZ (northern edge) and Sahel Savanna BZ of NW Africa. Range discontinuous in Algeria, Libya, Mali, Niger and Chad. Algeria (north): Mzab region, Oued Mya Bassin, Tademait. Algeria (south): Mouydir, Tefedest, Hoggar (= Ahaggar) Mts and Tassili n’Ajjer. Libya: Maghidet Plateau, Akakus, Massak Mallat, Massak Mastafat, Al Haruj al Aswad, Djebel Sawda, Djebel Al Hasawinah. Mali: Adrar des Ifoghas. Niger: Aïr Massif. Chad: Tibesti Massif. The distribution in Libya is not shown on the map.

Massoutiera mzabi

Because of the close phylogenetic relationship of this species to Felovia vae, and its behavioural characteristics (Gouat 1991a), it is highly probable that Massoutiera mzabi evolved in the Sahel and spread northwards across the Sahara when the climate was less arid. The present geographical range of the species is discontinuous and is probably a relict of a formerly more widespread distribution (Jaeger 1977b, George 1988). Habitat Mountainous regions of deserts and semi-deserts where rocks are present, but also in rolling hills of Mzab, and on the edges of hamadas in the Wadi Mya basin and Tademait. Abundance Patchily distributed in family groups (see below), and never abundant. Isolated animals are common, mainly during the summer. Density in the Mzab region (Algeria): 0.3–3.2 animal/ ha (George 1981a, Gouat 1988a). Adaptations

See family profile.

Foraging and Food Herbivorous. Animals forage individually but may forage close to each other. In the Mzab during winter (Nov–Mar), they forage continuously from sunrise to sunset. In contrast, during the summer (Apr–Sep) they are partly nocturnal – foraging begins before sunrise and continues until 10:00h, when animals retreat into the shade or into their rocky shelters; foraging resumes in late afternoon and may continue until after sunset (Gouat 1991a).There is no specific place for foraging; animals collect food items while they travel through their territory. Principal foods are the leaves, flowers, seeds and stalks of several species of herbs and grasses including Moricandia arvensis, Reseda villosa, Launea angustifolia, Stipa retorta, Peganum harmala, Chrysanthemum macrocarpum, Centaurea incana, Limonium sinuatum, Scabiosa arenaria, Odontospermum pygmaeum (George 1988). Social and Reproductive Behaviour In the Mzab region of Algeria, adult !! are sedentary and territorial. An adult ! occupies a permanent territory (1000–2000 m2) throughout the year. In autumn (Sep–Dec), " joins ! in her territory, and stays with her until the first mating (Nov–Dec) and the birth of the first litter (Jan– Mar). Depending on food availability, he remains with the ! until at least the time of the second mating (Mar–Apr), and at most until the beginning of summer (Jun). During the period of cohabitation, " and ! cooperate to actively defend the territory against conspecific intruders. Mzab Gundis may modify their use of space from one day to another; night and day shelters may differ every day. An alarm call may be emitted in response to a potential predator. This vocalization is a whistle that begins with a sudden decrease of frequency from an ultrasonic frequency far exceeding 16 KHz, and then reaches a plateau at 11–13 KHz. When the animal is worried by an unusual situation, the frequency of the plateau may fluctuate between 8 and 12 KHz. Both adults and juveniles emit alarm calls. The same vocalization is used during agonistic encounters. After the birth of the young, " may watch them from a distance but rarely interacts with them during their first month of life. During this period, the young remain in the vicinity of their shelter. During the day or at night, ! may carry her young, one at a time, in her mouth from one shelter to another, holding the young across its body. During 637

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daytime, the mother comes occasionally to feed the young and seems to spend at least a part of the night with them.This maternal behaviour appears to serve as a defence against predators (Gouat 1988a). In the presence of their mother, young gundis emit a ‘modulated whistle’, part of which is a ‘hiss’. The whistle starts at 2 KHz and ends as an ultrasonic vocalization far exceeding 16 KHz, and is emitted in successive pulses, each lasting 0.15–0.35 sec.The ‘modulated whistle’ appears to have the same function as the ‘extension trill’ of the Atlas Gundi. When the young grow older, they become more mobile and follow the mother when they wish to suck. During exploration, juveniles emit low frequency chuckles (1–4 KHz) (J. Gouat pers. comm.). Juveniles interact amicably with both adult "" and !!. With the arrival of the second litter, ! becomes aggressive towards her first young in order to avoid contact between juveniles and her new young. Juveniles are not aggressive to the young. Young of both litters remain within the territory of their mother until the beginning of the summer. Juvenile "" then disperse, while juvenile !! tend to stay with their mother for a longer time and try to establish themselves in a vacant territory near their mother. The formation of a group of related !! seems to be possible. Reproduction and Population Structure Two litters each year in the Mzab region, Algeria. First mating: Nov–Dec. Birth of first litter: Jan–Mar; second litter: Apr–May. Gestation: ca. nine weeks (captive animal; Gouat 1988a). Litter-size: 2 (1–3), n = 17 litters (combined data from George 1978a, Gouat 1988a). Data from other localities is inconclusive: at Mt Baguezan (Aïr, Niger), three !! with foetuses were captured in May; at Hoggar (Algeria), one ! with two well-developed foetuses obtained in mid-Apr (Thomas & Hinton

1921); at Al Haruj al Aswad (Libya), one ! with a ca. two-week old young in late Dec. Predators, Parasites and Diseases Preyed on by humans, canids, birds of prey and snakes (Gouat 1988a). Conservation

IUCN Category: Least Concern.

Measurements Massoutiera mzabi HB: 176 (125–210) mm, n = 16 T: 52 (33–85) mm, n = 14 HF: 35 (33–36) mm, n = 10 E: 17 (15–17) mm, n = 10 WT: 200 (132–234) g, n = 7 GLS: 45.5 (39.8–48.5) mm, n = 24 GWS: 29.3 (25.3–32.5) mm, n = 24 M1–M3: 8.4 (7.8–9.2) mm, n = 23* Algeria, Chad and Mali (MNHN) and Berriane region (Mzab, Algeria), and from their descendants born in captivity (J. Gouat & P. Gouat unpubl.) *P4 may occur in some adults; if so, cheekteeth measurement is P4– M3 Key References al. 1984.

George 1988; P. Gouat 1988a, 1991a; Gouat et Patrick Gouat

GENUS Pectinator Speke’s Pectinator Pectinator Blyth, 1856. Journ. Asiatic Soc. Bengal, for 1855, (2) 24: 294 [1856]. Type species: Pectinator spekei Blyth, 1855.

Monotypic genus distributed in desert and semi-desert rocky habitats in Ethiopia, Eritrea, Djibouti and Somalia. The characters of the genus include: ears partially flattened on top of head (all other species ‘flattened’); palate does not extend posteriorly to the cheekteeth (cf. all other genera of gundis), toothrows more or less parallel and not converging anteriorly; upper molar teeth simple

with narrow infoldings on both sides, and tail about double the length (ca. 224%) of the hindfoot (Misonne 1974); further details are given in the species profile. The single species is Pectinator spekei. Patrick Gouat

Pectinator spekei SPEKE’S PECTINATOR Fr. Pectinator de Speke; Ger. Buschschwanz-Gundi Pectinator spekei Blyth, 1856. J. Asiat. Soc. Bengal for 1855 (2) 24: 294. Between Goree Bunder and Nogal, Somalia.

Taxonomy Synonyms: legerae, meridionalis. Subspecies: none. Chromosome number: 2n = 40 (George 1979b). Description General appearance of a small guinea-pig with ears only partially flattened on the head, and a short, movable hairy tail. Pelage very dense and soft. Dorsal pelage ashy-grey, suffused with black or brown. Flanks greyish. Ventral pelage greyish-white. Head flat and broad. Muzzle short with long black vibrissae. Ear broadly

ovoid, almost naked, with a fringe of whitish hairs on anterior margin. Fore- and hindlimbs short; four digits on each foot. The digits of the hindfeet have three-tiered combs (George 1978a).Tail hairy and long for a gundi (ca. 224% of HF); white on the basal half, black on terminal half with white tip; tail conspicuous even when gundi is resting. The tail appears whitish along its middle, with two lateral black lines externally fringed with dull white. Skull: see family and genus profiles; each upper incisor without groove; angular process of mandible

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Pectinator spekei

Pectinator spekei.

short, not reaching level of coronoid process; auditory bullae moderately inflated (ca. 14.9 mm, 33% of GLS) (Figure 102). Mean length of faecal pellets: not known. Nipples: 2 + 0 = 4. Geographic Variation None recorded. Figure 102. Skull and mandible of Pectinator spekei (BMNH 14.2.9.2).

Similar Species No other species of gundi lives on the Horn of Africa. Distribution Endemic to Africa. Somalia–Masai Bushland BZ. Recorded from rocky habitats in Ethiopia, Eritrea, Djibouti and Somalia. Habitat In Ethiopia, Speke’s Pectinator inhabits volcanic and limestone rocky cliffs in desert or semi-desert areas. Altitudinal range: 0–1200 m (Yalden et al. 1976). Also lives in man-made rocky escarpments bordering roads (George 1974). Recorded up to 1950 m in N Somalia (N. Redman pers. comm.).

Abundance Speke’s Pectinators live at much higher densities than any other species of gundi. In the Danakil Desert of Ethiopia, densities range from 24 to 237 animals/ha (George 1981a). Adaptations Terrestrial and diurnal. Emerges from rocky shelters at sunrise; maximum activity occurs 0–3 hours after sunrise when ambient temperature is 23–29 °C; retreats to shelters during the daytime when temperature reaches 33–34 °C and emerges again in late afternoon. Pectinators climb trees and may rest on branches in the shade (George 1974). (See also family profile.) Foraging and Food Herbivorous. Feeds on dry grass (stalks and seeds), and on leaves of Caboda rotundifolia, Acacia senegal and A. seyal trees (George 1974). Food resources fluctuate seasonally from abundant during the wet season to scarce during the dry season. Social and Reproduction Behaviour Sociable, living in colonies based on extended family units. When there is a perceived danger, animals emit an alarm call that begins with a chirp, then a long whistle and finally three to six chirps (duration: 0.6–1.5 s, frequency: 1–4 KHz) (George1981a). Pectinators share their habitat with Rock Hyraxes Procavia capensis. Reproduction and Population Structure In the Danakil desert, births occur from late Aug to mid-Sep after the short wet season. Elsewhere in Ethiopia (no precise locality) births occur in ca. Jan. Litter-size: 1.2 (1–2, n = 6). Birth WT: ca. 20 g; young precocial, fully furred at birth. Time to attain adult weight: ca. 174 days. Growth rate is regular (George 1978b). Predators, Parasites and Diseases Gabar Goshawks Micronisus gabar attempt to catch pectinators (George 1974).

Pectinator spekei

Conservation

IUCN Category: Data Deficient. 639

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Measurements Pectinator spekei HB: 170 (155–190) mm, n = 8 T: 74 (60–80) mm, n = 10 HF: 33.1 (30–36) mm, n = 11 E: 19.0 (16–21) mm, n = 11 WT: 178.2 ± 0.9 g, n = 4 GLS: 44.9 (42.5–47.6) mm, n = 13

GWS: 26.5 (25.1–28.9) mm, n = 11 P4–M3: 8.2 (6.9–9.0) mm, n = 13 Body and skull measurements: Djibouti, Somalia and Ethiopia (MNHN) Weight: George (1978b) Key References

George 1974, 1978a, b, 1981a. Patrick Gouat

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Family BATHYERGIDAE

Family BATHYERGIDAE MOLE-RATS Bathyergidae Waterhouse, 1841. Ann. Mag. Nat. Hist., ser. 1, 8: 81. Bathyergus (2 species) Cryptomys (10 species) Georychus (1 species) Heliophobius (1 species) Heterocephalus (1 species)

Dune Mole-rats Mole-rats Cape Mole-rat Silvery Mole-rat Naked Mole-rat

p. 644 p. 648 p. 662 p. 664 p. 667

The Bathyergidae is a polygeneric family, distributed throughout most of tropical and sub-tropical sub-Saharan Africa, in habitats ranging from open forest to savannas and semi-deserts. All species in the family are subterranean, living in soils that range from soft coastal sands to soils that are extremely hard when dry. There are five genera and at least 15 species, all endemic to Africa (Ellerman 1940, De Graaff 1964a, 1981, Skinner & Smithers 1990, Faulkes et al. 1997b, Bennett & Faulkes 2000). The genera Georychus and Bathyergus are restricted to South Africa, Heterocephalus and Heliophobius occur only in eastern Africa, and Cryptomys occurs in eastern, western and southern Africa. Several genera are sympatric and syntopic, especially in southern Africa. The Bathyergidae is one of the most specialized families of African rodents, and has been the subject of many studies. All members of the family show morphological adaptations to life underground. Head and body cylindrical in shape, without distinct neck. Limbs short. Fore- and hindfeet have five digits each and naked soles; the radiale (scaphoid) and intermedium (lunar) carpal bones are separate, a feature unique to Bathyergidae and Ctenodactylidae. Pelage short and thick; longer sensory hairs are present over much of the body and are especially numerous on the head. Skin very loosely attached and, in those genera with hair, can be shaken vigorously (20– 25 vibrations/sec) to clean pelage of soil. Digits and outer borders of the feet fringed with stiff hairs. Tail very short and fringed with stiff hairs in all genera except Heterocephalus. Eyes very small and unable to form images, usually kept closed. Lacks ear pinnae, opening of external auditory meatus is slightly raised; hearing is good. Nostrils housed in a flat horseshoe-shaped nasal area situated above prominent extra-buccal incisors. Muscular folds, covered with medially directed stiff hairs, meet behind the incisors and keep soil out of the mouth when digging. Skull tends to be stoutly built; infraorbital foramina small and secondarily reduced, varied in shape and dimensions and in the thickness of the bone on the outer wall; cheekteeth (premolars and molars) strongly hypsodont and rooted, number varying in number in the different genera. Upper incisors pro-odont and sharp, with either a single groove on the anterior surface of each incisor (Bathyergus) or ungrooved (all other genera). Upper incisors of Bathyergus are rooted above the anterior cheekteeth, those of other genera more procumbent and rooted posteriorly to the cheekteeth (Ellerman 1940, De Graaff 1964a, 1981). Lower incisors ungrooved. The angle of the lower jaw strongly flared outwards to allow passage of a specialized portion of the superficial masseter lateralis muscle.The two halves of the mandible not firmly ankylosed, permitting lateral splaying of the lower incisors (Jarvis & Bennett 1991; contra De Graaff 1981). Dental formula varied, but usually I 1/1, C 0/0, P 1/1, M 3/3 = 20. Heliophobius is unusual because the complete cheekteeth formula (P and M) i s 6/6, but at any one time it is 5/4 or 4/4, and occasionally 4/5 or 6/6; the

anterior premolars are usually shed before the posterior molars have erupted. Heterocephalus only has three molariform teeth in each ramus and on occasion this is reduced to two. In Cryptomys, M3 erupts early in life, and in Georychus M3 erupts late in life (De Graaff 1981). All cheekteeth are molariform in structure and differ only in size. All genera are strictly subterranean (see Table 15); individuals live and feed in an extensive network of burrows, the majority of which are superficial foraging burrows running at depths close to those of their food. Deeper burrows link foraging areas with nest, toilet area and, where present, a food store. Loose skin facilitates turning in the tight confines of the burrow. All genera, except Bathyergus, are chiseltooth diggers, biting at the soil face with their sharp rapidly growing incisors. All genera use fore- and hindfeet to push the loosened soil behind them; fringes of hairs on feet and tail help contain the soil as a mole-rat reverses down the burrow and up a side-branch where the soil is disposed of as a mound (or mounds) at the surface (Jarvis & Sale 1971). Except at the inception of mound formation, all genera, except Heterocephalus, have a plug of soil between the burrowing animal and the outside. Once the mound is fully formed, the sidebranch is packed with soil thereby sealing the burrow system from the surface. Mole-rats are most vulnerable to predation by snakes, birds and small carnivores during mound formation, the only time when an above-ground predator can accurately locate them. Activity patterns appear to vary seasonally, largely in response to temperature changes in superficial foraging burrows; mole-rats show poor ability to entrain their circadian rhythms to light. Peak burrowing activity, and mound formation, occur after rainfall when soil is easily worked. When the soil is very dry, excavated soil is packed into disused sections of the burrow system and not disposed of on the surface (Jarvis & Sale 1971, Jarvis & Bennett 1991). Mole-rats have little exposure to sunlight and exhibit signs of vitamin D3 deficiency. Nevertheless, unlike most mammals that need vitamin D3 for calcium uptake, mole-rats absorb calcium in the intestine and kidneys via specialized vitamin D3-independent paracellular processes (Buffenstein et al. 1994). The diet is high in cellulose and fibre, and is digested by symbiotic micro-organisms in a large caecum and hindgut. Digestive efficiency is high (>80%), facilitating maximum returns for foraging effort (Buffenstein & Yahav 1991a, 1994, Bennett & Jarvis 1995), and is further enhanced by re-ingesting partly digested faecal pellets.This autocoprophagy contributes to digestive efficiency, reinoculates the mole-rat with endosymbionts and provides an additional source of protein and energy from digestion of the microbes themselves. For their size, members of the Bathyergidae have lower than predicted metabolic rates (McNab 1966, Lovegrove 1986, Buffenstein & Yahav 1991b) and long maximum life-spans: >26 years for captive Heterocephalus glaber and >12 years for Cryptomys damarensis (O’Connor et al. 2002, Sherman & Jarvis 2002, J. U .M. Jarvis & N. C. Bennett unpubl.). Bathyergus, Heliophobius and Georychus are solitary, each individual aggressively defending its burrow system against conspecifics. All breed seasonally. Bathyergus and Georychus communicate through the soil by drumming with their hindfeet. Occupancy of a burrow by 641

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more than one animal occurs briefly at mating; young leave the maternal burrow at about two months; 1–2 litters are born during the breeding season (Jarvis & Bennett 1991, Sumbera et al. 2003a). In contrast, Heterocephalus and Cryptomys are social: colonies consist of family units usually with a single reproductive !, 1–3 consort "" and a number of non-reproductive (but not infertile) helpers (Jarvis 1981, Bennett 1988, Jarvis & Bennett 1991, Jarvis et al. 1994). New colonies of Cryptomys are formed from an out-bred pair and their offspring; the colony fragments on the death of the reproductive animals (Jarvis & Bennett 1993, Jarvis et al. 1994, Bishop et al. 2004, Burland et al. 2004). Heterocephalus is strongly xenophobic, frequently inbreeds, and replacement of reproductive animals often occurs from within the colony; occasionally out-breeding may occur (O’Riain et al. 1996, O’Riain & Braude 2001). Some Cryptomys breed seasonally (e.g. C. h. hottentotus) and others aseasonally (e.g. C. damarensis; Bennett 1988, Spinks et al. 1999). Breeding in Heterocephalus is aseasonal. Gestation is long: 56–111 days in Cryptomys and Heliophobius, and 66–74 days in Heterocephalus (Bennett et al. 1991). In seasonal breeders, a maximum of two litters are born annually, whereas aseasonal breeders have 3–4 litters each year. In all genera, testes lie abdominally or in inguinal pockets, and there is no development of a scrotum. Non-reproductive !! have small nipples and a vagina closure membrane. Litter-sizes are usually 2–6 young/litter, but reach 28 young/litter in Heterocephalus. Maximum recorded colony sizes are 41 animals for Cryptomys damarensis (Jarvis & Bennett 1993) and >300 for Heterocephalus (Brett 1991a, S. Braude unpubl.). Bathyergus dig extensive foraging burrows with their strongly clawed forefeet; other genera bite at the soil with large, rapidly growing, extra-buccal incisors. All feed on underground portions of plants, particularly roots, bulbs, corms and tubers located while digging extensive, superficial foraging burrows. Bathyergus and Georychus also eat aerial parts of plants by loosening the soil under the roots, and then pulling the entire plant into the burrow. Most genera store small food items in a chamber or blind-ending burrow situated close to the nest. Larger items are eaten in situ, often being partly eaten and then left to regenerate (Jarvis & Bennett, 1991). Molerats do not drink free water. Solitary genera are usually restricted to mesic habitats where food items are spaced close together and where rainfall is frequent allowing frequent opportunities to burrow. Sociality has enabled Cryptomys and Heterocephalus to also inhabit arid regions where food is widely dispersed and patchy and where rainfall is sparse and unpredictable, allowing for only limited opportunities to burrow (Jarvis & Bennett 1993, Jarvis et al. 1994). After rainfall, colonies can dig >1 km of foraging burrows in a month, rapidly expanding their home-range while the cost of digging is relatively low. Most of the food needed to sustain the colony until the next rains is located at this time. All species peel their food, holding onto small items of food with their forefeet while eating, with frequent pauses to shake it, or hold it, between their incisors and to brush it with the forefeet. All except Heterocephalus balance on the hindfeet while feeding on small items; Heterocephalus rests on the elbows. The Bathyergidae is a monophyletic group currently placed with the suborder Hystricomorpha, infraorder Hystrignathi. The Bathyergidae are the most species-rich of the four African families of the Hystricognatha, which also includes the families Hystricidae, Petromuridae and Thryonomyidae. Bathyergids show a number of unique features. All five genera have a highly flared angle of the lower

jaw, secondarily reduced infraorbital foramina, and unfused carpal bones. Few genera of other families of rodents have such a variable number of cheekteeth (3–6 in both upper and lower jaws) (Ellerman 1940, De Graaff 1981). Closest, but still distant, relatives are the other phiomorph families of the Hystricognathi: Rock Rats (Petromuridae), Cane Rats (Thryonomyidae) and Old World Porcupines (Hystricidae). Fossils from two extinct genera of bathyergids and of Heterocephalus date from the early Miocene (ca. 25 mya) in East Africa and Namibia. Molecular evidence also indicates early divergence of Heterocephalus and Heliophobius from extant members of the family (Allard & Honeycutt 1992, Faulkes et al. 1997b, 2004). Taxonomy of the Bathyergidae, particularly of Cryptomys, is under review. Cryptomys spp. from C Zambia and around Pretoria, South Africa, show much genetic divergence and several, as yet undescribed, species have been found. Little is known of the relationships of Cryptomys spp. from West Africa and Uganda. Additionally, genetically divergent Heliophobius have been found in W Tanzania and Malawi, and divergent Heterocephalus in Ethiopia (Faulkes et al. 1997b, 2004).The significance of these genetic divergences is unclear. The family is traditionally divided into two subfamilies, Bathyerginae and Georhychinae (De Graaff 1981, Allard & Honeycutt 1992) and currently 15 species are recognized (Table 45). The Bathyerginae has grooved upper incisors whose roots originate above the cheekteeth, enlarged forefeet and claws and large body size (up to ca. 2000 g), and contains one genus (Bathyergus) and two species. The Georhychinae has ungrooved upper incisors with roots originating behind the molars, forefeet and claws which are not enlarged, and a smaller body size (2 m deep) into which an animal retreats if alarmed, blocking the burrow behind it. Blind-ending toilet burrows occur also near the nest chamber, and food stores are sometimes present. Burrow configuration is constantly changing but animals remain in one area (Davies & Jarvis 1986). Burrows may overlap with those of Georychus capensis and/or Cryptomys h. hottentotus, the burrows of each species being at different

Social and Reproductive Behaviour Solitary and aggressively territorial. Cape Dune Mole-rats communicate through soil by drumming with both hindfeet simultaneously on the ground (two beats, pause, two beats, etc.; J. U. M. Jarvis unpubl.). They make snorting grunts and drum when threatened or alarmed. Vacant burrows are quickly taken over by neighbours. Little is known about reproductive behaviour, and there have been no successful matings in captivity. During courtship, "" and !! drum in unison, move soil and lock incisors; ! raises her tail and vocalizes while " follows and attempts to mount (J. U. M. Jarvis unpubl.). Marked sexual dimorphism and thick protective skin on the neck suggests "" fight for !!. Males seem to have more linear burrows than neighbouring !!, possibly providing access to several !! (Davies & Jarvis 1986). Mean home-range of adults at one site in coastal fynbos was 0.27 ha (0.14–0.35 ha) (Davies & Jarvis 1986); home-range can be much smaller elsewhere when densities are high. Reproduction and Population Structure Reproduction is seasonal, occurring during the wet winter (Apr–Nov), with pregnancies peaking in Aug. During the reproductive season, testes in "" change from abdominal to inguinal, but there is no real development of a scrotum. Usually one (occasionally two) litter/season. Gestation: 2 months (estimate). Litter-size: 3.3 (1–6), n = 99. At birth, young weigh 27–52 g (n = 10). Eyes open Day 7. First solid foods eaten ca. Day 15. Weaned ca. Day 30. Inter-sibling sparring begins at Day 12, and later escalates to fighting. Young disperse either above or below ground ca. Day 60–65, when weight is ca. 300 g. Sexual dimorphism beginning to be evident at one year of age, when weight is 420–638 g (n = 4). Smallest wild sexually active individuals at weight of 529 g ("") and 494 g (!!) (J. U. M. Jarvis unpubl.). Growth continues for several years. Longevity: >6 years. Sex ratio is parity (M. J. O’Riain & J. U. M. Jarvis unpubl.). Predators, Parasites and Diseases Despite their large size, even adult Cape Dune Mole-rats are eaten by Mole-snakes Pseudapsis cana and probably also by the Cape Cobra Naja nivea. Jackals, caracals, other small carnivores and even herons and some raptors, will capture animals working close to the surface or wandering above ground. During the winter, Cape Dune Mole-rats occupying low-lying areas such as temporary vleis, sometimes get flooded out of their burrow systems. They have little resistance to cold and are easy prey to these predators. Ectoparasites include three species of mites, one species of flea, one species of sucking louse and two species of ticks. Three species of endoparasitic worms have been recorded (De Graaff 1964b, 1981). Close to habitations, Cape Dune Mole-rats are often infested with the cysts of the dog tapeworm. 647

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Family BATHYERGIDAE

Conservation IUCN Category: Least Concern. Cape Dune Mole-rats may be pests.They undermine roads, damage earthen dam walls, chew through underground communication cables and irrigation pipes. Their large mounds damage combine harvesters. They are trapped extensively, and eaten by local people. Measurements Bathyergus suillus HB (""): 281 (240–330) mm, n = 39 HB (!!): 256 (204–300) mm, n = 45 T (""): 52 (30–70) mm, n = 28 T (!!): 46.6 (25–61) mm, n = 32 HF (""): 51.9 (45–65) mm, n = 37 HF (!!): 46.9 (42–55) mm, n = 43 E (""): 0 mm E (!!): 0 mm

WT (""): 896 (529–2200) g, n = 208 WT (!!): 670 (494–900) g, n = 257 GLS (""): 65.1 (56–76.7) mm, n = 27 GLS (!!): 55.1 (39.4–69.3) mm, n = 32 GWS (""): 40.1 (34.2–47.3) mm, n = 28 GWS (!!): 38.1 (28.5–41) mm, n = 32 P4–M3 (""): 11.4 (10.6–12.5) mm, n = 27 P4–M3 (!!): 10.9 (9.3–13) mm, n = 32 South Africa (TM, AM, MM) Weights: J. U. M. Jarvis unpubl. Key References Davies & Jarvis 1986; De Graaff 1981; Jarvis & Bennett 1991; Skinner & Smithers 1990. J. U. M. Jarvis

GENUS Cryptomys Mole-rats Cryptomys Gray, 1864. Proc. Zool. Soc. Lond. 1864: 124. Type species: Georychus holosericeus Wagner, 1842 (= Bathyergus hottentotus Lesson, 1826).

Cryptomys hottentotus.

The genus Cryptomys contains ten species distributed throughout western, central and southern Africa, but is absent from the Horn of Africa, tropical rainforests of central and West Africa, and the Sahara. It is distributed in a wide range of soil types – fine clays to coarse sand and occasionally brecciated soils – and occurs in a variety of biomes, from mesic to arid. These mole-rats are of intermediate size, being larger than Heterocephalus but smaller (except for C. mechowi) than Bathyergus, Georychus and Heliophobius. Pelage colour is cinnamon, fawn, grey and black amongst the different species.The muzzle is flat; tail is shorter than the hindfeet; the toes of fore- and hindfeet, and their claws, are short.The skull is less robust than in Bathyergus. Cheekteeth are simple folds in adults.The premaxilla bones, housing the incisors, do not bulge out laterally as much as in Bathyergus and Georychus. The jugal fits into an elongate groove on the outer upper side of the zygomata. Upper incisors, without grooves, are rooted in the pterygoid bones posterior to the cheekteeth (Figure 104). Cryptomys mole-rats are social, occurring in colonies with reproductive division of labour, overlap of generations, cooperative

Figure 104. Skull and mandible of Cryptomys hottentotus (BMNH 98.4.4.23).

care of young and high reproductive skew. All species studied to date are obligate outbreeders. In mesic areas, Cryptomys spp. are loosely social, whereas species in arid areas have well-developed social structures. Each colony is characterized by having a single reproductive ! and between one and two reproductive "". Non-reproductive animals exhibit socially induced infertility ranging from strict incest

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Cryptomys anselli

avoidance in mesic species through to physiological suppression in arid species. In the clade (see below) containing C. hottentotus, courtship and copulation is usually initiated by ", whereas in the C. damarensis clade it is initiated by !. All species are subterranean and chiseltooth diggers; they excavate extensive burrow systems by disposing of excavated soils as mounds on the surface Approximately 49 forms have been named (Ellerman 1940). Size, colour and external appearance are poor criteria for making comparisons between species in this genus. The genus Cryptomys is composed of two highly divergent clades, whose genetic distance can exceed that between some of the other genera in the family (Faulkes et al. 1997b), a division also supported by morphological characters of the skull. DNA sequence analysis has gone some way in clarifying the taxonomic problems of the genus (e.g. Allard & Honeycutt 1992, Faulkes et al. 1997b, Bennett & Faulkes 2000, Walton et al. 2000). The first clade contains all species except for C. hottentotus, has thickwalled infraorbital foramina and is distributed widely throughout savanna habitats in southern, central and eastern Africa, with two species with restricted distributions in West Africa. This clade is karyotypically very diverse (Burda et al. 1999), ranging from 2n = 40 in C. mechowi to 2n = 74 in C. damarensis. The second clade contains only C. hottentotus, has thin-walled elliptically shaped infraorbital foramina, and is karyotypically conserved (2n = 54). The smallest genetic distance between Cryptomys species is 8.4% for C. mechowi and C. bocagei. Cryptomys anselli from Zambia and C. darlingi from Zimbabwe have traditionally been classified as subspecies

of C. hottentotus, but are highly divergent from this species and are indeed distinct species. Within the C. hottentotus clade, C. h. nimrodi from Zimbabwe, C. h. natalensis from South Africa (KwaZulu–Natal) and C. h. pretoriae from South Africa (Gauteng) are also genetically divergent from one another and should possibly be considered as separate species. There are no data for the genetic relationships between C. foxi, C. zechi and C. ochraceocinereus; all three species have a small thick-walled infraorbital foramina, as in C. damarensis. These differences within Cryptomys have shown that the genus, as currently defined, may be composed of two genera. The genus Fukomys has been proposed (Kock et al. 2006) to contain all species (anselli, bocagei, damarensis, darlingi, foxi, kafuensis, mechowi, ochraceocinereus, zechi) except for hottentotus, which remains in the genus Cryptomys. Fukomys cannot be separated from Cryptomys on the grounds of morphological or morphometric characters. Fukomys is distinguished from Cryptomys by nuclear and mitochondrial DNA (Faulkes et al. 2004, Ingram et al. 2004) and high karyotypic diversity with diploid karyotypes ranging from 2n = 40 to 80, as opposed to a very conservative 2n = 54 in Cryptomys (van Daele et al. 2004, Deuve et al. 2008).The separation into two genera is supported by reciprocal monophyly of the two lineages based on nuclear and mitochondrial data sets and the level of sequence divergence observed between the two lineages for nuclear and mitochondrial DNA (e.g. Bathyergus and Georychus) (Faulkes et al. 1997b, Ingram et al. 2004). Nigel C. Bennett

Cryptomys anselli ANSELL’S MOLE-RAT Fr. Rat-taupe d’Ansell; Ger. Ansells Graumull Cryptomys anselli Burda, Zima, Scharff, Macholan and Kawalika, 1999. Z. Säugetierkunde 64: 36–50. Chainama Hills, Lusaka, Zambia.

Taxonomy Prior to 1999, this species was included in C.hottentotus (e.g. Ansell 1978). It was referred to as C. hottentotus or Cryptomys ‘population Lusaka, karyotype 2n = 68’ in papers published between 1987 and 1999. This species has been placed in the genus Fukomys by Kock et al. (2006) (see genus Cryptomys). Synonyms: none. Allozyme profile (Filippucci et al. 1994, 1997) and DNA sequences (Ingram et al. 2004) clearly separate this species from other species of Cryptomys. Chromosome number: 2n = 68, FN = 79–82 (Burda et al. 1999). Description Medium-sized mole-rat. Pelage colour is age- and weight-dependent: dark slate-grey (neonates), greyish-brown (weaned young), brown (juveniles and subadults) and golden-ochre (adult animals). Head with conspicuous white patch on forehead in most (but not in all) individuals. Tail very short (ca. 14% HB). Skull: infraorbital foramina elliptical, wide at base, thick-walled. Upper incisors ungrooved. Nipples: 2 + 1 = 6. Geographic Variation None recorded. Similar Species Cryptomys kafuensis. Similar in appearance, head spot white, usually larger; chromosome number 2n = 58.

Cryptomys anselli

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Family BATHYERGIDAE

Distribution Endemic to Africa. Zambezian Woodland BZ. Recorded only from several localities in C Zambia (Lusaka and its surroundings, to a maximum of ca. 100 km from Lusaka, but not across the Kafue R.).

follicular development; tertiary follicles luteinize or atrophy so that ovulation does not occur (Willingstorfer et al. 1998).

Habitat Savanna bushland, cultivated fields, gardens and golf courses where mean annual rainfall is ca. 820 mm.

Predators, Parasites and Diseases Apart from humans, no predators are known that have specialized on Ansell’s Mole-rats. Ectoparasites have not been found. Low infestation with a nematode (Protospirura muricola) has been reported (Scharff 1998).

Abundance Detailed assessment of abundance not available, but the species appears to be rather abundant in the Lusaka and Central Provinces.

Conservation IUCN Category: Near Threatened. Ansell’s Mole Rats are considered to be agricultural and horticultural pests; they are hunted by local people and are a highly valued food.

Adaptations Subterranean. Ansell’s Mole-rat has a low resting metabolic rate of 0.63 ± 0.06 ml O2/g/h (67% of expected). Has a low body temperature of 33.8 °C and a high thermal conductance of 0.12 ml O2/h/ °C (Bennett et al. 1994a).

Measurements Cryptomys anselli HB (""): 121.5 ± 10.9 (109–135) mm, n = 20 HB (!!): 119.3 ± 8.0 (108–132) mm, n = 30 T (""): 17.9 ± 1.9 (15.6–21.7) mm, n = 20 T (!!): 18.3 ± 2.3 (13.9–22.9) mm, n = 30 HF (""): 22.6 ± 3.1 (21.8–25.8) mm, n = 20 HF (!!): 23.3 ± 0.9 (21.8–25.2) mm, n = 30 E (""): 0 mm E (!!): 0 mm WT (""): 96.1 ± 14.7 (80–145) g, n = 40 WT (!!): 79.1 ± 12.7 (65–122) g, n = 100 GLS (""): 33.9 ± 1.9 (29.0–38.8) mm, n = 10 GLS (!!): 31.8 ± 1.7 (29.8–34.7) mm, n = 10 GWS (""): 26.5 ± 2.3 (22.6–30.0) mm, n = 10 GWS (!!): 23.9 ± 1.8 (22.1–26.1) mm, n = 10 P4–M3 (""): 6.1 ± 0.3 (5.6–6.9) mm, n = 10 P4–M3 (!!): 6.0 ± 0.4 (5.4–6.8) mm, n = 10 Zambia (H. Burda unpubl.)

Foraging and Food Herbivorous; feeds on rootstocks, tubers, bulbs, corms and rhizomes (Scharff 1998). Social and Reproductive Behaviour Social. Lives in family groups of about 12 animals (range 2–25, n = 12) (Scharff 1998). A colony consists of a founding reproductive pair and their offspring from several litters. These offspring do not reproduce due to incest avoidance based on individual recognition of the family members. Offspring are (in many cases probably lifelong) helpers at the nest (Burda 1995, Burda et al. 2000). Reproduction and Population Structure Reproduction is aseasonal and one or two (rarely three) litters can be produced each year in captivity. Gestation: 98 (84–112) days. Litter-size: 2 (1–5), n = 102 litters. Sex ratio of neonates: 1 : 1.4 (n = 159). Weight of neonates: 7.9 (5.7–10.7) g. Pelage first visible on Day 8–10. Eyes open at weight 12.9 g on Day 23 (13–50). Solids first eaten ca. Day 22. Weaned at weight 34 g, on ca. Day 82 (75–105) (Burda 1989, 1990, Begall & Burda 1998). Non-reproductive !! have reduced

Key References Burda 1989.

Begall & Burda 1998; Bennett et al. 1994a; Nigel C. Bennett & Hynek Burda

Cryptomys bocagei BOCAGE’S MOLE-RAT Fr. Rat-taupe de Bocage; Ger. Bocages Graumull Cryptomys bocagei (de Winton, 1897). Ann. Mag. Nat. Hist., ser. 6, 20: 323. Hanha, Angola.

Taxonomy Originally described in the genus Georychus. Included within C. hottentotus by De Graaff (1981) and Smithers (1983), but now considered as a valid species by Woods (1993) and Woods & Kirkpatrick (2005). Recognized as a distinct species by Honeycutt et al. (1991) based on the character of the infraorbital foramina.This species has been placed in the genus Fukomys by Kock et al. (2006) (see genus Cryptomys). Synonyms: kubangensis. Subspecies: none. Chromosome number: 2n = 58 (G. H. Aguilar unpubl.). Description Medium-sized mole-rat with white patch on head. Dorsal and ventral pelage drab-grey to silvery-grey. Head blunt, usually with variably shaped white patch on forehead; incisor teeth visible outside lips. Long vibrissae. Eyes very small. External ear absent. Limbs short, feet pink and naked. Tail very short (ca.

7% of HB). Skull broad, strong; zygomatic arches slightly bowed anteriorly; infraorbital foramina small, teardrop-shaped (1.5– 2 mm), thin-walled. Upper incisors ungrooved. Males slightly larger than !!. Nipples: 2 +1 = 6. Geographic Variation

None recorded.

Similar Species Cryptomys mechowi. On average much larger in size. Distribution Endemic to Africa. Zambezian Woodland BZ. Recorded in C and S Angola, N Namibia (particularly Ondjeva, Ongha and Ondongera in Ovamboland Province), extreme S DR Congo and W Zambia.

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Cryptomys damarensis

Habitat

No information.

Abundance No information. Probably common. Remarks Recorded to feed on geophytes (G. H. Aguilar unpubl.). Social behaviour: four animals were caught from one colony that was not completely trapped-out. Conservation

IUCN Category: Data Deficient.

Measurements Cryptomys bocagei HB (""): 151 (141–165) mm, n = 4 HB (!!): 155 (150–165) mm, n = 4 T (""): 10 (7–12) mm, n = 4 T (!!): 11 (6–15) mm, n = 4 HF (""): 20 (19–22) mm, n = 4 HF (!!): 22 (20–24) mm, n = 4 E (""): 0 mm E (!!): 0 mm WT (""): n. d. WT (!!): n. d. GLS (""): 31.9 (29.9–34.4) mm, n = 4 GLS (!!): 33.5 (32.5–35.0) mm, n = 4 GWS (""): 22.5 (21–24.6) mm, n = 4 GWS (!!): 23.6 (23.3–24.4) mm, n = 4 P4–M3 (""): 5.3 (5.0–5.7) mm, n = 4 P4–M3(!!): 5.1 (4.8–5.3) mm, n = 4

Cryptomys bocagei

Throughout geographic range (De Graaff 1964a) Key Reference De Graaff 1964a. Nigel C. Bennett

Cryptomys damarensis DAMARALAND MOLE-RAT Fr. Rat-taupe du Damara; Ger. Damaraland Graumull Cryptomys damarensis (Ogilby, 1838). Proc. Zool. Soc. Lond. 1838: 5. Damaraland, Namibia.

Taxonomy Originally described in the genus Bathyergus. Included within C. hottentotus by Ellerman et al. (1953), De Graaff (1981) and Smithers (1983), but now considered as a valid species by Woods (1993) and Woods & Kirkpatrick (2005). Described as a distinct species by Honeycutt et al. (1991) on characters of the infraorbital foramina. Faulkes et al. (1997b) confirmed specific status by showing that an 11% sequence divergence in 12S rRNA occurred between haplotypes of C. hottentotus and C. damarensis.This species has been placed in the genus Fukomys by Kock et al. (2006) (see genus Cryptomys). Synonyms: lugardi, micklemi, ovamboensis. Subspecies: none. Chromosome number: 2n = 74 or 2n = 78, aFN = 92 (Nevo et al. 1986). Description Medium-sized colonial mole-rat with white patch on dorsal region of head. Pelage short, thick, with a sheen. Dorsal and ventral pelage fawn, or very dark brown to black. In a single colony, a single colour morph, or both colour morphs, may be present. Middorsal and mid-ventral white stripes in some individuals. Isolated tactile hairs protrude from pelage, especially on the face. Head with large white patch on forehead, sometimes with flecks of pelage colour. Incisor teeth visible outside lips. Eye small. External ear

pinnae absent. Limbs short and feet pink and naked. Tail very short (ca. 18% of HB), with stiff bristles that radiate from tail. Skull: dorsoventrally flattened; sagittal crest well developed; zygomatic arch strongly bowed outwards; infraorbital foramina small, teardropshaped (1.5–2 mm), thin-walled; upper incisors ungrooved; angular process on mandible not extending far backwards. Nipples: 2 + 1 = 6. Geographic Variation Individuals from South Africa (Kalahari), Botswana (Okavango swamps) and Zimbabwe are black in colour. Those from C Namibia (Dordabis and Rheoboth) may be either black or fawn in colour. Similar Species C. darlingi. On average smaller; pelage fawn (similar to the fawncoloured morph of C. damarensis); chromosome number: 2n = 54; Zimbabwe and Mozambique. C. hottentotus. On average smaller; pelage grey; chromosome number: 2n = 54; Western Cape, Northern Cape, Eastern Cape, KwaZulu–Natal, Free State, Gauteng and Mpumalanga Provinces. Occurs in sympatry with C. hottentotus in Van Zyl’s Rus, Northern Cape Province. 651

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Foraging and Food Herbivorous. Superficial foraging burrows are excavated as the animals search for underground storage organs of geophytes (bulbs, tubers and rootstocks).There is selective storage of geophytes: geophytes that are proportionately larger may possibly have a longer ‘shelf life’, or it may be more cost-effective to transport large geophytes to the store than small ones. Geophytes include various genera of Hyanthaceae (e.g. Dipcadi, Ledebouria, Ornithogalum), Portulaceae (e.g. Talinum) and Cucurbitaceae (e.g. Acanthosicyos), a number of which are toxic to livestock but not to mole-rats. Portable bulbs and corms are stored whereas the very large tubers (e.g. Acanthosicyos) are partly eaten in situ, and will often regenerate (Bennett & Faulkes 2000, Jarvis et al. 2000).

Cryptomys damarensis

Distribution Endemic to Africa. Zambezian Woodland BZ and South-West Arid BZ (Kalahari Desert). Recorded from most of Botswana (except extreme east), NW South Africa, C and N Namibia, SW Zimbabwe and extremeW Zambia. Distribution closely associated with red Kalahari arenosols, but also occurs in coarse sandy soils. Habitat Semi-arid thorn scrub, woodland savanna and grasslands associated with red Kalahari sands and sandy soils; rainfall is typically low and sporadic (200–400 mm/annum) and burrow temperatures range from a mean of 30 °C in summer to 19 °C in winter (Bennett et al. 1988). Abundance Localized, but may be abundant in suitable habitats where population numbers can exceed 380 individuals/km2. However, abundance of subterranean mammals is not easy to estimate. Adaptations Subterranean. Activity is discontinuous throughout the day and night. Damaraland Mole-rats excavate extensive burrow systems that can extend for more than 1 km and appear to radiate from a central nest and nearby food store (Jarvis et al. 1998). The nest is deep, sometimes exceeding 2.4 m below the surface, and has two to three entrances. Toilet areas have not been found. Individuals within a colony remain resident in the same home-range for many years (>8 years; N. C. Bennett & J. U. M. Jarvis unpubl.). When threatened, a Damaraland Mole-rat has a unique defensive posture, rolling onto its back, exposing its belly, with the mouth agape and incisors bared. It braces itself with its limbs and rolls from side to side (Bennett 1990). Like other species of Cryptomys, it can also throw its head back with mouth agape and produce a threatening grunt. Damaraland Mole-rats have one of the lowest resting metabolic rates of all mammalian species: 0.66 ± 0.07 ml O2/g/h (85% of expected), a low body temperature (35.1 °C) and a low thermal conductance 0.065 ml O2/g/h/ °C (Bennett et al. 1992).

Social and Reproductive Behaviour Social. Lives in mediumsized colonies of around 12 animals (range 2–41) (Bennett & Jarvis 1988, Jarvis & Bennett 1993). The colony consists of a founding reproductive pair and their progeny from several litters; these younger individuals do not breed while in the natal colony. Sex ratios of captured colonies range from 0.8 to 2.1 in favour of "". Mean body mass of individuals in the colony may vary from 103 to 202 g in "" and 88 to 145 g in !! (n = 6 colonies, n = 107 individuals) depending upon the ages of the adult non-reproductive animals in the colony (Jacobs et al. 1990).The reproductive pair consists of the most dominant individuals; the non-reproductive "" are more dominant than non-reproductive !!. The non-reproductive members of the colony can be placed into work-related groups based on body mass: there is a tendency for smaller (not necessarily younger) animals to perform more burrow maintenance than larger (but not necessarily older) animals. An oestrus ! solicits the " prior to mating. Multiple mating occurs following a ritualized courtship of tail-to-tail chasing, vocalizations and head mounting by the reproductive ! (Bennett & Jarvis 1988). Reproduction and Population Structure In most "", the testes are abdominal and the penis is contained in a penile sheath; in a reproductive ", the penis is usually visible beyond the sheath and the testes are often in inguinal pockets. Females have external labial flaps and an os-clitoris, that it only exposed during sexual activity, and is the same length as the penis of the ". A vaginal closure membrane is present in non-reproductive !!. The reproductive ! can be identified by her open vagina and prominent nipples. Reproduction is aseasonal, with up to three litters per annum. Ovulation is spontaneous. Gestation: 78–92 days (Bennett & Jarvis 1988). Litter-size: 3 (1–6), n = 8 litters. At birth, young weigh 8–9 g and are mobile. Pelage develops from Day 6. Solid foods first eaten ca. Day 6–8. Eyes open on Day 18. Weaned Day 28. Intersibling sparring begins at Day 18–25.Young do not disperse but join the natal colony (Bennett et al. 1991). Non-reproductive "" have functional gonads but are oligospermic and occasionally azoospermic, although they have similar hormone profiles to the reproductive " (Maswanganye et al. 1999). In contrast, the non-reproductive !! do not ovulate and show reduced follicular development with tertiary follicles luteinizing or atresing. The hormone profiles of non-reproductive !! have lower concentrations of oestrogen, progesterone and LH than the reproductive ! (Bennett et al. 1993, 1994c). Reproductive suppression is due to two components: a suppressive action from the

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social environment, and incest avoidance in the form of obligatory out-breeding. Removal of a non-reproductive ! from the colony releases her suppression and her gonads become active. In a functional colony, the oldest non-reproductive animals are the primary dispersers; however, when either a reproductive " or ! dies, the entire colony will fragment. In either instance, dispersal occurs after rain when the costs of digging are lowest (Bennett et al. 1996). Predators, Parasites and Diseases The Damaraland Mole-rat is at particular risk from predation during mound formation. Molesnakes Pseudapsis cana appear to detect such freshly turned soil. They seize a mole-rat from behind, constrict and kill it. Cobras Naja nivea also enter the open holes of mole-rat burrows. Dispersing animals are particularly vulnerable to owls and small carnivores such as jackals, mongooses, Caracals Caracal caracal and occasionally Brown Hyaenas Hyaena brunnea. Damaraland Mole-rats have conspicuously few parasites; the predominant parasites are intestinal nematodes. Conservation IUCN Category: Least Concern. There is little conflict with agriculture (unlike some other species of mole-rats) because most of the habitat is arid.

Measurements Cryptomys damarensis HB (""): 164 (150–185) mm, n = 20 HB (!!): 151 (141–164) mm, n = 4 T (""): 25 (23–30) mm, n = 20 T (!!): 28 (25–32) mm, n = 4 HF ("") 27 (26–30) mm, n = 20 HF (!!): 27 (26–28) mm, n = 4 E (""): 0 mm E (!!): 0 mm WT (""): 161 (56–234) g, n = 17 WT (!!): 119 (49–206) g, n = 25 GLS (""): 36.2 (32.2–44.1) mm, n = 20 GLS (!!) 35.3 (31.6–38.2) mm, n = 4 GWS (""): 25 (23.3–33.0) mm, n = 20 GWS (!!): 27.1 (25.1–30.3) mm, n = 4 P4–M3 (""): 5.8 (5.2–6.7) mm, n = 20 P4–M3 (!!): 6.0 (5.5–6.4) mm, n = 4 South Africa and Namibia (De Graaff 1964a; Bennett et al. 1990) Key References Bennett & Jarvis 1988; Bennett et al. 1994c; Jarvis & Bennett 1993; Jarvis et al. 1998. Nigel C. Bennett

Cryptomys darlingi DARLING’S MOLE-RAT (MASHONA MOLE-RAT) Fr. Rat-taupe de Darling; Ger. Darlings Graumull Cryptomys darlingi (Thomas, 1895). Ann. Mag. Nat. Hist., ser. 6, 16: 239. Salisbury, Rhodesia (now Harare, Zimbabwe).

Taxonomy Originally described in the genus Georychus. Included within C. hottentotus by Ellerman et al. (1953), De Graaff (1981), Smithers (1983) and Woods (1993) but now considered as a valid species (Woods & Kirkpatrick 2005). This species has been placed in the genus Fukomys by Kock et al. (2006) (see genus Cryptomys). Synonyms: beirae, nimrodi, zimbitiensis. Subspecies: none. Chromosome number: 2n = 54, aFN = 80 (Aguilar 1993).

number: 2n = 74; South Africa, Namibia, Zimbabwe, Botswana and Zambia.

Description Medium-sized mole-rat with a longitudinal white stripe on ventral surface. Pelage short and thick. Dorsal and ventral pelage blackish, seal-brown, slate- or silvery-grey. White stripe or patch on ventral pelage; variable in width and length, may extend for length of body. Head blunt, sometimes with white patch on forehead; incisor teeth visible outside lips. Isolated tactile hairs protrude from pelage, especially on the face. Eye small. External ear absent. Foreand hindfeet naked with soft pink skin. Tail very short (ca. 7% of HB), naked with coarse vibrissae. Pelage of juveniles is darker than in adults. Skull broad, braincase large and rounded; infraorbital foramina small, teardrop (1.5–2 mm), thin-walled; upper incisors ungrooved. Nipples: 2 + 1 = 6. Geographic Variation

None recorded.

Similar Species Cryptomys damarensis. Fawn-coloured morph very similar; on average larger; pelage fawn, or very dark brown to black; chromosome

Cryptomys darlingi

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Distribution Endemic to Africa. Zambezian Woodland BZ. Recorded only in N and E Zimbabwe, extending into C Mozambique. Mainly restricted to the Mashonaland plateau at altitudes above 1000 m. Limits of geographic range not known. Habitat Miombo woodland predominated by Brachystegia and Julbernardia. Also found in valley grasslands, and on sandstone and granitic derived soils in areas of relatively high and predictable rainfall (ca. 700 mm/annum) (Genelly 1965, as C. hottentotus).

Reproduction and Population Structure Reproduction is aseasonal, with up to four litters per annum. Gestation (estimated): 56–61 days. Litter-size: 2 (1–3), n = 7 litters. Sex ratio at birth is parity. At birth, young weigh 6.9–8.2 g (n = 4). Pelage first appears Day 4. First solid foods eaten Day 14. Eyes open Day 14. Weaned ca. Day 45. Non-breeding "" are smaller than breeding "", the penis is enclosed by a sheath, and they lack prominent bulging inguinal testes. Non-reproductive !! have an imperforate vagina. Predators, Parasites and Diseases No information.

Abundance Common in miombo woodlands and grasslands. Conservation Adaptations Subterranean. Mashona Mole-rats have a low resting metabolic rate of 0.98 ± 0.14 ml O2/g/h (97% of expected) and a low body temperature of 33.3 ± 0.5 °C. They have strong poikilothermic tendencies in body temperature below ambient temperatures of 25 °C, whereas above 25 °C they are endothermic (Bennett et al 1993a). Foraging and Food Herbivorous. These Mole-rats sometimes burrow around the root systems of Brachystegia and Julbernardia trees, where they feed on geophytes and on the swollen roots of these trees (N. C. Bennett unpubl.). Social and Reproductive Behaviour Social. Mashona Molerats live in small colonies of 5–9 individuals consisting of a founding reproductive pair and the offspring from several litters, who remain non-reproductive while resident in the natal burrow. Sex ratios of all individuals from captured colonies appear to be biased towards "". Mean body mass of all individuals in a colony varies from 52 to 75 g (n = 5 colonies), this depending upon number of adults in the colony. The reproductive animals are the most dominant, thereafter "" are more dominant than !! (Gabathuler et al. 1996). Nonreproductive members of the colony cannot be placed into clearly defined work-related groups based on body mass (cf. C. damarensis). When in oestrus, ! solicits " prior to mating. Multiple copulations occur over two days (Bennett et al. 1994b). Reproductive inhibition in the non-breeding animals appears to be maintained by incest avoidance alone (Bennett et al. 1994b).

IUCN Category: Least Concern.

Measurements Cryptomys darlingi HB (""): 145 (125–165) mm, n = 38 HB (!!): 141 (135–150) mm, n = 18 T (""): 10 (8–13) mm, n = 38 T (!!): 10 (10–10) mm, n = 18 HF (""): 23 (21–30) mm, n = 38 HF (!!): 22 (20–24) mm, n = 18 E (""): 0 mm E (!!): 0 mm WT (""): 76.4 (60–88) g, n = 11 WT (!!): 77.0 (54–92) g, n = 7 GLS (""): 33.3 (30.6–37.9) mm, n = 38 GLS (!!): 32.6 (31–36.5) mm, n = 18 GWS (""): 24.4 (20.8–28.2) mm, n = 38 GWS (!!): 23.3 (21.3–27.2) mm, n = 18 P4–M3 (""): 5.3 (4.7–5.6) mm, n = 38 P4–M3 (!!): 5.1 (4.7–5.5) mm, n = 18 Harare and Goromonzi, Zimbabwe (De Graaff 1964a, N. C. Bennett unpubl.) Key References Bennett et al. 1994a; Gabathuler et al. 1996; Genelly 1965. Nigel C. Bennett

Cryptomys foxi FOX’S MOLE-RAT Fr. Rat-taupe de Fox; Ger. Foxs Graumull Cryptomys foxi (Thomas, 1911). Ann. Mag. Nat. Hist., ser. 8, 7: 462. Panyam, Jos Plateau, Nigeria.

Taxonomy Originally described in the genus Georychus. De Graaff (1975) placed foxi as a subspecies of C. ochraceocinereus, an arrangement followed by Happold (1987). Currently considered as a valid species. Detailed analysis of skull characters provided by Williams et al. (1983). This species has been placed in the genus Fukomys by Kock et al. (2006) (see genus Cryptomys). Synonyms: none. Chromosome number: 2n = 66, FN = 122 (n = 6), 2n = 70, FN = 130 (Cameroon; Williams et al. 1983). See Rosevear (1969) for further information. Description Sepia-coloured mole-rat, sometimes with whitish markings on head or body. Pelage soft and short, with velvet-like

texture. Dorsal and ventral pelage sepia-brown. Head blunt, usually with sub-circular white patch on forehead; incisor teeth visible outside lips. Long vibrissae. Eyes very small. External ear absent. Limbs short and feet naked. Fore- and hindfeet well-developed. Five digits on fore- and hindfeet. Tail very short (ca. 8% of HB), covered with stiff bristles. Skull: widest part of nasal bones anteriorly; upper incisors ungrooved and comparatively narrow (ca. 2.4 mm); infraorbital foramina teardrop-shaped (1.5–2 mm), thin-walled; upper cheekteeth comparatively sh ort (mean 7.6 [range 7.2– 8.3] mm). Individuals from Cameroon are larger than those from Nigeria (see Measurements). Nipples: 2 + 1 = 6.

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Geographic Variation None recorded. Similar Species C. zechi. Pelage pale cinnamon; on average larger; widest part of nasal bones posteriorly; upper cheekteeth longer (ca. 8.3 mm); incisor teeth comparatively wider (ca. 3.2 mm); Ghana and Togo only. Distribution Endemic to Africa. Afromontane–Afroalpine BZ. Recorded only from near Panyam (1220 m) on the Jos Plateau, N Nigeria (Thomas 1911b, Rosevear 1969), and Ngaoundere, Cameroon (Williams et al. 1983). Occurs only at altitudes above 1000 m. Habitat On the Jos Plateau, Nigeria (ca. 1000–1800 m), typical habitats are extensive grasslands, rocky areas and riverine forest along streams (Happold 1987). Abundance Uncertain; very localized and probably uncommon. Remarks Virtually nothing is known about this species. Subterranean, living in colonies. Twelve animals were trapped from one colony but it is not known whether these animals constituted the whole colony. The Rev. G. T. Fox, who collected the holotype, recorded that these mole-rats ate earthworms in captivity (Thomas 1911b), an unusual situation for members of a genus that is primarily vegetarian. Conservation

IUCN Category: Data Deficient.

Measurements Cryptomys foxi HB: 145 (135–159) mm, n = 10 T: 14 (11–17) mm, n = 10 HF: 29 (26–31) mm, n = 10 E: 0 mm WT: n. d. GLS: 40.7 (39.3–43.0) mm, n = 10 GWS: 28.2 (25.7–30.0) mm, n = 10 P4–M3: 7.7 (7.2–8.3) mm, n = 10 Nigeria (Rosevear 1969)

Cryptomys foxi

T (""): 20.1 (16–23) mm, n = 7 T (!!): 20.5 (17–25) mm, n = 10 HF (""): 32.4 (30–36) mm, n = 7 HF (!!): 31.2 (29–34) mm, n = 10 E (""): 0 mm E (!!): 0 mm WT (""): n. d. WT (!!): n. d. GLS (""): 44.8 (42.0–48.4) mm, n = 7 GLS (!!): 42.7 (39.7–46.4) mm, n = 10 GWS (""): 31.0 (28.9–33.1) mm, n = 7 GWS (!!): 29.8 (28.7–32.8) mm, n = 10 P4–M3 (""): 7.5 (6.8–8.4) mm, n = 7 P4–M3 (!!): 7.3 (6.7–8.0) mm, n = 10 Cameroon (Williams et al. 1983) Key References

Thomas 1911b; Williams et al. 1983.

HB (""): 176.6 (161–191) mm, n = 7 HB (!!): 174.8 (162–191) mm, n = 10

Nigel C. Bennett

Cryptomys hottentotus COMMON MOLE-RAT (HOTTENTOT MOLE-RAT) Fr. Rat-taupe hottentot; Ger. Hottentotten-Graumull Cryptomys hottentotus (Lesson, 1826). Voyage Monde Coquille, Zool. 1: 166. Paarl, Cape Province, South Africa.

Taxonomy Originally described in the genus Bathyergus. Cryptomys hottentotus is a geographically variable polytypic species (hence the many synonyms), and is in need of revision. Meester et al. (1986) refer to five subspecies (each with many synonyms): hottentotus (17 synonyms), damarensis (3), darlingi (3), bocagei (1) and natalensis (16). Corbet & Hill (1991) recognize natalensis as a valid species without comment. The taxa damarensis, darlingi and bocagei are currently regarded as three valid species and are placed in the same clade as C. hottentotus (see genus

profile), and natalensis is placed as a synonym of C. hottentotus. Cryptomys hottentotus is placed within its own clade within the genus (see genus profile). All the available evidence indicates that the C. hottentotus clade is chromosomally conserved (2n = 54), and is distributed throughout South Africa, extending into Mozambique and parts of Zimbabwe. Mitochondrial DNA sequence analysis consistently resolves six groups within the C. hottentotus clade as follows: natalensis (KwaZulu–Natal and Mpumalanga Provinces), pretoriae (Gauteng, NorthWest and Limpopo 655

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Provinces), nimrodi (SW Zimbabwe), hottentotus (Western Cape, Northern Cape and Eastern Cape Provinces), mahali (Gauteng Province) and aberrans (Eastern Cape Province) (Faulkes et al. unpubl.). The first four forms are considered as subspecies (see below), but the status of mahali and aberrans is uncertain (Faulkes et al. unpubl.) Synonyms: abberrans, albus, amatus, arenius, beirae, bigalkei, caecutiens, cradockensis, darlingi, exenticus, holosericius, jamesoni, jorisseni, kopmotiensis, langi, lugwigii, mahali, melanoticus, montanus, natalensis, nemo, nimrodi, orangiae, pretoriae, rufulus, stellatus, talpoides, transvaalensis, valschensis, vandami, vrybergensis, whytei, zimbitiensis, zuluensis. Subspecies: four. Chromosome number (C. h. hottentotus): 2n = 54, FN = 106; (C. h. natalensis): 2n = 54, FN = 104 (Nevo et al. 1986). Description Medium-sized mole-rat with flattened body. Pelage short, thick and silky. Dorsal pelage cinnamon-buff, fawn or dark grey but with considerable geographical variation. Ventral pelage paler. Flanks may or may not be paler than dorsal pelage. Head similar in colour to body, with small white patch on forehead in some individuals. Isolated tactile hairs protrude through the pelage, more numerous on face than on body. Limbs short. Feet naked and pink.Tail very short (ca. 15% of HB), pink, with hairs fringing from the tail itself. Body size is variable, depending on habitat, status within the colony and reproductive state. Skull with strong convex curvature dorsally; sagittal crest poorly developed; infraorbital foramina elliptical (ca. 3 mm), thin-walled; upper incisors ungrooved; four cheekteeth without re-entrant folds and decreasing in size from P to M3. No (or very little) sexual dimorphism; sexes difficult to distinguish. Nipples: 2 + 1 = 6. Geographic Variation C. h. hottentotus: Western, Eastern and Northern Cape Provinces and Free State of South Africa; 2n = 54; pelage cinnamon-buff to fawn with or without head-patch. C. h. natalensis: KwaZulu–Natal and Mpumalanga Provinces of South Africa; 2n = 54; pelage dark grey to black with characteristic black colouration around snout (usually without distinct head-patch). C. h pretoriae: Gauteng, North West, Limpopo and Mpumalanga Provinces of South Africa; 2n = 54; pelage silvery-fawn to grey, with or without head-patch. C. h. nimrodi: Limpopo Province, South Africa, and S Zimbabwe; pelage silvery-fawn to cinnamon-buff, with or without head-patch.

BZ, and parts of Coastal Forest Mosaic BZ). Recorded from South Africa, Lesotho, Swaziland and S Zimbabwe. In South Africa, known from Stellenbosch and Somerset West on the Cape Peninsula, northwest to Steinkopf and inland to Prieska and Calvinia (Northern Cape Province); also in consolidated sands of the east coast and throughout Eastern Cape Province and southern parts of Free State; and in grasslands and savanna woodlands of KwaZulu–Natal, Mpumalanga, North-West and Gauteng Provinces in a range of soils from granitic sandstones to coarse clays. In Zimbabwe, found in granitic sandstones and in brecciated soils. Habitat Fynbos, grassland, savanna and Succulent and Nama Karoo with rainfall of 200–1000 mm per annum. Occurs in a range of substrates from friable sandy loams to exfoliated schists and sandy soils. Does not occur in heavy clay or very brecciated soils. Abundance Uncertain. In preferred habitats, density may exceed 150/km2.Abundance of subterranean mammals is not easy to estimate. Adaptations Subterranean. Capable of running forwards and backwards with equal ease in the burrow (Bennett 1992). Common Mole-rats have a low resting metabolic rate 0.92 ± 0.1 ml O2/g/h (90% of expected), low body temperature of 34.4 °C and high thermal conductance of 0.14 ml O2/g/h/ °C (Bennett et al. 1992). Body size varies with environment: in arid habitats, the mean body size (mean weight ± 1 SE) is much lower (57.8 ± 0.7 g; n = 722) than in mesic habitats (75.2 ± 1.0 g; n = 865) (Spinks 1998, Spinks et al. 2000). This reduction in body size in arid environments may be an adaptation to burrowing and to dispersed food, which requires less soil to be excavated per metre of burrow (Bennett et al. 1992). Foraging and Food Herbivorous. Common Mole-rats specialize on bulbs, corms and tubers, especially those of Albuca, Lachenalia,

Similar Species Bathyergus suillus. HB much larger; pelage without sheen; skull much flatter with large sagittal and occipital ridges; forefeet with elongated claws; upper incisors grooved. B. janetta. HB larger; dorsal pelage silvery-fawn with sheen; skull with smaller sagittal and occipital ridges; auditory bullae markedly swollen; palate extends posteriorly to level of M3; forefeet with elongated claws; upper incisors grooved. Georychus capensis. On average larger; dorsal pelage russet; head blackish with white muzzle and large white patch around ear opening; forefeet without elongated claws; upper incisors ungrooved. Distribution Endemic to Africa. Widespread in several BZs (South-West Cape BZ, South-West Arid Zone BZ [Karoo], Highveld

Cryptomys hottentotus

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Ornithogalum (Hyacinthaceae), Romulea, Micranthus, Homeria (Iridaceae) and Oxalis (Oxalidaceae). They select the larger-sized geophytes and place them in a store situated close to the nest. The stored geophytes are packed in soil and disbudded when they shoot. In areas with seasonal flooding, the store and nest are often sited in raised areas. In arid areas, where some geophytes are too large to carry to the store, they are left growing and hollowed out in situ; these partly eaten geophytes can often regenerate. Small quantities of above-ground vegetation may also be consumed. In mesic parts of their range, where food occurs in closely spaced clumps and rainfall is frequent, mole-rats are able to search for new resources for many months of the year. In arid regions, food is more dispersed and there are few opportunities to greatly extend the foraging burrows. A number of geophytes containing cardiac glycosides that are extremely toxic to livestock (e.g. Ornithogalum spp., Homeria spp., Morea spp.) are eaten by Common Mole-rats (Davies & Jarvis, 1986). Social and Reproductive Behaviour Social. Lives in small colonies of around five animals (2–14, n = 109). A colony consists of a founding reproductive pair and number of non-reproductive offspring. Sex ratios of captured colonies (n = 4) ranged from 0.5 to 1.6. Sex ratio of a population is 1.28 : 1 in favour of "" (n = 1053). Reproductive animals are the dominant animals in the colony; of the non-reproductive animals, "" are more dominant than !!. The non-reproductive members of the colony cannot be placed into clearly defined workrelated groups based on body mass; there is, however, a tendency for smaller (not necessarily younger) animals to perform more burrow maintenance than their larger (not necessarily older) counterparts. Sexual differences are minimal and Common Mole-rats are difficult to sex. In "", the testes are abdominal and the penis is retracted into a penile sheath. A vaginal closure membrane is present in all !! that are not actively reproducing. In the reproductive !, the vagina is open, and axillary and inguinal nipples are prominent. The gonads of nonreproductive "" are active and incest avoidance prevents the nonreproductive " offspring from breeding. The reproductive " solicits the ! prior to mating. Multiple copulations occur between the pair following ritualized courtship by the ", who grasps the hind region of the ! with his incisors, sometimes also urinating on her head (Bennett 1989, Spinks 1998, Malherbe et al. 2003). Common Mole-rats are strictly obligate outbreeders. A colony will remain resident in the same area for a number of years, and will aggressively defend its home-range against invasion by other colonies. In the more mesic parts of their distribution, there is some mixing of animals from different colonies, whereas in the arid parts, mixing rarely occurs (Bishop et al. 2004). Dispersal in the Common Mole-rat occurs more frequently in "" than in !!. Reproduction and Population Structure Reproduction is seasonal, occurring in summer (Oct–Jan), with up to two litters per annum. Litter-size: 3 (1–6), n = 6 (Bennett 1989, Bennett & Faulkes 2000). However, there is no regression of gonads in winter, perhaps because this is when adult non-reproductive animals can disperse and find mates. Elevated levels of reproductive hormones (testosterone, progesterone and oestrogen) at this time would promote bonding, and prepare a new reproductive pair for breeding during the following summer. Ovulation is induced (Spinks et al.

1999). Gestation: 59–66 days (Bennett 1989, Malherbe et al. 2003). At birth, young weigh 8–9 g and are altricial. Pelage developed from Day 8. First solid foods eaten ca. Day 10. Eyes open on Day 13. Weaned at Day 28. Inter-sibling sparring begins on Day 10–14. Young do not disperse but remain in the natal colony (Bennett 1989). Predators, Parasites and Diseases Common Mole-rats are at particular risk to predation during mound formation. They are eaten by a wide range of predators, including Mole-snakes Pseudapsis cana, Shield-nosed Snakes Aspidelaps scutatus, Cobras Naja naja, Barn Owls Tyto alba, Marsh Owls Asio capensis, Grey Herons Ardea cinerea and a number of small carnivores. Ectoparasites include eight species of fleas (one of which, Cryptopsylla ingrami, is specific to this mole-rat), one species of tick, one species of louse and ten species of mites (De Graaff 1981). Conservation IUCN Category: Least Concern. In parts of its range, Common Mole-rats are agricultural pests. Measurements Cryptomys hottentotus hottentotus HB (""): 122 (90–190) mm, n = 70 HB (!!): 118 (100–160) mm, n = 62 T (""): 17.4 (8–27) mm, n = 69 T (!!): 18 (10–25) mm, n = 54 HF (""): 21.3 (18–33) mm, n = 58 HF (!!): 21.5 (17–25) mm, n = 48 E (""): 0 mm E (!!): 0 mm WT (""): 65.8 (56–79) g, n = 19 WT (!!): 46.8 (41–54) g, n = 19 GLS (""): 30.6 (29–38.6) mm, n = 44 GLS (!!): 30.6 (27.2–34.3) mm, n = 42 GWS (""): 22.0 (19.6–28.0) mm, n = 47 GWS (!!): 21.9 (18.4–25.3) mm, n = 43 P4–M3 (""): 5.1 (4.5–6.1) mm, n = 39 P4–M3 (!!): 5.1 (4.6–5.9) mm, n = 36 Cape Province, South Africa Body measurements: De Graaff (1964a) Weights: Bennett (1989) C. h. pretoriae HB (""): 143.4 (103–172) mm, n = 19 HB (!!): 139.9 (123–165) mm, n = 20 T (""): 13.6 (10–15) mm, n = 19 T (!!): 13.8 (10–17) mm, n = 20 HF (""): 25.6 (19–27) mm, n = 19 HF (!!): 25.8 (23–29) mm, n = 20 E (""): 0 mm E (!!): 0 mm WT (""): 87.9 (59–148) g, n = 19 WT (!!): 80.6 (51–135) g, n = 20 GWS (""): 25.4 (19–31.6) mm, n = 71 GWS (!!): 24.1 (18.6–29.3) mm, n = 123 GLS (""): 38.7 (31.2–44.4) mm, n = 7 GLS (!!): 37.1 (29.2–41.2) mm, n = 123 657

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Family BATHYERGIDAE

P4–M3 (""): 6.7 (5.1–7.7) mm, n = 71 P4–M3 (!!): 6.6 (4.7–7.6) mm, n = 123 Gauteng Province, South Africa (Van Rensburg 2000)

Key References Bennett 1989; Davies & Jarvis 1986; Spinks et al. 1999, 2000. Nigel C. Bennett

Cryptomys kafuensis KAFUE MOLE-RAT Fr. Rat-taupe du Kafue; Ger. Kafue Graumull Cryptomys kafuensis Burda, Zima, Scharff, Macholan and Kawalika, 1999. Z. Säugetierkunde 64: 36–50. ‘Hot Springs’ in Itezhi-Tezhi, Kafue National Park, Zambia.

Taxonomy Prior to 1999, this species was included in C. hottentotus (e.g. Ansell 1978). Allozyme profile (Filippucci et al. 1994, 1997), chromosome number (Burda et al. 1999) and DNA sequences (Ingram et al. 2004) clearly separate this species from other species of Cryptomys. This species has been placed in the genus Fukomys by Kock et al. (2006) (see genus Cryptomys). Synonyms: none. Chromosome number: 2n = 58, FN = 82 (Burda et al. 1999). Description Medium-sized mole-rat. Pelage dark slate-grey (young) to golden-ochre (adults). Head blunt with conspicuous white patch on forehead (in most but not in all individuals), variable in shape. Tail very short (15% of HB). Skull: infraorbital foramina elliptical, thin-walled; upper incisors ungrooved. Nipples: 2 + 1 = 6. Geographic Variation None recorded. Similar Species C. anselli. Similar in appearance; white patch on the head tends to be smaller; chromosome number: 2n = 68.

HB (!!): 104 (96–115) mm, n = 5 T (""): 16.6 (14.3–19.0) mm, n = 4 T (!!): 16.7 (13.5–20.0) mm, n = 5 HF (""): 23.1 (19.9–25.0) mm, n = 4 HF (!!): 23.0 (18.9–24.3) mm, n = 5 E (""): 0 mm E (!!): 0 mm WT (""): 95 ± 32 g, n = 6 WT (!!): 75 (61–93) g, n = 6 GLS (""): 33.5 (31.5–35.6) mm, n = 5 GLS (!!): 32.3 (30.3–34.2) mm, n = 5 GWS (""): 23.7 (21.9–26.0) mm, n = 5 GWS (!!): 22.0 (20.6–23.9) mm, n = 5 P4–M3 (""): 6.0 (5.6–6.8) mm, n = 5 P4–M3 (!!): 5.9 (5.5–6.2) mm, n = 5 Zambia (H. Burda unpubl.) Key Reference Burda et al. 1999. Nigel C. Bennett & Hynek Burda

Distribution Endemic to Africa Zambezian Woodland BZ. Recorded only in Itezhi-Tezhi, Southern Province, Zambia. (Degree square of 1526C of Ansell 1978.) Habitat Grasslands and cultivated fields near villages where mean annual rainfall is 787 mm. Abundance Abundant in this restricted area. Remarks Little is recorded about the biology of this species. Subterranean and social. In many aspects of ecology, reproductive biology and behaviour apparently similar to C. anselli. Apart from humans (mole-rats are considered agricultural pests and are hunted for food), no predators are known to specialize on Kafue Mole-rats. Ectoparasites have not been found on this species. Endoparasites include two species of cestodes (Inermicapsifer madagascariensis and an undetermined species) and a nematode (Protospirura muricola); in one study, proportion of individuals with endoparasites was low (three out of 18) (Scharff et al. 1997). Conservation

IUCN Category: Vulnerable.

Measurements Cryptomys kafuensis HB (""): 112 (105–129) mm, n = 4

Cryptomys kafuensis

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Cryptomys mechowi

Cryptomys mechowi GIANT MOLE-RAT Fr. Rat-taupe géant; Ger. Riesiger Graumull Cryptomys mechowi (Peters, 1881). Senckenberg. Ges. Natuurw. Freunde Berlin, p. 133. Malanje, North Angola.

Taxonomy Originally described in the genus Georychus. Recently placed in the genus Fukomys by Kock et al. (2006) (see genus Cryptomys). Synonyms: ansorgi, blainei, mellandi. Subspecies: none. Chromosome number: 2n = 40, FN = 80 (Macholan et al. 1993). Description Large, stout mole-rat, the heaviest of all species in the genus (but not as large and heavy as Bathyergus spp.). Pelage short and dense, almost woolly in texture. Pelage colour age- and weightdependent: dark slate-grey (neonates), greyish-brown (weaned young), brown (juveniles and subadults) and golden-ochre (adult animals); hairs pale brown at base, variable as dorsal pelage at tip.Ventral pelage pale brown. Head large, without white spot on forehead (except very small spot in some individuals). Eyes small. No external ears (small ears hidden in pelage in specimens from Mt Moko). Mouth rustystained (some individuals). Fore- and hindfeet large and broad, naked with fringe of whitish bristles on outer border of each foot. Tail very short (up to 15 mm); longish white bristles, which extend beyond end of tail. Males on average larger than !!. Skull: robust and dorsoventrally flattened; infraorbital foramina elliptical (ca. 5 mm), thin-walled and slender; upper incisors ungrooved, large and broad. Nipples: 2 + 1 = 6. Cryptomys mechowi

Geographic Variation Individuals from Mt Moko, Angola are larger than those from elsewhere (see below). Similar Species Several sympatric species; none are usually as large or weigh as much as C. mechowi. Distribution Endemic to Africa. Zambezian BZ and Southern Rainforest–Savanna Mosaic. Recorded from N Zambia, S and E DR Congo, E Angola (and perhaps N Malawi). Habitat Savanna bushland, cultivated and abandoned fields, gardens, dambos (temporary swamps), pine plantations and dense Acacia woodland. Found in a variety of soil types from quite stony to pure sand and clay. The area of distribution is characterized by an annual rainfall of more than 1100 mm (Scharff et al. 2001a). Abundance Detailed assessment of abundance not available, but appears to be rather abundant in the Copperbelt Province of Zambia. Adaptations Subterranean. Extensive burrow system comprises a deep nest (60–160 cm deep) with three or four entrances, food stores and toilet areas (Scharff et al. 2001a). Giant Mole-rats have a low resting metabolic rate 0.6 ± 0.08 ml O2/g/h (96% of expected), low body temperature of 34 ± 0.4 °C, and a low thermal conductance 0.09 ± 0.01 ml O2/g/h/ °C (Bennett et al. 1994a) Foraging and Food Predominantly herbivorous. Food in noncultivated areas includes grass rhizomes, roots, bulbs and tubers of diverse weeds, shrubs and trees; in cultivated areas, they probably

feed on crop plants such as sweet potatoes, cassava and groundnuts (Scharff et al. 2001a). Giant Mole-rats are unusual amongst bathyergids, because they apparently supplement their diet with invertebrate and vertebrate commensals found in their burrows (Burda & Kawalika 1993, Scharff et al. 2001a). Social and Reproductive Behaviour Social. Lives in colonies of 2–20+ (probably up to 40 or more) individuals (Burda & Kawalika 1993, Scharff et al. 2001a). Sex ratio within colonies is in favour of !! (Scharff et al. 2001a). The colony consists of a founding reproductive pair and non-reproductive offspring from several litters (Burda & Kawalika 1993, Wallace & Bennett 1998, Scharff et al. 2001a). The reproductive animals are the most dominant, and the non-reproductive "" are more dominant than !!. The nonreproductive members of the colony cannot be placed into clearly defined work-related groups based on body mass (Wallace & Bennett 1998). Giant Mole-rats are very vocal compared to other species of Cryptomys (Burda & Kawalika 1993, Credner et al. 1997). Reproduction and Population Structure Giant Mole-rats breed aseasonally both in the laboratory and in the field, producing up to three litters per annum (Burda & Kawalika 1993, Bennett & Aguilar 1995, Scharff et al. 1999). Gestation: 112 (89–118) days. Litter-size: 2.6 (1–5), n = 41. Sex ratio (mean of 41 litters) at birth biased in favour of !! (1 : 1.9). At birth, young weigh 19.6 (12.6– 27.7) g. Thin pelage appears within the first week. Eyes open on Day 6. First solid foods eaten ca. Day 14.Weaned ca. Day 90. Inter-sibling sparring begins at Day 10 (Bennett & Aguilar 1995, Scharff et al. 659

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1999). Growth is comparatively slow: weight increases in an almost linear fashion until ca. 120 g at age ca. Day 170; thereafter increase in weight continues until ca. 250–300 g at ca. Day 450 (n = 1; Scharff et al. 1999). A colony consists of a founding reproductive pair and several litters of their offspring. In the laboratory, genetic studies have shown that female offspring may sometimes mate with the reproductive " (M. J. O’Riain & J. Bishop unpubl.). Offspring can be denoted as (in many cases probably lifelong) helpers at the nest (Burda & Kawalika 1993, Bennett & Aguilar 1995, Scharff et al. 1999, 2001a, Burda et al. 2000). Predators, Parasites and Diseases Giant Mole-rats are eaten by Mole-snakes and other large snakes, and also by humans. Ectoparasites have not been found on animals or in the nests of Giant Molerats. Endoparasites include three species of cestodes (Inermicapsifer madagascariensis, Raillietina sp., Inermicapsifer madagascariensis) and two species of nematodes (Protospirura muricola, Cappilaria sp.). Proportion of individuals with endoparasites is relatively low (12 out of 35) compared to most other rodents (Scharff et al. 1997, Scharff et al. 2001a). Conservation IUCN Category: Least Concern. Giant Mole-rats are detrimental to agriculture and horticulture and are considered as pests. They are an important source of protein for local people and their meat is highly valued in some areas of Zambia. Measurements Cryptomys mechowi HB (""): 190 ± 22 (156–262) mm, n = 10

HB (!!): 165 ± 18 (135–205) mm, n = 10 T (""): 27.3 ± 2.3 (23–31) mm, n = 10 T (!!): 27.8 ± 3.8 (23–33.7) mm. n = 10 HF (""): 35.3 ± 2.0 (30.6–37.8) mm, n = 10 HF (!!): 32.2 ± 1.0 (31–34) mm, n = 10 E (""): 0 mm E (!!): 0 mm WT (""): 370 ± 94 (250–560) g, n = 20 WT (!!): 245 ± 34 (200–295) g, n = 22 GLS (""): 52.0 (45.6–59.2) mm, n = 5 GLS (!!): 42.2 (34–49.7) mm, n = 5 GWS (""): 46.7 (40.3–53.2) mm, n = 5 GWS (!!): 33.2 (28.6–37.0) mm, n = 5 P4–M3 (""): 9.1 (7.9–10.2) mm, n = 5 P4–M3 (!!): 7.8 (6.9–9.2) mm, n = 5 Zambia (H. Burda unpubl.) HB: 241.1 (222–260) mm, n = 5 T: 26.0 (20–32) mm, n = 4 HF: n. d. E: 46.8 (42–50) mm, n = 5 WT: n. d. GLS: 55.7 (53.8–57.2) mm, n = 5 GWS: 38.0 (35.7–39.1) mm, n = 5 P4–M3: 9.0 (8.8–9.4) mm, n = 5 Mt Moko, Angola (BMNH) Key References Burda & Kawalika 1993; Bennett & Aguilar 1995; Scharff et al. 1999, 2001a. Nigel C. Bennett & Hynek Burda

Cryptomys ochraceocinereus OCHRE MOLE-RAT Fr. Rat-taupe ocre; Ger. Ockerfarbiger Graumull Cryptomys ochraceocinereus (Heuglin, 1864). Nova Acta Acad. Caes. Leop., Dresden 31: 3. Upper Bahr-el-Ghazal, Sudan.

Taxonomy Originally described in the genus Georychus. Recently placed in the genus Fukomys by Kock et al. (2006) (see genus Cryptomys). Synonyms: kummi, lechei. Subspecies: none recognized here (but see Geographic Variation). Chromosome number: not known. Description Medium-sized mole-rat. Pelage short, dense and soft. Dorsal pelage medium-brown, sometimes with silvery tinge. Ventral pelage similar in colour to dorsal pelage. Conspicuous roundish white head-spot (5–15 mm diameter) on forehead in some individuals, although absent in others (e.g. in parts of Uganda). Eyes blue. Conspicuous white eye-ring. Fore- and hindfeet small, broad and naked. Forefoot with five digits, hindfoot with five digits; all with sharp claws. Small fringe of pale white hairs around base of hindfoot. Tail (ca. 10% of HB) barely visible, mostly obscured by hair-like bristles. Skull: infraorbital foramina round-oval (ca. 2 mm diam.), thick-walled; upper incisors ungrooved. Nipples: 1 + 1 = 4.

GeographicVariation De Graaff (1975) records two subspecies: C. o. ochraceocinereus from Upper Bahr el Ghazal, Sudan, and C. o. oweni from S Sudan. Distribution Endemic to Africa. Primarily Northern Rainforest– Savanna Mosaic of central Africa. Recorded from S Sudan, N Uganda, N DR Congo and Central African Republic. Locality records are widely distributed. Presence in W Cameroon and NW Kenya (as shown on map) is uncertain. Habitat Wooded savanna (e.g. Isoberlinia woodlands) and cultivated land. Abundance Uncertain; widespread but localized. In Didinga Hills, S Sudan, recorded as ‘Plentiful in mountain-meadows’ (Setzer 1956).

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Cryptomys zechi

groups of three or four close together. An enlarged area in a passage forms the nest, which is lined with grasses on the outside and with roots of trees and leaves on the inside (Verheyen & Verschuren 1966). Mounds are 7–10 cm high and 20–25 cm at the base (Hatt 1940a). Herbivorous. In Garamba N. P. (NE DR Congo) feeds on roots, principally on those of Diascorea abyssinica (Verheyen & Verschuren 1966, as C. lechei). Conservation

IUCN Category: Least Concern.

Measurements Cryptomys ochraceocinereus HB: 169 (157–200) mm, n = 14 T: 18 (14–27) mm, n = 24 HF: 31 (27–35) mm, n = 14 E: 0 mm WT: n. d. GLS: 43.2 (39.7–48.2) mm, n = 16 GWS: 29.4 (27.1–32.9) mm, n = 16 P4–M3: 7.2 (6.9–8.8) mm, n = 16 N DR Congo (Hatt 1940a, as C. lechei) Cryptomys ochraceocinereus

Key References

Setzer 1956; Verheyen & Verschuren 1966.

Remarks Subterranean. The burrow is a network of tunnel passages, which may attain an overall length of 315 m with up to 32 mounds of excavated soil. The mounds are irregularly placed and in

Nigel C. Bennett

Cryptomys zechi TOGO MOLE-RAT Fr. Rat-taupe du Togo; Ger. Togo Graumull Cryptomys zechi (Matschie, 1900). Sber. Ges. naturf Freunde, Berlin, p. 146. Near Kete Krachi, Middle Volta, Togo. (Old Kete Krachi, now innudated by L. Volta, is in present-day Ghana, not Togo [Grubb et al. 1998]).

Taxonomy Originally described in the genus Georychus. Recently placed in the genus Fukomys by Kock et al. (2006) (see genus Cryptomys). Synonyms: none. Chromosome number: not known. See Rosevear (1969) for further information. Description Pale-coloured mole-rat, sometimes with whitish markings on head or body. Pelage soft and short, with velvet-like texture. Dorsal pelage pale cinnamon to buff (and slightly variable); hairs unicoloured, may be white very close to base.Ventral pelage not recorded. Head blunt, usually with white patch on forehead; incisor teeth visible outside lips. Long vibrissae. Eyes very small. External ear absent. Limbs short. Fore- and hindfeet well-developed and naked. Five digits on fore- and hindfeet.Tail very short (ca. 8% of HB), covered with stiff bristles. Skull: widest part of nasal bones posteriorly; infraorbital foramina round-oval (ca 2 mm diam.), thick-walled; upper incisors ungrooved and comparatively wide (ca. 3.2 mm). Nipples: 2 + 1 = 6. Geographic Variation None recorded. Similar Species C. foxi. On average smaller; pelage dark sepia-brown; widest part of nasal bones anteriorly; P4–M3 shorter (ca. 7.6 mm); incisor teeth comparatively narrow (ca. 2.4 mm). Nigeria only.

Cryptomys zechi

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Distribution Endemic to Africa. Guinea Savanna BZ. Recorded from NW Ghana and between the Oti and Volta rivers in Togo. One specimen (BMNH) allocated to this species from SW Sudan (Rosevear 1969) is probably C. ochraceocinereus (Grubb et al. 1998). Habitat Grasslands with a few scattered trees (typically Khaya senegalensis, Butyrospermum parkii, Adansonia digitata, Parkia clappentoniana and Ceiba pentandra) (Yeboah & Dakwa 2002). Abundance Unknown; presumably rare with a rather restricted distribution. Remarks Colony size 1–7 (average four animals). Sex ratio of colony members is parity. It is believed that animals occur in small colonies comprising a reproductive pair and their litter. Reproduction restricted to the reproductive pair (Yeboah & Dakwa 2002). Conservation

Measurements Cryptomys zechi HB: 167 mm, n = 4 T: 15 mm, n = 4 HF: 27 mm, n = 4 E: 0 mm, n = 4 WT: 217 (155–283) g, n = 48 GLS: 42.7 mm, n = 4 GWS: 31.0 mm, n = 4 P4–M3: 8.3 mm, n = 4 Throughout geographic range (Rosevear 1969; mean values only, range not given) Weight: Atebubu District, Ghana (Yeboah & Dakwa 2002) Key References Dakwa 2002.

Grubb et al. 1998; Rosevear 1969; Yeboah & Nigel C. Bennett

IUCN Category: Least Concern.

GENUS Georychus Cape Mole-rat Georychus Illiger, 1811. Prodr. Syst. Mamm. Avium., p. 87. Type species: Mus capensis Pallas, 1778.

Georychus capensis.

A monotypic genus with a restricted distribution in South Africa. The genus occurs from the Cape Peninsula of Cape Province eastwards through Eastern Cape Province to KwaZulu–Natal and Mpumalanga Provinces. Preferred habitat is loose sandy soils and loams in mesic regions of South Africa, usually where mean annual rainfall is >500 mm. Georychus mole-rats are of intermediate size, being larger than most species of Cryptomys but smaller than Bathyergus. The pelage is distinctive: head black with flat white muzzle, white lips and eye-ring; and large white patch around auditory meatus. Dorsal pelage of body varies from dark grey, russet to orange-cinnamon; ventral pelage whitish. Digits of fore- and hindfeet are short, with short claws. Tail is short and white. The skull is less robust than in Bathyergus and is characterized by upper incisors without grooves, which have their roots in the pterygoid bones behind the cheekteeth. Each upper cheektooth has one narrow inner and one outer fold that persist in adults (the only genus without simplified ovate cheekteeth in adults). The jugal bone dovetails into a backward projection of the zygomatic arch (Figure 105). Other characters of the genus are given in the species profile. The single species is Georychus capensis.

Figure 105. Skull and mandible of Georychus capensis (BMNH 95.9.3.18).

Nigel C. Bennett

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Georychus capensis

Georychus capensis CAPE MOLE-RAT (BLESMOL) Fr. Rat-taupe du Cap; Ger. Kap-Blessmulle Georychus capensis (Pallas, 1778). Nova. Spec. Quad. Glir. Ord., 76: 172. Cape of Good Hope, South Africa.

Taxonomy Originally described in the genus Mus, but later placed in Marmota (see de Graff 1981). In 1811, Illiger proposed the genus Georychus with Mus capensis as the type. In 1832, Smuts referred to the species as Bathyergus capensis. Modern authors, e.g. De Graaff (1981) and Smithers (1983), refer to the species as Georychus capensis. The populations from Western Cape Province and from KwaZulu–Natal could represent two different species as allozyme and mitochondrial DNA RFLP analyses suggest they are divergent (Honeycutt et al. 1987; Nevo et al. 1987). Synonyms: buffonii, canescens, leucops, yatesi. Subspecies: none. Chromosome number: 2n = 54, aFN = 100 (Taylor et al. 1985, Nevo et al. 1986). Description Medium-sized mole-rat with distinctive black and white colouration on head. Pelage thick and woolly. Dorsal pelage russet, often with a brownish tinge. Ventral pelage silvery-white. Unlike other genera, tactile hairs do not project from the pelage on the body. Head large and blunt, black, charcoal or deep russet, with a pattern of white markings unlike that of any other mole-rat. Muzzle and lips white. Eyes black (a little larger than other bathyergids) with large white eye-ring. Large white patch around auditory meatus. Fore- and hindfeet usually white. Tail very short (ca. 13% of HB), pink with a number of coarse white hairs radiating from it. Skull: dorsoventrally flattened; old animals possess distinctive sagittal and nuchal crests; jugal dovetails into backward projection of the zygomatic arch; infraorbital foramina small, round (ca. 1–1.5 mm), thick-walled. Upper cheekteeth each with one narrow inner and outer fold that persist in adults. Upper incisors ungrooved. No sexual dimorphism. Nipples 2 +1 = 6. GeographicVariation

None except for some genetic characters.

Similar Species Bathyergus suillus. Larger; pelage cinnamon-brown, and sometimes with darker mid-dorsal band; forefeet with elongated claws; upper incisors with grooves. Cryptomys hottentotus. On average much smaller; pelage fawn to grey. Distribution Endemic to Africa. South-West Cape BZ and some parts of the Highveld BZ. Recorded only from South Africa; distribution is disjunct in mesic regions where mean annual rainfall is >500 mm. Recorded from Cape Peninsula in SW Western Cape Province northwards to Citrusdal and Niewoudtville, and eastwards to Port Elizabeth and W Eastern Cape Province (formerly Transkei). Isolated populations occur in KwaZulu–Natal Province near the border with Lesotho, and at Belfast, Wakkerstroom and Ermelo in Mpumalanga Province (formerly Eastern Transvaal). In S Western Cape Province, found at 12–510 m and in KwaZulu–Natal at 1372–1700 m. Habitat Coastal and montane fynbos, forest and savanna grassland, where mean annual rainfall is 500–800 mm. Occurs in sandy loams, alluvium and clay soils. In coastal regions, occurs where annual rainfall

Georychus capensis

is 279–728 mm (mean ca. 500 mm) and in KwaZulu–Natal where annual rainfall is 954–1278 mm (mean ca. 800 mm). Sympatric with Bathyergus suillus and Cryptomys hottentotus in coastal areas where the sands are more consolidated. Abundance Uncommon and localized, but densities may exceed 30 animals/km2 in Cape Town. Adaptations Strictly solitary and highly territorial. Multiple occupancy of burrows only during the breeding season, when the mate is present or when ! has young. Territorial drumming, using both hindfeet simultaneously, is frequently performed to advertise presence in the burrow system.The burrow system can exceed 130 m in length, and the burrow diameter is typically 100 mm. The burrow system comprises a central nest, a food store and a toilet area located away from the nest area. The food store consists of geophytes packed with soil; stored geophytes are disbudded when they sprout. Mounds are thrown up after rain when the soil is moist. Cape Mole-rats have been observed above ground on occasions, particularly after dusk. It is thought that in mountainous regions, where there are pockets of soil interspersed with rocky barriers, they disperse above ground. Cape Mole-rats have a low basal metabolic rate of 0.59 ml O2/ g/h, and low body temperature (36 °C). Foraging and Food Herbivorous. Food is located as the mole-rats excavate their superficial (5–25 mm deep) foraging tunnels of diameter 7–8 cm. Most extensions to the burrows occur after rainfall when the costs of burrowing are lowest.The diet consists predominantly 663

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of geophytes – bulbs, corms and tubers – but also ca. 6% aboveground vegetation. The geophytes include species of Hyacinthaceae (e.g. Albuca, Lachenalia and Ornithogalum), Iridaceae (e.g. Romulea, Micranthus and Homeria) and Oxalidaceae (e.g Oxalis). A number of these contain cardiac glycosides and are toxic to livestock but not to mole-rats. Cape Mole-rats selectively store the larger-sized geophytes of many species, and food stores may exceed 5000 items. The food in the store is probably eaten when ! has young or during unfavourable periods when soils are difficult to excavate (Lovegrove & Jarvis 1986). Social and Reproductive Behaviour Solitary, aggressive and territorial. Seismic signalling (which differs from territorial drumming) occurs at the onset of reproduction. Males and !! drum at different frequencies. The onset of courtship seismic signalling in "" is accompanied by raised urinary testosterone concentrations, and enlargement of the testes and accessory reproductive glands. Hindfoot seismic signalling by "" is extremely fast, each drum pulse in "" lasting two minutes with a beat length of 0.035 seconds. Females do not drum as fast, with a beat length of 0.05 per second. Courtship is usually initiated by " and copulation is brief, involving multiple intromissions of 2–3 thrusts per second, interspersed by short periods during which the animals are involved in bouts of grooming, particularly around the genitalia (Bennett & Jarvis 1988, Narins et al. 1992). Reproduction and Population Structure Reproduction is seasonal; young are born in summer (Aug–Dec), and ! produces a maximum of two litters/summer. Gestation: ca. 44 days. Litter-size: 6 (3–10), n = 19. At birth, young naked and blind. Pelage with the distinctive markings Day 7. Eyes open Day 9. First solid foods eaten ca. Day 17.Weaned Day 28. Pup development is comparatively rapid compared with that of social species. Inter-sibling aggression begins around Day 35, and young disperse (either below or above ground) at about Day 50.

when moving above ground. Preyed upon by Mole-snakes Pseudapsis cana, cobras, owls, herons and small carnivores. Few parasites are associated with these mole-rats: ectoparasites includes four species of mites, a tick and a flea; endoparasites include a tapeworm (Echinococcous sp.) and a nematode (Trichurus sp.). Mole-rats are susceptible to bubonic plague (De Graaff 1981). Conservation IUCN Category: Least Concern. Cape Mole-rats are occasionally an agricultural pest, and can cause problems for horticulturists and green-keepers. Measurements Georychus capensis HB (""): 189 (177–200) mm, n = 12 HB (!!): 182 (155–204) mm, n = 29 T (""): 31 (25–40) mm, n = 12 T (!!): 26 (20–33) mm, n = 29 HF (""): 32 (30–35) mm, n = 12 HF (!!): 29 (27–35) mm, n = 29 E (""): 0 mm E (!!): 0 mm WT (""): 181 g, n = 51* WT (!!): 180 g, n = 37* GLS (""): 48.3 (44.1–53.3) mm, n = 12 GLS (!!): 45.1 (41–51.2) mm, n = 29 GWS (""): 37.9 (35.1–39.9) mm, n = 12 GWS (!!): 32.8 (30.1–40.4) mm, n = 29 P4–M3 (""): 8 (7.6–9.4) mm, n = 12 P4–M3 (!!): 7.8 (6.8–8.5) mm, n = 29 Western Cape Province, South Africa (De Graaff 1964a, Taylor et al. 1985) *Range not recorded Key References

Predators, Parasites and Diseases Cape Mole-rats are particularly vulnerable to predation during mound formation and

Bennett & Jarvis 1988; Taylor et al. 1985. Nigel C. Bennett

GENUS Heliophobius Silvery Mole-rat Heliophobius Peters, 1846. Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin 11: 259. Type species: Heliophobius argenteocinereus Peters, 1846.

Heliophobius argenteocinereus.

Currently considered to be a monotypic genus distributed from Kenya southwards through Tanzania to E DR Congo, N Zambia, Malawi and Mozambique north of the Zambezi R. Occurs in a variety of savannas and woodlands, and in many soil types from ‘black cotton’ soil (very sticky when wet, hard when dry) to well-drained sandy soils. The genus is distinguished from other genera in the family by long, silky pelage, up to six simple upper and lower cheekteeth (not usually all present at the same time), and a narrow palate that does not extend posteriorly beyond the level of the cheekteeth. Cheekteeth show re-entrant folds in young animals. Unlike Bathyergus, and like the other genera of mole-rats, the angular process of the lower jaw does not extend far posteriorly to the skull (Figure 106).

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Heliophobius argenteocinereus

Biological characters of the genus include solitary behaviour, seasonal breeding, and a long gestation (87–101 days) that limits !! to having one litter a year (up to five young/litter).The other solitary genera (Bathyergus, Georychus) have shorter gestations that allow two litters in the breeding season after winter rainfall (late winter, early spring). Heliophobius has a wider geographic distribution than the other solitary genera (Jarvis & Bennett 1991, Sumbera et al. 2003a). Ellerman (1940) recognized two species, H.argenteocinereus and H. spalax (from Kilimanjaro, Tanzania) although Honeycutt et al. (1991) consider the genus to be monotypic with variations that are age-related. Recent studies have shown large genetic divergencies between animals from Kenya and from Malawi (up to 15.4% HKY85 corrected cytochrome b sequence differences; C. G. Faulkes pers. comm.). Furthermore, there are chromosomal differences between the Zambian populations (2n = 62, Scharff et al. 2001b) and those in Kenya (2n = 60, George 1979a). The systematics of this genus requires further study. The single species is Heliophobius argenteocinereus. J. U. M. Jarvis

Figure 106. Skull and mandible of Heliophobius argenteocinereus (HC 2438). Subadult: only three cheek-teeth visible (cf. 4–5 in adults).

Heliophobius argenteocinereus SILVERY MOLE-RAT Fr. Rat fouisseur gris-argent; Ger. Silbergrauer Erdbohrer Heliophobius argenteocinereus Peters, 1846. Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin 11: 259. Tete, Mozambique. (See also below.)

Taxonomy Ellermannn (1940) listed two species of Heliophobius: H. argenteocinereus and H. spalax, as well as eight subspecies of H. argenteocinereus (Honeycutt et al. 1991). All forms currently considered to be synonyms of H. argenteocinereus. However, genetic differences between animals from Kenya and from Malawi suggest that populations in different parts of the geographic range may be specifically or subspecifically different (see genus profile). Molecular studies suggest that there is a good case for recognizing five genetically divergent clades within the species: robustus/mottuolei species complex (clade 1), H. kapiti (clade 2a), H. spalax (tentatively clade 2b), H. emini (clade 3), argenteocinereus/angonicus complex (clade 4) and H. marungensis (clade 5) (C. Faulkes and N. Bennett, pers. comm.). Synonyms: albifrons, angonicus, emini, kapiti, marungensis, mottoulei, pallidus, robustus, spalax. Subspecies: none currently recognized (but see above and Geographic Variation). Chromosome number: 2n = 60 (George 1979a), 2n = 62 (Scharff et al. 2001b). Description Medium-sized silvery-grey mole-rat with soft silvery pelage. Pelage silky and long (20–25 mm). Dorsal pelage silvery-grey to tan. Nose, eye region, lateral parts of head, limbs and ventral pelage paler. Head similar to dorsal pelage, paler around muzzle, eyes and nose; small white patch or fleck on forehead in some individuals (ca. 50% of those in Malawi, R. Sumbera pers. comm.).

Incisor teeth long, slightly curved and lying outside the lips. Limbs short. Feet, pale-coloured, not enlarged, outer edges fringed with stiff hairs. Five digits, not strongly clawed. Tail very short (8–9% of HB), pale-coloured, with fringe of stiff hairs. Sexual dimorphism occurs in some regions (see below). Infraorbital foramen round (diam. 1.5 mm), thick-walled. Upper incisors ungrooved. (See also genus profile.) Nipples: 2 + 1 = 6. Geographic Variation Silvery Mole-rats from Malawi are larger than those recorded elsewhere; they also have shorter pelage (10–15 mm) and show sexual dimorphism with "" being about 15% larger than !! (Sumbera et al. 2003a). Tail length is shorter than in specimens from Kenya, and !! have significantly longer tails (12.9 ± 1.8 mm) than the "" (12.4 ± 1.7) (R. Sumbera pers. comm.). There is a white patch in the axilla of animals from Athi Plains, Kenya, but not from Malawi (J. Jarvis & R. Sumbera unpubl.). The pelage is darker and the head-patch is larger in animals from Morogoro, Tanzania (C. Faulkes pers. comm.). Molecular genetic studies also show considerable geographic variation (see above). Similar Species Tachyoryctes splendens. Similar size; ear pinnae present, but small; eyes larger; incisors orange. (Although subterranean in habit, as are 665

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47 m (range 39–61, n = 4) in length (Jarvis & Sale 1971). In Malawi, mean burrow lengths are 73 m (range 22–138) at Blantyre, and 105 m (range 39–188) at Mulanje (Sumbera et al. 2003b, R. Sumbera unpubl.). Nest chambers are slightly deeper (300 mm) and contain a hollow ball of nesting material (grass, corm husks, skins of tubers); deeper still is a bolt-hole (up to 540 mm) where a mole-rat can retreat when threatened. Toilet areas are in blind-ending tunnels (Sumbera et al. 2003b).When the black cotton soil is wet, the animals use balls of mud to repair damaged burrows. Burrow systems often associated with slightly raised areas with better drainage (J. U. M. Jarvis unpubl.). Individuals show no particular activity pattern, and are active during the day and night (Jarvis 1973b).

Heliophobius argenteocinereus

bathyergid mole-rats, Tachyoryctes splendens belongs to a different family of rodents – Spalacidae: Tachyoryctinae.) Distribution Widespread in Zambezian Woodland BZ and southern part of Somalia–Masai Bushland BZ. Recorded from C and S Kenya, Tanzania, Malawi, E Zambia, C and N Mozambique, and E DR Congo. Occurs from ca. 100 m to >2000 m and where annual rainfall is usually 250–900 mm, and up to 1500 mm (in Malawi). The type locality is given as Tete (which is on the southern (right) bank of the Zambezi R.); however, the type description records that the holotype (and specimens of other species) came from ‘Tette’ without saying precisely where they were caught. Smithers & Lobão Tello (1976) show that all records in Mozambique come from north of the Zambezi R., although Smithers (1983), which considers only Africa south of the Zambezi, shows a small range south of the Zambezi. As pointed out by Skinner & Smithers (1990), it seems unlikely that the species occurs south of the Zambezi. Habitat Combretum and Brachystegia woodlands, rocky hillsides and sub-montane grasslands. In Malawi, frequently invades crops of banana, cassava and sweet potato (R. Sumbera pers. comm.). Occurs in a wide range of soils from well-drained easily worked sandy soils to heavy ‘cotton soils’ that are very hard when dry but sticky and waterlogged when wet (Kingdon 1974, Ansell & Dowsett 1988, J. U. M. Jarvis unpubl.). Abundance Little information. Densities are low on the Athi Plains, Kenya (J. U. M. Jarvis unpubl.). In Malawi, density is 5.2/ha at Blantyre (Sumbera et al. 2003a, b). Sex ratio is parity (Sumbera et al. 2003a). Adaptations Subterranean. Silvery Mole-rats dig using their chisel-like incisor teeth. On the Athi Plains, Kenya, foraging burrows are 12–23 mm below the surface, 50 mm in diameter and average

Foraging and Food Herbivorous. Little known of the diet in the wild. In Kenya and Malawi, partly eaten tubers of Dolichos sp. and Vigna sp., still growing in situ, were apparently harvested as needed by the mole-rats (Jarvis & Sale 1971, R. Sumbera pers. comm.). Of the two tubers, Vigna is the preferred food and appears to be the key species determining the occurrence of Silvery Mole-rats near Blantyre (R. Sumbera pers. comm.). In Malawi, other species such as Gladiolus dallenii, Imperata cylindrica and Hypoxis sp., as well as a variety of cultivated root crops, have also been found in food stores (either in special chambers or blind-ending tunnels) (R. Sumbera pers. comm.). In captivity, animals are strongly selective, preferring bulbs and tubers, which they peel before eating. They practise coprophagy, seizing faecal pellets as they are voided from the anus with their incisors, and eating up to 12 pellets at one time (Jarvis 1969b). Social and Reproductive Behaviour Silvery Mole-rats aggressively defend burrows against conspecifics. Animals emit a snorting hiss when cornered, assume a rigid stance with the feet braced and head thrown up, and with mouth and eyes wide open. When fighting, opponents lock incisors and sometimes roll over while still maintaining a grip on the opponent. Territorial drumming has not been reported (Jarvis 1969a, Sumbera et al. 2003a), but short, fast, repeated drumming with the forelimbs occurs during highly aggressive encounters (Sumbera 2001). In Kenya, animals are unusual amongst bathyergids in emitting an almost continuous cry (similar to a newborn baby) when alarmed (such as being transported by car; Jarvis 1969a). Silvery Mole-rats from Malawi are not very aggressive and will often coexist in captivity. Courtship behaviour (Sumbera 2001) is initiated by ", who follows !, sometimes sniffing her anogenital region. The animals face each other, gently nibble each other or lock incisors. Sometimes one of them (of either sex) lies on its back. Both animals vocalize, ! more so than ".The " frequently urinates on vertical surfaces. Repeated intromissions (10–30) and rapid thrusting (7.3/sec) occur during copulation. Aggression has not been observed at the end of mating (Sumbera 2001). Reproduction and Population Structure Reproduction is seasonal. In Malawi, mating occurs during the beginning of the cold dry season (Apr–Jun) and young are born when it is hot and dry (Aug–Oct) (Sumbera et al. 2003a), whereas limited data from Kenya indicate that young are born at the onset of the long rains (Apr– Jun) (Jarvis 1969a). Gestation: 87–101 days (n = 3 litters; Jarvis & Bennett 1991, Sumbera et al. 2003a). Litter-size: 3.2 (2–5), n = 27 litters (Sumbera et al. 2003a). At birth, mean weight of young 12.8 g

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(n = 5). First solids eaten Day 8–11. Eyes open Day 13–14. Weaned ca. 2 months. Mean adult weight ca. 12 months. The ability to engage in serious sparring/fighting appears to be correlated with attainment of a specific body weight (and not with age), ranging from ten weeks (" WT: 94 g) to 22 weeks (" 80 g) (Sumbera et al. 2003a). The long gestation precludes the ability of ! to have more than one litter each breeding season. The lack of seismic signalling, ‘relatively high pacifism in captive animals’, capture of mature animals above ground, and no evidence of interlinking burrows suggest that this species may find its mates by moving above ground (Sumbera et al. 2003a). Predators, Parasites and Diseases Little information available. However, predators are probably snakes, raptors and small carnivores. Endoparasites include four species of Eimera (Koudela et al. 2000, Modry et al. 2005), Protospirura muricola and Inermicapsifer arvicanthidis (Tenora et al. 2003). Ectoparasites include an unidentified species of mite (Trombiculidae, Acarina) (R. Sumbera unpubl.). Conservation

IUCN Category: Least Concern.

Measurements Heliophobius argenteocinereus HB (""): 161 (131–195) mm, n = 69 HB (!!): 158 (131–191) mm, n = 129 T (""): 12.4 (6–16) mm, n = 64 T (!!): 12.9 (8–17) mm, n = 62

HF (""): 28.3 (18–31) mm, n = 69 HF (!!): 28.4 (23–34) mm, n = 129 E (""): 0 mm E (!!): 0 mm WT (""): 145 (107–220) g, n = 70 WT (!!): 153 (110–259) g, n = 128 GLS (""): 39.1 (33.9–45.0) mm, n = 68 GLS (!!): 38.1 (32.9–43.0) mm, n = 124 GWS (""): 30.3 (26.8–35.3) mm, n = 69 GWS (!!): 30.6 (25.3–34.8) mm, n = 124 P4–M3 (""): 7.5 (6.1–9.4) mm, n = 69 P4–M3 (!!): 7.7 (6.1–9.3) mm, n = 125 Morogoro and Mlali, Tanzania (W. Verheyen unpubl.) HB (""): 155.3 (106–193) mm, n = 64 HB (!!): 148.8 (108–180) mm, n = 62 HF (""): 29.8 (24–36) mm, n = 64 HF (!!): 28.7 (23–35) mm, n = 62 WT (""): 190.1 (63–331) g, n = 70 WT (!!): 162.1 (51–271) g, n = 74 Malawi (Sumbera et al. 2003a) Key References Jarvis 1973b; Jarvis & Sale 1971; Sumbera 2001; Sumbera et al. 2003a. J. U. M. Jarvis

GENUS Heterocephalus Naked Mole-rat Heterocephalus Rüppell, 1842. Mus. Senckenbergianum Abh. 3 (2): 99. Type species: Heterocephalus glaber Rüppell, 1842.

Heterocephalus glaber.

Heterocephalus is a monotypic genus distributed only in NE Africa (parts of Kenya, Ethiopia and Somalia). The habitat is semi-desert where the annual rainfall is 50). Population size difficult to estimate because mole-rats are subterranean.

Figure 107. Skull and mandible of Heterocephalus glaber (BMNH 7.4.1.12).

Adaptations Subterranean, but may very occasionally venture above ground. Naked Mole-rats run forwards and backwards with equal ease, using their tail and sensory hairs to navigate. Burrow

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of food encountered. Burrows branch finely when patches of food are encountered, but not when they encounter large tubers, as these occur singly. Large tubers are partly eaten in situ, and the cavities are then packed with soil; many tubers are not totally destroyed and will regenerate (Brett 1991a, b). Non-reproductive animals of both sexes cooperate to dig foraging burrows: one animal bites the earth face, others collect excavated soil and form a chain to kick it back along the burrow to a larger animal who kicks it onto the surface through an open hole (Jarvis & Sale 1971, Braude 1991). Active mounds resemble an erupting volcano and are the site where most predation occurs. Most mounds are produced between 02:00 and 08:00h, times when predatory snakes are least active (Brett 1991a, b). In the laboratory, foragers (the non-reproductive animals) returning to the nest leave scent trails back to the food (Judd & Sherman 1996). Small items of food may be carried back to the nest where they are eaten by the young or by other colony members. Naked Mole-rats do not store food and all colony members (including the reproductive animals) will visit the primary source of food to feed.

Heterocephalus glaber

system length depends on colony size and local food abundance. One completely excavated system containing 60 animals was 595 m long (Jarvis 1985), another with 85–90 animals was estimated (from telemetry studies over 12 months) to be 3 km long (Brett 1991b). Burrow systems consist of superficial networks (2–20 cm deep) of narrow (2.5–3 cm) foraging burrows, which are blocked off once the animals have finished foraging in the area. Superficial burrows lead down to wide (4–5 cm), long and relatively un-branched main burrows (>30 cm deep), which are dug to locate large tubers and patches of food; they also lead to nests (with several entrances), blind-ending toilet burrows and deep ‘bolt-holes’ – these may descend steeply to ca. 2 m (Jarvis 1985, Brett 1991b). Temperatures in the main burrows vary little diurnally and seasonally (ca. 28–32 °C), and the humidity is high (ca. 80%). Naked Mole-rats are poikilothermic over most ambient temperatures, using behavioural means (huddling, basking close to the surface) to maintain body temperature close to ambient temperature (Buffenstein & Yahav 1991b). The well-vascularized hairless skin facilitates rapid heat exchange; and the high humidity in the burrow reduces water loss but prevents evaporative cooling. The basal metabolic rate is exceptionally low (40% of expected; McNab 1966); other energydemanding features such as eyes and the optic regions in the brain are reduced to conserve energy. Energy-saving features help reduce life-time expenditure of energy, reduce cumulative oxidative damage and may possibly promote longevities of >26 years for captive breeders and non-breeders (O’Connor et al. 2002, Sherman & Jarvis 2002). In the wild, longevity is 10 years for reproductive animals (S. Braude unpubl.). Foraging and Food Herbivorous, feeding exclusively on roots and swollen underground storage organs of plants. Food is widely dispersed in patches of small items or single large tubers (up to 50 kg). After rain, Naked Mole-rats dig long prospecting foraging burrows (up to 1 km/month) whose pattern is modified depending on the type

Social and Reproductive Behaviour Naked Mole-rats are highly social, indeed they resemble eusocial insects in having reproductive division of labour, overlap of generations and cooperative care of young (Jarvis 1981, Sherman et al. 1991). Only one ! per colony is reproductively active (occasionally two); she is the dominant ! and suppresses reproduction in subordinates, not through pheromones but through stress-inducing behavioural interactions (shoving, passing over, etc.). She solicits mating with 1–3 consorts, who may remain her consorts throughout her life, occasionally continuing as reproductive "" to the succeeding reproductive !. When in oestrus, she emits a distinctive call and crouches in front of the " or "" with tail to one side. Multiple paternity may occur in a litter (Faulkes et al. 1997a). There are no castes of workers but smaller non-reproductive animals (both adults and subadults) tend to perform more maintenance and foraging tasks than larger adults and also devote more time caring for the young. Larger adults tend to act as defenders in times of danger. The reproductive ! may remain as the only reproductive ! for many years (sometimes >20 years in the laboratory and >10 years in the wild).The oldest, largest !! (irrespective of relatedness) typically fight viciously for reproductive succession when a vacancy occurs (O’Riain 1996, Van der Westhuizen 1997, Jarvis & Sherman 2003). This is usually when the dominant reproductive ! dies, but also sometimes if she is sick or in some other way unable to dominate her colony; on such occasions she is usually killed by rivals.The lumbar vertebrae of the new dominant ! elongate during her first pregnancies, rendering her morphologically distinct from other !!; this unusual situation is the first mammalian example of a behaviourally and morphologically distinct caste (O’Riain et al. 2000). Her enlarged abdomen enables her gut to hypertrophy to cope with the greatly increased energy demands of having large litters. It also enables her to remain mobile in the narrow confines of the burrow and thereby to maintain dominance over her colony. Males and !! remain fertile for >23 years (Buffenstein & Jarvis 2002, O’Connor et al. 2002, Sherman & Jarvis 2002). Colonies are strongly xenophobic, recognizing colony members by a cocktail of odours acquired from huddling in the communal nest and grooming in the communal toilet (O’Riain & Jarvis 1997). Because reproductive succession often occurs within the colony, many colonies are highly 669

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inbred (Faulkes et al. 1997a). Occasionally fat, highly sexed, dispersers appear who, in laboratory experiments, will actively solicit foreign animals but not members of their own colony (O’Riain et al. 1996). These individuals may disperse to found new colonies, and perhaps even enter other colonies, but it is not known if dispersal is above or below ground. Small incipient and out-bred colonies are occasionally found in the wild; such colonies rarely last more than a year, indicating the high risks taken in attempting to‘go it alone’(O’Riain & Braude 2001, Jarvis & Sherman 2003). The vocal repertoire of Naked Mole-rats is unusually large for a rodent (>18 vocalizations). Most vocalizations are in the 1–9 kHz frequency range and include high-pitched contact and aggressive chirps, a prolonged alarm scream, and aggressive grunting towards outside sources of danger. Larger individuals are mobilized, through the alarm calls, to defend the colony. Some vocalizations are restricted to specific animals in the colony, e.g. juveniles, reproductive animals (Pepper et al. 1991). Although most activities in the colony are accompanied by frequent vocalizations, the primary modes of communication, odour and touch, are also used (Sherman et al. 1991, O’Riain & Jarvis 1997). Reproduction and Population Structure The one (occasionally two) reproductive !! in a colony breed throughout the year and have 3–4 litters/year. Gestation: 70–74 days. Litter-size 11.4 (1–28), n = 190. Postpartum oestrus Day 10. At birth, young weigh 1–2 g. First solids eaten by Day 14. Fully weaned at ca. 5 weeks (Jarvis 1991, O’Riain 1996).Young do not disperse but join the nonreproductive workforce. Only the reproductive ! lactates but, in many captive colonies, the entire colony ("" and !!) develop nipples prior to the birth of the young.This hormonal response to the pregnancy of the dominant ! by the colony may prime the animals to accept, and help care for, her young. The energetic costs of breeding are high; during pregnancy energy demands of the breeding ! increase by ca. 1300 kJ per gestation cycle, and lactation requires an additional 1515 kJ per day for an average-sized litter (Urison & Buffenstein 1995). Lactation lasts about five weeks. On weaning, the juveniles solicit caecotrophs from adults (special nutritious faeces routinely eaten by mole-rats as they are voided), and also eat food brought to the nest. The reproductive ! is the only adult that can solicit caecotrophs from other adults; they form an important dietary supplement to enhance her reproductive success. Of all the !! in the colony, the dominant (reproductive) ! is identified by her elongated body, prominent nipples and perforate vagina. Reproductive "", often small emaciated-looking animals, are strongly bonded to the ! through repeated mutual anonasal interactions. In colonies with >1 reproductive ", there is no apparent competition for mating rights (Jarvis 1991). In the wild, the

reproductive ! and reproductive "" are usually the oldest members of the colony, and are distinctive in being less counter-shaded than non-reproductive animals (S. Braude unpubl.). Non-reproductive !! are prepubescent, have a vaginal closure membrane, small inactive ovaries and low levels of reproductive hormones (Faulkes & Abbott 1997); they can begin to have oestrus cycles within a week of removal from the colony. Non-reproductive "" have lower sperm counts, lower levels of reproductive hormones, and the sperm have low motility (Faulkes et al. 1994, Faulkes & Abbott 1997). Testes of reproductive and non-reproductive "" are abdominal. Over 95% of non-reproductive animals never have opportunity to breed (Sherman et al. 1992, Jarvis & Sherman 2003). Predators, Parasites and Diseases Naked Mole-rats are preyed on by snakes (especially the Rufous-beaked Snake Rhamphiophis oxyrhynchus and Sand Boa Eryx columbrinus) and various raptors. They are especially vulnerable when they build mounds and eject soil onto the surface. Probably because of their nakedness, Naked Mole-rats have few ectoparasites; the most numerous are subcutaneous mites and chiggers (Parona 1895). Endoparasitic nematodes and cestodes are very rare (300 animals autopsied from Mtito Andei; J. U. M. Jarvis unpubl.) Conservation IUCN Category: Least Concern. Local populations often highly inbred, and therefore potentially at risk from extinction by disease. There is little conflict with farmers because much of their habitat is not suitable for agriculture. Measurements Heterocephalus glaber HB: 83.4 (66–110) mm, n = 131 T: 42.1 (34–50) mm, n = 123 HF: 20.1 (18–22) mm, n = 127 WT: 33.9 (25–80) g, n = 715 GLS: 21.4 (16.5–24.4) mm, n = 119 GWS: 16.8 (12.6–20.1) mm, n = 105 M1–M3: 3.4 (3.1–3.9) mm, n = 100 Mtito-Andei, S. Kenya (J. U .M. Jarvis unpubl.) Weight: Mtito-Andei, S Kenya (Brett 1991) In areas where food is limiting, body size is considerably smaller (see Jarvis, 1985) Key References Bennett & Faulkes 2000; Hill et al. 1957; Jarvis & Sherman 2003; Sherman et al. 1991, 1992. J. U. M. Jarvis

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Family HYSTRICIDAE

Family HYSTRICIDAE PORCUPINES

Hystricidae G. Fischer, 1817. Mem. Soc. Imp. Nat., Moscow 5: 372. Atherurus (1 species) Hystrix (2 species)

Brush-tailed Porcupine Crested Porcupines

p. 672 p. 674

The Hystricidae contains three genera and 11 species distributed in Africa, the Middle East, the Indian sub-continent and South Asia, and on various islands in Indonesia and the Philippines (Woods 1984, Woods & Kirkpatrick 2005). The family, as a whole, is represented in many habitats including rainforest, savanna and semi-desert, although each species is rather restricted in its requirements. Two genera and three species occur in Africa. Porcupines are best known for their large size and weight, and the possession of very coarse hair and spines on the back in some species. The spines (or quills) vary in size, shape, length and pattern according to the species and position on the body; however, all are pointed at the tip and provide a very effective defence against potential predators. The limbs are short, the fore- and hindfeet are plantigrade and each foot has five digits with claws. Locomotion is either a rather clumsy walk or a trot. The tail is short, and covered by quills, some of which may be modified into ‘rattle quills’. Most species are brown or black in colour, although white bands on the spines of some species result in a generally paler colouration. Size categories of species in the family (based on mean head and body length) are given in the order Rodentia profile. The skull is massive and high domed with an inflated nasal region in some species (Hystrix spp.), and with large pro-odont smoothfaced incisors, a large infraorbital foramen and relatively small zygomatic arches. Dental formula is I 1/1, C 0/0, P 1/1. M 3/3 = 20. Upper incisor teeth without grooves. The premolar is retained for most of the individual’s life, but is replaced late in life. The four cheekteeth have a unique wavy and complex pattern of enamel and dentine. On the upper molars, there are three labial (outer) folds and one lingual (inner) fold of the enamel. These folds usually become isolated early in life (due to wear) and then form ‘islands’ so that there are alternating lines and rings of enamel and dentine (Figure 108). Porcupines are vegetarians, feeding on a wide variety of grasses, fruits, bark, bulbs and roots. They are nocturnal and travel at night along well-defined paths, often covering long distances in search of food. During the day they rest in burrows, which they may dig themselves, caves, or cavities under forest trees (such resting places are sometimes referred to as ‘dens’). They are primarily terrestrial and unable to climb (except for one species in Asia). Porcupines are solitary or gregarious, according to the species. In the social species, groups of several individuals of mixed sexes rest together in a single burrow. In Hystrix, parental care is well developed: !! look after young in the nest and accompany them when they begin

to forage above ground. Development of porcupines is slow (by comparison with other rodents), mainly because of their large size: gestation is 90–120 days, suckling continues for about two months, weaning occurs at about four months and adult size and reproductive maturity are not attained until about one year (or longer in the largest species). Porcupines are long-lived; in captivity, African species have survived for at least 20 years. The family comprises genera and species that are very different from other rodents and the taxonomy of the family is not controversial. Hystricidae are placed in the suborder Hystricognathi (synonym: Hystricomorpha), together with the Bathyergidae, Petromuridae and Thryonomyidae. Fossil porcupines in Africa date back to the Pleistocene, but older fossils are known from India and Eurasia (Woods 1984). Porcupines have radiated into two major groupings, which are sometimes considered as subfamilies (e.g. by Rosevear 1969): Atherurinae (Athererus, Triochys) consists of the smaller forest-living Brush-tailed Porcupines with poorly developed quills, and Hystricinae (Hystrix) consists of the larger savanna-living Crested Porcupines with well-developed quills. Following Woods (1993) and Woods & Kirkpatrick (2005), subfamilies are not listed here, even though there are considerable differences in the morphology and biology of each subfamily. The two genera (Athererus, Hystrix) in Africa are distinguished by size, tail length, character and size of the quills on the tail, and by the length of the nasal bones. D. C. D. Happold

Figure 108. Cheekteeth of Atherurus africanus (left), Hystrix cristata (centre), Thryonomys swinderianus (right). White = enamel, stipple = dentine, black = infolding from outer surface, or space within enamel ‘island’.

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Genus Atherurus Brush-tailed Porcupines Atherurus F. Cuvier, 1829. Dict. Sci. Nat. 59: 483. Type species: Hystrix macroura Linnaeus, 1758.

The genus contains one African species and one Asian species. The genus is characterized by comparatively small size (cf. Hystrix), relatively long tail with a brush of ‘rattle-quills’ at the terminal end, and comparatively short dark quills on back which do not form a crest. The skull has small nasal bones which are shorter than the

frontal bones and end anteriorly to the anterior end of the zygomatic arches (cf. Hystrix). The genus is represented only in rainforest habitats. The single African species is Atherurus africanus. D. C. D. Happold

Atherurus africanus.

Atherurus africanus AFRICAN BRUSH-TAILED PORCUPINE Fr. Athérure d’Afrique; Ger. Afrikanischer Quastenstachler Atherurus africanus Gray, 1842. Ann. Mag. Nat. Hist, ser.1, 10: 261. Sierra Leone (exact locality not specified).

Taxonomy Synonyms: africanus and armatus (West Africa), centralis (DR Congo ) and turneri (East Africa). Subspecies: none. Chromosome number: not known.

Similar Species Hystrix cristata. Much larger, and with longer spines (up to 30 cm); nasal region of skull more inflated; rattle-spines of tail with only one large elongated hollow.

Description Extremely large dark rodent, with comparatively short spines (quills) on back and flanks, and long tail ending in ‘rattle quills’. Dorsal pelage dark brown; hairs rather sparse, modified into coarse thick spines, off-white at base, darkening towards the middle, blackish-brown (without alternating black and white bands as in Hystrix spp.) and sharply pointed at tip. Length of spines variable: 20 mm on neck, 35 mm on mid-back, up to 90 mm on rump and 25–45 mm on flanks. Fine hairs grow between the spines.Ventral pelage off-white to pale brown; ‘hairs’ softer and less spiny than dorsal hairs; length ca. 10–15 mm. Head sparsely covered with short dark coarse hairs; very long black vibrissae. Ears darkly pigmented, mostly naked. Fore- and hindfeet short and thick-set, covered with coarse dark brown hairs; five digits on each foot, all with small claws (except for short Digit 1 of forefoot).Tail short (ca. 40% of HB), thick and swollen at base with short black spines; tapers towards tip; terminal end with off-white or yellow ‘rattle-quills’, each with 4–5 hollow cavities along its length. When tail is shaken, the hollow cavities of the rattle-quills produce a rustling sound. Skull rather elongated (cf. Hystrix spp.) with ‘normal’ non-inflated nasal bones that end anteriorly to zygomatic arch; incisor teeth smooth and without grooves on outer surface (Figure 109, see also Figure 108). Nipples: 2 + 0 = 4. Geographic Variation None recorded.

Figure 109. Skull and mandible of Atherurus africanus (RMCA 15288).

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Brush-tailed Porcupines walk and trot on the forest floor, but they can also scramble and climb to a limited extent. While moving, the tail is held slightly upwards and backwards from the body, and sometimes at night the rustling of the ‘rattle quills’ can be heard even though the animal appears invisible. When alarmed or threatened, the spines on the back are raised and the rattle-quills are shaken; if attacked, the animal moves sideways and backwards with the pointed quills facing towards the opponent. Brush-tailed Porcupines usually defecate regularly at the same place, either under a rock or in the den (Rahm & Christiaensen 1963). Foraging and Food Vegetarian; feeds primarily on the leaves, flowers and fruits of forest trees (most of which have fallen from the higher storeys of the forest) and on some roots (Rahm 1962b).Where farms are adjacent to forest, maize, manioc (cassava), bananas and palm nuts are favourite foods. Occasionally, porcupines feed on carrion and earthworms (Rosevear 1969).

Atherurus africanus

Thryonomys swinderianus. Pelage coarse (without spines); tail without spines; each upper incisor tooth with three longitudinal grooves. Distribution Endemic to Africa. Rainforest BZ and Rainforest– Savanna Mosaic from Gambia and throughout West Africa to E DR Congo; small populations in Uganda, Kenya and S Sudan. Bioko I. (Eisentraut 1973). Habitat Evergreen rainforests and gallery forests where there are hollow trees, buttress roots and soft soil, especially close to streams in little valleys. May occur in farmlands adjacent to forest within Rainforest BZ. Occurs over a range of altitudes from near sea level to 1400 m in E DR Congo (on volcanoes and Rwenzori Mts) and 1500 m on Mt Cameroon. Abundance May be very abundant in some localities. Estimates vary according to locality (and method of calculation), e.g. 2.4–13.2/ km2 in Bayanga, Central African Republic (Noss 1998); 55/km2 in Equatorial Guinea (Fa et al. 1995); 174 kg/km2 = ca. 58/km2 in Gabon (Feer 1993 in Jori et al. 1998). Adaptations Nocturnal, terrestrial. Brush-tailed Porcupines do not dig burrows, but hide during the day in a den in hollow trees and hollow logs, or under roots of large trees. At Makokou, Gabon, they leave their hiding places at ca. 17:00h and return at about 05:30h (Emmons 1983).The activity pattern is usually trimodal, with two rest periods during the night, although bimodal on moonlit nights. Rest periods of 10–50 min are taken during each hour – these tend to be longer in the middle of the night than at the beginning or end, and when there is bright moonlight. On dark nights, "" do not normally take a rest period in the middle of the night (23:00–01:00h), although !! rest for some of the time. On nights close to full moonlight, the average time of inactivity/night was 3.5 hours, whereas on nights close to no moonlight, the average time of inactivity/night was 1.89 hours.

Social and Reproductive Behaviour At night, Brush-tailed Porcupines usually travel alone, although they may meet at feeding places. During the day, when resting in dens, they may be gregarious. In 22 dens investigated in Gabon, there were one (n = 12), two (n = 7) three (n = 1) or four (n = 1) animals in each den; associations were either !! and "", all !!, or all "" (Emmons 1983). When living together (in captivity), porcupines show mutual grooming and auditory displays of dominance and submission. In some instances, they may form family groups (! and " and young) (Rahm 1962b).The social organization of porcupines seems to be rather fluid, without the formation of monogamous pairs or cohesive groups. Home-ranges (measured by radio-telemetry in Gabon) of adults varied from ca. 11 ha to 22 ha, !! on average having larger homeranges than "". During the night, individuals travel rapidly along a well-defined network of pathways, averaging about 100 m in ca. 4 mins. Pathways connect dens with resting sites and foraging areas. Average nightly distances travelled are impressively large: !! covered 1697 m (on moonlit nights) and 2333 m (on dark nights), and "" covered 1646 m (on moonlit nights) and 1953 m (on dark nights) (Emmons 1983). Home-ranges of !! and "" tend to overlap. Most !! travel through most of the home-range every night (Emmons 1983), but do not display territorial behaviour. Reproduction and Population Structure In E DR Congo, young animals have been found in most months of the year (Rahm 1962a), and it is likely that this pattern of reproduction occurs at other localities within the Rainforest BZ (see e.g. Jeffrey 1975). Gestation: 100–120 days (Rahm 1962a). Litter-size: 1–2 (Rahm 1962a,Weir 1974); 1–4 (Kingdon 1974). Females probably polyovular, so litter-size of only one young/litter in captivity may be related to captive conditions. Weight at birth: 150 g. The young are precocious at birth: the eyes are open, there are very soft spine-like hairs on the back and flanks, and walking on all four legs is possible. Sucking continues for ca. 2 months; solid food is first eaten at 2–3 weeks. Growth rate is comparatively slow: adult size is attained in ca. 300 days; but adult weight and sexual maturity are not reached until about two years (Rahm 1962a, Rosevear 1969). Females probably have 2–3 litters/year. Breeds well in captivity. Longevity: up to 22 years (Fa et al. 1995). 673

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Family HYSTRICIDAE

Predators, Parasites and Diseases The main predators are humans and Leopards Panthera pardus. Brush-tailed Porcupines are vigorously hunted by humans because of their succulent flesh and because (to a lesser extent) they may cause damage to crops. Carcasses are seen for sale as ‘bushmeat’ in many parts of the Rainforest BZ (see also Thryonomys swinderianus). In countries where surveys have been conducted (e.g. Nigeria [Martin 1983, Anadu et al. 1988], DR Congo [Colyn et al. 1987], Equatorial Guinea [Colell et al. 1994, Fa et al. 1995] and NE Gabon [Lahm 1993, quoted by Fa et al. 1995]), porcupines formed 6–19% of carcasses for sale in local markets on an annual basis. They are usually the third or fourth most numerous ‘species’ after antelopes (Cephalophus spp.), Cane Rats (Thryonomys swinderianus) and giant rats (Cricetomys spp.). In S Cameroon, they comprised 61% of carcasses (Muchaal & Ndjangui 1995, quoted by Jori et al. 1998). Studies to assess the impact of hunting on populations suggest that, in most places at the present time, Brush-tailed Porcupines are not overhunted, although their supposed rate of recruitment (based on rather slow reproduction) seems contrary to this assessment. One study (Feer 1993), where biomass was estimated at 174 kg/km2 (= 58 individuals/km2), suggested that maximum sustained annual yield could be 44 kg/km2/year (= 14 individuals/km2/year). Brush-tailed Porcupines are host to a malarial parasite, Plasmodium atheruri (Van den Berghe et al. 1958).

Conservation IUCN Category: Least Concern. Brush-tailed Porcupines appear to be able to retain their population numbers except where hunting pressure is very high. However, loss of forest habitat as well as hunting are cause for concern in some parts of their range. Measurements Atherurus africanus HB: 534 (508–560) mm T: 204 (177–230) mm HF: 72 (71–73) mm E: 39 (38–39) mm WT: (2500–3400) g GLS: 98.0 (87.5–104.3) mm GWS: 49.2 (45.7–52.2) mm M1–M3: 18.5 (16.7–20.3) mm West Africa (Rosevear 1969) Weight: Rahm 1962a Sample sizes not stated Key References 1969.

Emmons 1983; Rahm 1956, 1962a, b; Rosevear D. C. D. Happold

Genus Hystrix Crested Porcupines Hystrix Linnaeus, 1758. Syst. Nat., 10th edn, 1: 56. Type species: Hystrix cristata Linnaeus, 1758.

The genus contains eight species (with many synonyms) widely distributed in Africa, the Middle East, India, South-East Asia and islands of the Indonesian archipelago (Woods & Kirkpatrick 2005). The genus may be divided into three subgenera: Thecurus, Acanthion and Hystrix, the latter subgenus containing both African species and being characterized

by having long quills along the length of the back, which can be erected to form a ‘crest’. Representatives of the genus occur in savannas, woodlands, rocky areas, forests and plantations, but not in rainforests. The genus is characterized by extremely large size (HB up to ca. 800 mm, WT up to ca. 20 kg, and the largest of African

Hystrix cristata.

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Family HYSTRICIDAE

Skeleton of Hystrix cristata.

rodents), coarse black bristles on head, limbs and ventral surface, long spines (quills) with alternating black and white bands on the back and flanks, short tail covered with black-and white banded ‘rattle quills’, and large thickset head. The hairs on the head and the quills on the back can be erected to form a backwardly facing crest when the animal is stressed or in danger. Quills are periodically lost (moulted) and replaced. Tail very short, with spines but without ‘brush’ of ‘rattle-quills’ (cf. Atherurus). The skull is large and thickset, rather narrow with a high domed forehead; nasal bones very large (extending over much of the top of the skull and exceeding in length the frontal and parietal bones combined) and ending posteriorly in line with the anterior or posterior end of the orbit (cf. Atherurus); zygomatic arch short; upper incisor teeth large, pro-odont and without grooves; and with four cheekteeth on each ramus (Rosevear 1969) (Figure 110). The common name, Crested Porcupines, refers to the erectile crest on the back (cf. Atherurus). Crested porcupines are nocturnal and terrestrial, and live in holes or caves during the day. They live singly or in small family groups. They are vegetarian, eating a wide range of roots, tubers, bark and fallen fruits. On occasion they may gnaw bones, presumably to obtain additional calcium.

Figure 110. Skull and mandible of Hystrix africaeaustralis (RMCA 6191).

Within Africa, Ellerman (1940) recognized four species, although Corbet & Jones (1965) reduced the number to two, placing stegmanni as a synonym of H. africaeaustralis and galeata as a synonym of H. cristata. The two species are distinguished by length of the nasal bone, the ratio of the frontal bone to the nasal bone (frontal : nasal ratio), the characteristics of the rattle-quills, and colour of the rump (Table 46). D. C. D. Happold

Table 46. Characters of H. cristata and H. africaeaustralis (mostly after Corbet & Jones 1965). Character

H. cristata

H. africaeaustralis

Length of rattle-quills Diameter of rattle-quills Colour of nuchal crest Colour of rump Relative length (%) of frontal bone to nasal bone Relative length (%) of nasal bone to occipital–nasal length

5 cm or less 2–5 mm Mostly dark Black 23–38%, i.e. nasal bone very long 58–68%

6 cm or more 5–7 mm Mostly white White 49–68%, i.e. nasal bone not especially long 51–58%

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Family HYSTRICIDAE

Hystrix africaeaustralis CAPE CRESTED PORCUPINE (CAPE PORCUPINE) Fr. Porc-épique de l’Afrique du Sud; Ger. Südafrikanisches Stachelschwein Hystrix africaeaustralis Peters, 1852. Reise nach Mossambique, Saügethiere, p. 170. Near Tette, Querimba coast, Mozambique (exact locality uncertain, but ca. 10° 30' to 12° 00' S, 40° 30' E, at sea level).

Taxonomy Synonyms: capensis, prittwitzi, stegmanni, zuluensis. Subspecies: none. Chromosome number: 2n = 66, NF = 114 (George & Weir 1974). Description Extremely large rodent, the largest in Africa (together with H. cristata), with extremely long stiff hairs and spines (quills).Very similar in external appearance to H. cristata. Dorsal pelage with long stiff black or white hairs anteriorly, and with long smooth quills on midback and rump. Quills with alternating wide black and narrow white bands (usually four or five of each), ending in long white pointed tip; length varies, maximum 30 cm on rump. Nuchal crest (on neck and shoulders) formed of long wiry hairs, up to 45 cm, mostly white with black base. Crest and quills erectile when animal is frightened or threatened. Head, neck and limbs with coarse dark bristles (up to 50 mm). Mid-line of rump white. Head rather broad with short muzzle, swollen nasal region and long dark vibrissae. Eyes small and dark. Ears short, darkly pigmented. Limbs with short black hairs, relatively short and broad; five digits on each foot (Digit 1 of forefoot greatly reduced), each with claws. Tail very short, covered with short weak quills, usually invisible beneath quills of rump; some tail quills modified to form ‘rattle-quills’ or ‘wine-glass quills’ (ca. 6 cm long, 5–7 mm diameter), which rattle when tail is shaken. Skull large and rounded. Upper incisor teeth smooth; cheekteeth with complex folds of enamel and dentine; nasal bones long, 51–58% of occipito-nasal length, wide, extending posteriorly almost to level of anterior end of orbit; frontal-nasal ratio 49–68%. Nipples: 2 or 3 + 0 = 4 or 6. Geographic Variation None recorded. Similar Species H. cristata. Nasal bones >57% of occipital-nasal length; frontal-nasal ratio 23–38%; mid-line of rump black or mottled; western and central Africa; sympatric with H. africaeaustralis in parts of S Uganda, S Kenya and Tanzania. Distribution Endemic to Africa.Widespread throughout southern Africa in four biotic zones (Zambezian Woodland, South-West Arid, Highveld and Coastal Forest Mosaic BZs). Recorded from South Africa northwards to Angola, S DR Congo,Tanzania, S Kenya and S Uganda. Parapatric with H. cristata in East Africa. Probably not present on Zanzibar I. (contra Kingdon 1974; see H. cristata). Habitat Savanna, semi-deserts and forested areas, but not in swamps, moist forests and barren deserts. Common in rocky hill country; shows preference for Burkea savanna (see below). Also occurs in farmlands and forest plantations. Abundance Relatively common in suitable habitats, although rarely seen because of nocturnal habits. Many records are based on the presence of discarded quills.

Hystrix africaeaustralis

Adaptations Nocturnal and terrestrial. Rests during daytime in caves, natural crevices and burrows. Cape Crested Porcupines are good diggers and may excavate their own burrows or may take over burrows made by other animals (e.g. Aardvarks). Burrows may be complex with several chambers. The gait is a either a shuffling plantigrade walk or (if frightened or in danger) a clumsy gallop. In South Africa, Cape Crested Porcupines may have a significant effect on their environment due to their habit of chewing the bark and roots of many species of trees, and digging for food.They are especially fond of the bark of Burkea africana and Dombeya rotundifolia and preferentially chew on these species (De Villiers & Van Aarde 1994); by removing the bark, trees are more susceptible to fire and may be killed during subsequent savanna fires. Chewing bark also results in coppicing and a reduction in the number of trees reaching maturity (Yeaton 1988).The combination of fire and Cape Crested Porcupines has a strong effect on the species composition and structure of Burkea savanna, and helps to maintain the mosaic of grasslands and woodland patches. Cape Crested Porcupines rarely feed on the bark of Acacia trees and do not prevent the development of mature Acacia woodlands. As a result, Acacia woodland has the potential to spread into regions where Burkea woodlands have been partly destroyed. Digging for food disturbs the soil and increases the chances of seed germination; seedling density and diversity may be higher where porcupines (as well as Aardvarks Oryceropus afer and Bat-eared Foxes Otocyon megalotis) have been digging than on flat hard soil surfaces (Dean & Milton 1991). Cape Crested Porcupines have a lower metabolic rate than expected for their size, and can maintain a constant body temperature (Tb) when

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Hystrix africaeaustralis

ambient temperature (Ta) ranges between 13 °C and 30 °C. When Ta is low,Tb is maintained at about 37 °C mostly by increasing metabolic rate and decreasing heat loss.When Ta is 30 °C, there is an increase in ventilation rate and loss of water through the lungs, which prevents body temperature from rising to a dangerously high level. If Ta is 37 °C – a condition not normally experienced by free-living porcupines because they are nocturnal – hyperthermia occurs. Low metabolic levels, efficient heat loss when ambient temperature is high and ability to regulate body temperature over a wide range of conditions, enable these porcupines to live in a wide variety of habitats (Haim et al. 1990a, b). Foraging and Food Omnivorous. Feeds primarily on roots, bark, bulbs, tubers, berries and other fruits, and shoots of herbs (De Graaff 1981, De Villiers et al. 1994). Cape Crested Porcupines forage above ground, digging for roots and bulbs. They may cause damage to crops and forestry plantations. In captivity, they eat more during winter (10 °C and short days) than during summer; an increase in food consumption (and hence energy production), together with a decrease in heat loss, appear to be mechanisms that allow porcupines to be active during cold weather (Haim et al. 1990b). Food consumption is correspondingly reduced during summer conditions. Social and Reproductive Behaviour Live in monogamous pairs, usually with their young. Family groups live together in burrows, although group may split up while foraging at night. Size and the duration of family group seem to depend on opportunities for young to disperse when they become mature. Family groups may, on occasion, be large – up to 14 individuals have been found in a single burrow – and composed of two (or more?) pairs with young (Van Aarde 1987a). Within a family group, only the monogamous pair reproduces. Offspring disperse when mature provided there are areas where territories may be established and where food is not limiting. If dispersal opportunities are limited, offspring may stay with parents, forming large family groups, but reproductive suppression prevents offspring from being reproductively active (Corbet & Van Aarde 1996). Home-ranges are large. In Burkea savanna in South Africa (Corbet & Van Aarde 1996), mean home-range area in summer was 215 ha, although only about 80 ha of the range (the ‘core area’) was used extensively. Home-ranges of different individuals overlapped, especially during winter, although the core area was used exclusively by a single individual. Hence individuals are probably territorial with a small territory, advertised by scent marking, within a larger homerange. Winter home-ranges and territories are smaller than those in summer (e.g. 142 ha and 55 ha, respectively). Each territory has 1–3 burrow systems (Van Aarde 1987b). Porcupines that live and feed in crop areas have larger ranges than those living in natural savanna, and without any seasonal differences in area. Porcupines communicate with each other with various piping calls and pig-like grunts. Members of a monogamous pair indulge in daily bonding behaviour (grooming, etc.), and " conceives only after living with her partner for at least 90–100 days. Parental care of young is well developed. In the burrow, mother suckles young while in a crouching position. Male lives in the burrow with " and young and, later, escorts them on foraging excursions and protects them from predators (Van Aarde 1997). Family group remains intact until young disperse at adulthood.

Reproduction and Population Structure Breeding season varies according to locality. In South Africa (30° S), in the wild, births occur mostly in spring and summer, from Aug to Mar with a peak in Jan (Van Aarde 1987a). Most "" more than 24 months of age are reproductively active (88–95%) during the breeding season.Younger "" (12–24 months) are less reproductively active (63–88%), and "" aged less than 12 months rarely breed. In the drier regions of the Karoo in South Africa, births coincide with peaks in rainfall (Skinner et al. 1984). Times of reproductive activity in northern parts of geographic range are uncertain, although there is some evidence that births may occur in all months of the year. Mean oestrus cycle: ca. 35 days (Weir 1974). In South Africa, captive "" are polyoestrous, with most "" cycling every 28–36 days (Van Aarde 1985). Gestation: 93–94 days. In captivity (in South Africa), litter-size 1.5 (1–3); of 165 litters, 58% had a single young, 32% had twins and 9% had triplets (Van Aarde 1985).Weight at birth 300–440 g. Each young (whether singletons or twins) weighs ca. 2% of maternal weight; overall litter weight (young, placenta, etc.) weighs ca. 10% of maternal weight (Weir 1974). Mammary glands are situated on the side of the thorax. Twins usually suck from opposite nipples. The average length of lactation is 101 days, but may continue for 163 days. Mean litter interval of captive "" is 385 (269–500) days; thus mothers give birth to only one litter/year (Van Aarde 1985). Young are precocious (although relatively small) at birth, with eyes open and with soft spines and soft quills on the back. They remain in the burrow for 7–9 weeks, a much longer period of time compared with other rodents, emerging for the first time when the quills have hardened.This behaviour (as well as huddling with siblings and parents) probably conserves energy, which can be channelled into growth rather than being used for activity above ground, and provides protection before the quills are fully developed. Growth in body weight is linear until a weight of 11–12 kg is attained at about 52 weeks; thereafter, there is a slow increase in weight to the full adult size of 12–18 kg (Van Aarde 1987a). Sexual maturity occurs when 12–24 months of age. The approximate age of an individual can be determined by tooth wear until adult weight is attained; thereafter age cannot be determined. Little is known about age structure of populations. In N South Africa, age structure varied markedly during a 2-year study (Van Aarde 1987b). When categorized into four age categories (24 months), the proportions of each age category during one of the years were 20– 40%, 10–25%, 57% of occipito-nasal length, wide, extending posteriorly almost to level of posterior margin of orbit; frontal : nasal ratio 23–38%. Nipples: 2 or 3 + 0 = 4 or 6.

Hystrix cristata

Geographic Variation Populations show differences in occipito-nasal length, zygomatic width, length of nasals and in overall size. North African (and Italian) individuals are the smallest in size, those from West Africa are intermediate, and those from eastern African (galeata) are the largest. However, there is considerable overlap, and differences are probably clinal (Corbet & Jones 1965). In West Africa, Rosevear (1969) refers to two races (subspecies) that differ in size: the largest is senegalica from the savanna woodlands (total length about 900 mm, HF 120 mm, GLS over 140 mm, and P4–M3 over 32 mm), and the smallest is aerula from the semi-desert (TL about 700 mm, HF 80–90 mm, GLS under 140 mm, and P4–M3 30 mm or less).

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Similar Species H. africaeaustralis. Nasal bones 51–58% of occipito-nasal length; frontal-nasal ratio 49–68%; mid-line of rump white; southern Africa; parapatric with H. cristata in parts of S Uganda, S Kenya and Tanzania. Distribution Widespread in Mediterranean Coastal BZ, Sudan Savanna and Guinea Savanna BZs, Northern Rainforest–Savanna Mosaic, and Afromontane–Afroalpine BZ of Ethiopia. Disjunct distribution in parts of Sahel Savanna and Sahara Arid BZs. Recorded from coastal regions and parts of Atlas mountains of Morocco, Algeria and Tunisia; coastal areas of Libya. Probably extinct in Egypt (Osborn & Helmy 1980). Isolated populations in semi-desert habitats in Aïr (Niger) and Adrar des Iforas (Niger). Widespread throughout West Africa from Senegal to Cameroon, and in NE DR Congo, Rwanda, Ethiopia, Uganda, Kenya and N Tanzania, Zanzibar I. Probably present in S Chad and Central African Republic. In Ethiopia, occurs from sea level to about 3550 m. Also occurs (as an introduced species) in Italy, Sicily, Albania and N Greece (Woods & Kirkpatrick 2005). Habitat Semi-desert, woodland and grassland savannas, especially where rocks and caves are present. In Algeria and Morocco, lives in forested hills and steppes, but not in the Sahara Desert. Appears to be very tolerant of a wide range of habitats and climates, including warm coastal scrub, dry semi-desert and cold grasslands on mountains. In Ethiopia, may be attracted to irrigated large-scale farmlands, where they can become pests on crops (Yalden et al. 1976). Abundance Uncertain because rarely seen; thought to be quite common in suitable habitats in most parts of range. Many records are based on the presence of discarded quills. Adaptations Nocturnal, although in captivity may be active in daylight. Terrestrial. During the day, these crested porcupines rest in caves, holes under trees or made by other animals, and in rocky crevices. They do not dig their own burrows. Locomotion is a walk or slow trot, and because of their large size, they are unable to climb. When frightened or threatened, the crest and quills can be erected (making the animal look much bigger than it really is) and the quills are rattled. If in danger, the animal moves sideways or backwards with the pointed tips of the erect quills facing the source of danger, and stamps its feet (Ewer 1968). If really provoked by a potential predator, an individual charges backwards forcing some of its quills into the predator; the quills are easily detached and may stick (like arrows) in the predator. Quite severe wounds can be caused by these quills. This behaviour, and the sharpness of the quills, provides the porcupine with a very effective defence mechanism (even against potential predators such as Lions Panthera leo). There are many reports of porcupines (this species and probably also Cape Crested Porcupines) gathering and chewing on bones, and dens being littered with bones.These habits are thought to be associated with wearing and sharpening the incisor teeth, perhaps with the added benefit of providing an additional source of calcium and minerals (Kingdon 1974).

Foraging and Food Herbivorous. Principal foods are fruits, roots, bulbs and bark. Cassava, sweet potatoes and groundnuts are eaten in savanna farmlands. Social and Reproductive Behaviour Crested Porcupines are social and gregarious. Secretions from anal glands are used to mark home-ranges and to indicate an individual’s presence, and vocal sounds are used (as in many species of hystricomorph rodents) for male–female interactions, to warn conspecifics of danger and during aggressive encounters. During courtship, ! approaches " using a ‘bipedal approach’ gait, and he also grooms the ". Females solicit copulation by a ‘tail-up rump’ display (Mohr 1965, Kleiman 1974). Several individuals may rest together in a burrow (Delany 1975). Reproduction and Population Structure Gestation: 112 days. Litter-size: 2 (1–4).Weight of young at birth ca. 1000 g. Ratio of litter weight/maternal weight is 10% – a low percentage compared with most hystricomorph rodents (6–60%) (Weir 1974).Young born with eyes open and soft spines (Rosevear 1969). Mother suckles while sitting because nipples are placed on the side of thorax. Males assist with retrieving and grooming young, and will rest with young in the burrow (Mohr 1965).Young weaned at 16 weeks of age. Predators, Parasites and Diseases North African Crested Porcupines have few predators because of their defensive behaviour (see above). Humans may hunt them for food, but they are not a major source of ‘bushmeat’ (cf. Atherurus africanus, Thryonomys swinderianus and duikers; for details, see references in profiles for these species). Two species of tsetse flies (Glossina submorsitans and G. tachinoides), which transmit sleeping sickness to humans, are recorded as feeding on the blood of porcupines. Conservation IUCN Category: Least Concern. North African Crested Porcupines are uncommon and populations are scattered (see above) and hence may be in need of protection. Measurements Hystrix cristata HB: 650–850 mm T: 120–170 mm HF: ca. 95 mm E: ca. 40 mm WT: ca. 20 kg GLS: 158 (152–170) mm GWS: 83 (81–88) mm P4–M3: 33.6 (32.3–34.4) mm Body measurements: Morocco (H. c. cristata; Aulagnier & Thévenot 1986; no sample sizes given) Skull measurements: West African savanna (H. c. senegalica; Rosevear 1969; no sample sizes given) Key References

Corbet & Jones 1965; Rosevear 1969. D. C. D. Happold 679

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Family PETROMURIDAE

Family PETROMURIDAE NOKI (DASSIE RAT)

Petromuridae Wood, 1955. J. Mamm. 3: 184. Petromus (1 species)

Noki

p. 681

The family contains only a single species, Petromus typicus. It is restricted to Namaqualand (South Africa), Namibia and extreme SW Angola, occurring only on mountains and rocky outcrops of the semi-arid western escarpment and adjoining areas of the Namib Desert. The single species is unusual in its external appearance, looking like a mixture of a rat and a squirrel. It is about 200 mm long, with brownish or brownish-grey coarse pelage, and longish tail covered with long dark bristle-like hairs.The head is relatively large, rather flattened with pointed muzzle and long vibrissae. Eyes large; ears moderate and rounded, not protruding above the line of the head. Limbs are short, and feet are broad and naked with well-developed pads; forefoot with four digits (Digit 1 rudimentary); hind-foot with five digits, all with short sharp claws. The ribs are particularly flexible so the body can be pressed flat (when under boulders and rock slabs) without injury. Pectoral nipples are situated laterally behind the shoulders (as in Cane Rats). Size categories of species in the family (based on mean head and body length) are given in the order Rodentia profile. The skull exhibits typical hystricomorph characters as well as those associated with a rupiculous life-style: infraorbital foramen enlarged,

Figure 111. Skull and mandible of Petromus typicus (BMNH 28.9.11.377). This specimen is from a young animal and shows infoldings on the labial side of the cheekteeth. See family profile for details.

jugal bone of zygomatic arch enlarged dorsoventrally, cranium flattened dorsoventrally (height above M1 about 43% of zygomatic width), rostrum narrow (especially in relation to the wide orbital area, zygomatic arches and braincase), and without an interorbital constriction. Dental formula: I 1/1, C 0/0, P 1/1, M 3/3 = 20. Upper incisors narrow, not grooved, opisthodont, yellowish. Anterior palatal foramina wide at both ends (similar in width to the upper molars), reaching posteriorly to between premolars; septum between each foramen very narrow. Cheekteeth show a unique structure: four cheekteeth (one premolar, three molars), hypsodont and four-rooted. Each upper cheektooth has deep infolding on the lingual side giving the impression that it consists of two sections. Similar infoldings occur on the labial side but, because of increased wear on this side, they are less obvious in older animals. Each lower cheektooth has deep infoldings on the labial side, and most wear on the lingual side.Auditory bullae considerably inflated, with a well-developed paraoccipital process on the posterior side, which does not project below the level of the bullae. Mandibles very wide posteriorly, with a distinct ridge on the lower outer side, stretching from below the first molar to the narrow angular process (Figure 111). Nokis are adapted for desert and semi-desert life on rocky habitats. They live in family groups amongst boulders of outcrops and mountainsides, or amongst rock slabs in broken terrain.They are diurnal, but less active during the warmer hours of the day, and they forage on grass and leaves of shrubs and trees up to 20 m away from the protection of rocks. The phylogenetic relationship of the Petromuridae to other hystricognaths is uncertain. Simpson (1945) grouped Petromuridae with the Octodontoidea, a superfamily proposed by him.Wood (1955) re-established the suborder name Hystricogmorpha for Old World taxa of the hystricognaths and proposed the superfamily name Thryonomyoidea (for the Cane Rats and Dassie Rat) as well as the use of Petromuridae to replace Petromyidae (see Woods & Kirkpatrick 2005). Lavocat (1974) supports the differentiation between Hystricognathi of the Old and New World by using the terms Phiomorpha and Caviomorpha respectively, but prefers to group them together into a higher clade Hystricognathi (suborder). Mess (1999), using the analysis of molecular data (Catzeflis et al. 1995), the structure of the rostrum (Ade 1998) and of the ethmoid- and orbital regions (Mess 1997), grouped Petromus, Hystrix and Thryonomys under the Hystricoidea – a name proposed by Gill in 1874. Patterson & Wood (1982) split the Petromuridae in three subfamilies: the Petromurinae for the extant Petromus, and the Phiomyinae and Diamantomyinae for extinct species. At present, there is one genus and one species, Petromus typicus. C. G. Coetzee

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Petromus typicus

Genus Petromus Noki (Dassie Rat) Petromus A. Smith, 1831. S. Afr. Quart. J. 1(5): 10. Type species: Petromus typicus A. Smith, 1831.

A monotypic genus occurring only in the South-West Arid BZ. Characters of the genus are given in the family profile above. The single species is Petromus typicus. C. G. Coetzee

Petromus typicus.

Petromus typicus NOKI (DASSIE RAT) Fr. Rat des rochers du Namibie; Ger. Felsenratte Petromus typicus A. Smith, 1831. S. Afr. Quart. J., ser. 1, 5: 11. Mountains towards the mouth of the Orange River, Little Namaqualand, Northern Cape Province, South Africa.

Taxonomy Originally described in the genus Petromus, but in 1834 Smith referred to the genus as Petromys. This mis-spelling prevailed for some time and influenced the spelling of the higher categories – Petromyidae (e.g. Roberts 1951) and Petromyinae (Ellerman 1940). The erroneous spelling was used also in the descriptions of 11 of the 13 described forms (based mainly on coat colour). Synonyms: ausensis, barbiensis, cinnamomeus, coetzeei, cunealis, greeni, guinasensis, karasensis, kobosensis, majoriae, namaquensis, pallidior, tropicalis, windhoekensis. Subspecies: none recognized here (but see Geographical Variation). Chromosome number: not known. Description Large, squirrel-like diurnal rodent with longish pelage and hairy tail. (The term ‘large’ is ambivalent here; the species is large by murid or sciurid standards, but it is the smallest of non-fossorial hystricomorpha rodents; see Order profile.) Dorsal pelage dark blackish-brown, dark grey or pale buffy-yellow (depending on locality; see below); hairs pale grey at base. Ventral pelage slightly paler than dorsal pelage. Head dorsoventrally flattened, similar in colour to dorsal pelage, paler on nose, lips and around the eyes. Vibrissae long. Eyes large. Ears slate-gray to black, sparsely covered with extremely short hair, small, oval, not protruding above the line of the head. Limbs short; feet broad, well covered with hair on upper surface, naked on undersurface. Forefoot with four digits (Digit 1 rudimentary), three plantar pads at the base of digits and two on palm. Hindfoot with five digits, three plantar pads at base of the toes and only one pad on sole. All digits with short sharp claws. Tail long (ca. 85% of HB), densely covered with long hairs (but not as long as in a squirrel), similar in colour to rump at base, black on terminal three-quarters; tail normally rests on (or is held close to) the ground, not squirrellike over the body. The scrotum is not conspicuous. Skull: see family profile. Nipples: 2 + 1 = 6; anterior nipples placed laterally behind shoulders; inguinal nipples often absent.

Geographic Variation Although no subspecies are recognized here, three groups may be distinguished, based on the colour of the dorsal pelage, which varies from dark brown in the south to pale brown in the north, and pale to dark grey towards the higher rainfall area of the Otavi Mts complex, leading to south–north and west– east coat clines in colour. The geographical groups may be linked to named forms, as follows: (a) Southern buffy-brown group (Namaqualand, along the Orange R., and S Namibia) corresponding to the nominate form (typicus) and to ausensis, barbiensis, cinnamomeus, karasensis and namaquensis. (b) Pale brown group (C and NW Namibia to SW Angola) corresponding to coetzeei, greeni, koboscensis, marjoriae, pallidor, tropicalis and windhoekensis. (c) Grey group (Otavi Mountain area towards the eastern Kaokoveld and northwards to the Kunene R. valley below the Rua Cana Falls and east of the Baines Mts) corresponding to cunealis and guinasensis. However, preliminary cranial measurements do not support these divisions (C. G. Coetzee & C. Chimimba unpubl.). Similar Species Procavia spp. Similar in appearance, especially when the Noki has lost its tail, as is the case in about 10% of animals; sympatric and often syntopic. Xerus princeps. Larger; comparatively longer HB; tail with long bushy hairs; may be sympatric and syntopic. Graphiurus spp. Smaller HB; pelage woolly, nocturnal; upper incisors form an inverted V-shape. Distribution Endemic to Africa. South-West Arid BZ (Namib Desert). Recorded in rocky habitats in extreme SW Angola, W Namibia and extreme NW South Africa where mean annual rainfall 681

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Family PETROMURIDAE

Petromus typicus

is ca. 50–400 mm. Found in canyons of the Namib where the mean annual rainfall can be as low as 35 mm/year. In Angola, limited to the outcrops and broken terrain on the eastern side of the Namib, approximately as far north as 16° S, 12° E and along the Kunene R. inland to the Rua Cana Falls. Habitat Occurs amongst granite, schist slabs and sedimentary rocks; and amongst the boulders of outcrops, canyons and mountain slopes where rock crevices provide shelter and nesting sites. May also occur amongst boulders and rock accumulations that are some distance away from mountains and outcrops. Normally absent on steep, rather bare rock faces although such rock faces may be used during the day for basking. Habitats with deciduous and evergreen trees and perennial shrubs provide a more stable food resource on outcrops and mountains of the Namib (Rathbun & Rathbun 2005). Also recorded in marginal habitats where annual rainfall is 80% during 10 months of the year, and in N Ghana, pregnancy rate was 26–95%, with >80% during the four months of the wet season (May–Aug) (Asibey 1974b). No information available on months when young are born, but seasonality of birth rate is probably related to seasonality of rainfall and grass growth. Information from elsewhere is inconclusive: young recorded in Jun and Aug in Botswana (Smithers 1971), and in Aug and Nov in Zimbabwe but information is lacking for other months (Smithers & Wilson 1979). Gestation 155 days (137–172), n = 33 litters (Asibey 1974b). Postpartum oestrus likely. Embryo number: 3 (1–5); n = 18 females (South Africa; van der Merwe 1999); 3.8 (2–5), n = 6 litters (van der Merwe & van Zyl 2001); 4–6, mode 4, n = 480 litters (Ghana; Asibey 1974b). Some evidence of embryo reabsorption (Asibey 1974b). Mean weight of young at birth: 128 (75–190) g (n = 9 litters). Total litter weight at birth is ca. 15–20% of maternal weight (n = 112 litters), and there is no relationship between litter weight and weight of the mother (Asibey 1981). Great variation in weight of young within and between litters. Mean litter-size varies according to age of mother: 3.1 for primiparous "" and 3.9 for parous mothers (Asibey 1974b). At birth, young are fully furred with eyes open, and can follow the mother within an hour of birth. In captivity, growth rates and adult weights greater for "" than for !!; "" reach adult size at ca. Day 300, and !! at ca. Day 390 (van der Merwe & van Zyl 2001). Sexual maturity is attained at about seven months and a " gives birth to her first litter when about one year old. Females probably produce two litters each year (Booth 1960, Asibey 1974b). In captivity, in South Africa, "" may have two litters per year (van der Merwe & van Zyl 2001). Histological changes during the oestrous cycle are described by Adjanohoun (1992). Longevity up to four years in captivity. Predators, Parasites and Diseases Leopards, servals, hunting dogs, eagles, eagle-owls and pythons are potential natural predators (De Graaff 1981). Greater Cane Rats are vigorously hunted by humans

because of their succulent flesh. Carcasses are seen for sale as‘bushmeat’ in many parts of Africa, especially in the Rainforest BZ (see also Atherurus africanus). Greater Cane Rats are hunted with dogs and by burning savanna grasses, and are caught by snares. In Bendel State, S Nigeria, carcasses of cane rats formed 20–34% of all bushmeat for sale, and were either the most abundant or second most abundant species in local markets (Martin 1983, Anadu et al. 1988). In Accra, Ghana, about 110,000 kg of Greater Cane Rats were traded during a 12-month period in 1970–71 (Asibey 1974a) – equivalent to 40,000–55,000 individuals. In less suitable habitats, e.g. Equatorial Guinea, they form only a small proportion of ‘bushmeat’ (Juste et al. 1995). The meat of Greater Cane Rats has a higher percentage of protein and less fat per unit weight than does rabbit and chicken, and is also rich in calcium and phosphorus (Jori et al. 1995). Because of the desirability of these animals for food, studies in several African countries are investigating the possibility of domesticating and farming them commercially (Ajayi & Tewe 1980, Hardouin 1995, Jori et al. 1995). Many species of ticks have been recorded (De Graaff 1981, Aeshlimann 1967), as well as gastrointestinal cestode and nematode worms (De Graaff 1981). Conservation IUCN Category: Least Concern. In spite of the very high hunting pressure on Greater Cane Rats, in some parts of the geographic range, numbers do not appear to be threatened at present. However, there is some evidence that population numbers are falling near large urban centres. Measurements Thryonomys swinderianus TL (!!): 715 (670–792) mm, n = 5 TL (""): 666 (654–670) mm, n = 3 T (!!): 188 (180–192) mm, n = 6 T (""): 183 (165–195) mm, n = 3 HF (!!): 94 (80–100) mm, n = 6 HF (""): 89 (88–90) mm, n = 3 E (!!): 33 (30–35) mm, n = 6 E (""): 35 (34–45) mm n = 3 WT (!!): 4.5 (3.2–5.2) kg, n = 6 WT (""): 3.6 (3.4–3.8) kg, n = 3 GLS: 90.6 (86.5–95.1) mm, n = 4 GWS: 58.0 (55.6–61.3) mm, n = 4 P4–M3: 18.8 (18.1–19.5) mm, n = 4 Body measurements: southern Africa (Smithers 1983) Skull measurements: Nigeria (forest; Rosevear 1969) Key References Smithers 1983.

Asibey 1974a, b; Ewer 1969; De Graaff 1981; D. C. D. Happold

Thryonomys gregorianus.

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Family MYOCASTORIDAE

Family MYOCASTORIDAE COYPU

Myocastoridae Ameghino, 1904. Anales Soc. Cient. Argentina 56–58: 103.

This family of South American aquatic rodents contains only a single genus and single species, Myocastor coypus. The family belongs to the suborder Hystricomorpha (together with the African endemic families Bathyergidae, Hystricidae, Petromyidae and Thryonomyidae). This species is not indigenous to Africa, and only one introduced

population (now feral) is known to exist in Africa; therefore details of the family and genus are not given here (see Woods et al. 1992 [and references therein] andWoods & Kirkpatrick [2005] for further details). D. C. D. Happold

Genus Myocastor Coypu Myocaster Kerr, 1792. In: Linnaeus, Anim. Kingdom, p. 23. Type species: Mus coypus Molina, 1782.

The genus is monotypic. Further information is given in the family and species profiles.

Myocastor coypus COYPU (NUTRIA) Fr. Ragondin; Ger. Nutria Myocastor coypus (Molina, 1782). Sagg. Stor. Nat. Chile, p. 287. Bio Maipo, Chile.

Taxonomy No information available for African populations. See Woods & Kirkpatrick (2005) for details of synonyms and subspecies in natural geographic range of species. Unless otherwise stated, this profile refers to the introduced population in Kenya – the only feral population known on the African continent. Synonyms: eight (world-wide). Subspecies: none in Africa. Chromosome number: 2n = 42, FN = 76. Description Extremely large shaggy rodent with dark lustrous pelage, long tail and webbed hindfeet. Pelage thick with long coarse guard hairs (dull or shiny) and dense underfur. Dorsal pelage brown, tending to blackish-brown in some individuals; guard hairs thin and long (up to 50 mm), pale or dark brown, usually with pale brown band(s) below tip.Ventral pelage similar to dorsal pelage in colour and texture. Underfur dark brown to black; dense and woolly. Head broad and thickset, similar in colour to dorsal pelage; muzzle and chin with some hairs pale or white or with white tip. Vibrissae very long and coarse. Ears dark, small and rounded. Fore- and hindlimbs short with dark brown or black hairs. Forefeet with five digits; Digit 1 short, Digits 2–5 long, each with long claw and without webbing between digits. Hindfeet long with webbing (skin) between digits, especially between Digits 1–2, 2–3 and 3–4; long claw on all digits.Tail short to long (ca. 72% of HB), with scales, sparsely covered with short dark brown hairs. Skull large and strong; zygomatic arches deep; infraorbital foramen very large; large pointed paraoccipital processes; mandible deep with angular process extending far posteriorly to condylar process and paraoccipital process; dental formula: I 1/1, C 0/0, P 1/1, M 3/3 = 20; incisor teeth smooth, without grooves, usually orange in colour on outer surface; cheekteeth flattened, converge anteriorly and camber outwards; each cheektooth wth complex foldings of dentine and enamel on outer and inner surfaces (Figure 113) . Nipples: 4–5 pairs situated dorsolaterally.

Figure 113. Skull and mandible of Myocastor coypus (BMNH 60.1948).

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Family MYOCASTORIDAE

Geographic Variation None recorded in Africa. Similar Species Thryonomys swinderianus. HB of similar size, but shorter T (mean 183– 188 mm); without webbing on hindfeet; smaller skull (GLS, GWS and P4–M3); terrestrial or semi-aquatic. Thryonomys gregorianus. Smaller in size (mean HB: ca. 375 mm), with much shorter tail (mean 132–144 mm; ca 38% of HB); without webbing on hindfeet; skull smaller; terrestrial or semi-aquatic. Distribution Introduced. In Africa, recorded only from aquatic habitats in C Kenya. Introduced (as captive animals) at Nanyuki in ca. 1947 (see also below). Later (ca. 1950), some animals were released (or escaped) and gradually spread throughout rivers, dams and swamps of the Central Highlands east of the Rift Valley (Ian Parker in litt.), as well into the Rift Valley at L. Naivasha (ca. 1960). They have been recorded, at various times, from the Ewaso Nyiro and Ewaso Narok rivers on the Laikipia Plateau, Ol Pejeta Conservancy and Mt Kenya Safari Club near Nanyuki, the Ark waterhole in the eastern Aberdare Ranges, near Sagana (possibly), L. Naivasha, L. Ol-Bolossat and near Kiserian (south of Nairobi) (data from many local sources). They have not been recorded from the alkaline lakes Elmenteita and Nakuru in the Rift Valley. Nor, as yet, have they been recorded from the highlands west of the Rift Valley (where the climate is similar to that east of the valley) and where they might be expected to occur. Map not given. De Vos (1965) recorded that ‘coypu have been established in the wild in Zambia’ (see also Haltenorth & Diller [1980], Lever [1985] and Long [2003]). However, Ansell (1978), commenting on De Vos’ statement, wrote: ‘I am unaware of the basis for this assertion, but even if correct no more appears to have been heard of them.’ Local sources (in litt. 2005/2006) state that there is no evidence for the presence of coypu in Zambia. Haltenorth & Diller (1980) record that ‘from 1960 [coypu] has gone feral in the coastal swamps of Tanzania’ (see their map showing small range in the extreme NE corner of the country). Lever (1985) and Long (2003), both following Haltenorth and Diller (1980), also record the presence of Coypu in Tanzania. However, local sources (2005, with records back to the 1960s) have no knowledge of Coypu in Tanzania. On the basis of the present evidence, it appears that Coypu do not occur in the wild in Zambia or Tanzania. Lever (1985) records, without detail, that Coypu have been farmed in Zimbabwe and South Africa, and have not become feral in these countries. Other authors have also commented on the presence of Coypu in southern Africa. Aliev (1967) records Coypu in Botswana and Zimbabwe (as dots on a map) without comment or reference, and Carter & Leonard (2002) show the presence of Coypu in Zambia, Zimbabwe and Botswana (map, mostly following Aliev 1967) but also comment (as pers. comm. J. du Toit) that the species is not feral anywhere in southern Africa. Local sources in Zimbabwe and South Africa (M. van der Merwe, F. P. D. Cotterill, D. Spears pers. comm. 2007) record that the species is not present (farmed or feral) in these countries. The overall evidence suggests that, at the present time, Coypu are feral only in Kenya. The natural distribution of the species is South America, but it has been widely introduced into North America, Europe and N Asia (Lever 1985, Long 2003,Woods & Kilpatrick 2005). (Map not given.)

Habitat Rivers, lakes, streams and swamps. Coypu are able to disperse from one aquatic habitat to another when conditions are favourable. Abundance In Kenya, distribution is patchy. Very numerous in some habitats, e.g. the rivers and dams on the Laikipia plateau (N. Gregory in litt.). One dam at Ol Pejeta Conservancy contained eight individuals (ca. 4/ha; Butynski pers. comm.). Presence and abundance varies seasonally and annually; local populations may become extinct, but recolonization of a habitat may occur when conditions are suitable. Remarks Aquatic and nocturnal. There are no detailed studies on the species in Kenya; the following remarks refer to populations extralimital to Africa. The webbed hindfeet are adaptations for swimming, and the thick water-repellent pelage helps to maintain a more or less constant core body temperature, especially when the water is cool or cold. Other adaptations for aquatic life include the ability to stay submerged for at least 10 minutes, and to preferentially maintain blood flow, during a dive, to the brain and heart while restricting blood flow to the muscles, intestines and kidney. Vegetarian, feeding on a large variety of aquatic and terrestrial herbs, stems and roots. Often gregarious. Litter-size: usually 3–6 (1–12); young precocial at birth; weaned at Week 8; attain adult weight at ca. 16–18 months (Britain; Southern 1964); maximum longevity in the wild probably 5–6 years. See Woods et al. (1992) for a review. Conservation IUCN Category: Least Concern (worldwide). Introduced into Kenya to provide pelts (skins) for making coats (see above). Population numbers in Kenya are kept in check by many predators (N. Gregory in litt.). Introduction of Coypu to L. Naivasha, together with the introduction of crayfish and Floating Water-fern Salvinia molesta in the 1960s and 1970s, has had dramatic adverse effects on the indigenous aquatic flora (Harper et al. 1990); additionally unwise water usage and inappropriate land management of the water catchment of the lake are having deterimental effects on the lake ecosystem. Geographic range is unlikely to expand in Kenya to any great extent because aquatic habitats in a cool climate are required for the survival of the species. Measurements Myocastor coypus HB: 521 (472–575) mm T: 375 (340–405) mm HF: 135 (120–150) mm E: 27 (25–30) mm WT (!!): ca. 6.7 kg WT (""): ca. 6.36 kg GLS: 114.2 (102–106) mm GWS: 68.1 (60–76) mm P4–M3: 27.8 (22.5–29.3) mm Locality not stated, presumably North America; sample sizes not recorded (Woods et al. 1992) Key References

Woods et al. 1992; Woods & Kilpatrick 2005. D. C. D. Happold

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Order LAGOMORPHA

Order LAGOMORPHA – Hares, Rock-hares, Rabbits and Pikas Lagomorpha Brandt, 1855. Leporidae (5 genera, 13 species)

Hares, Rock-hares, Rabbits

p. 694

The order Lagomorpha contains two extant families (Leporidae, Ochotonidae), 13 extant genera and about 87 species (Hoffmann & Smith 2005). Of these, only the Leporidae, with five genera and 13 species, is represented in Africa. The order was once considered to be a suborder of rodents (because of the superficial similarity of the teeth). The modern view is that the Lagomorpha is an order in its own right. Based on molecular and morphological evidence, the Lagomorpha and Rodentia may be grouped together in the cohort Glires (see for example Scally et al. 2001, Bronner et al. 2003). The order is represented naturally in all continents except Antarctica, South America and Australia. The principal characteristics of the order include: two pairs of upper incisors, the second pair being very small and located behind the first principal pair where they have no cutting function; incisor teeth which grow throughout life and are rooted in the premaxilla bone; no canine teeth; a diastema between the incisors and the cheekteeth (as in rodents and artiodactyls); five or six high-crowned cheekteeth (P1, P2, P3, M1, M2, M3), which may or may not have roots (depending on the family); large caecum; no baculum in the penis; and testes that are anterior to the penis. In size and habitat, members of the two families are rather different. Species of Leporidae are

the largest members of the order (details below), have long narrow upright ears, small fluffy tails, and live in grasslands and scrublands. In contrast, species of Ochotonidae (no longer present in Africa) are small (HB: ca. 125–300 mm, WT: 125–400 g), with small rounded ears close to the head, and most species are associated with rocks and talus (Nowak 1999). All members of the order are terrestrial and vegetarian. The fossil record of the Lagomorpha in Africa is relatively poor and fragmentary (Cooke 1972). The earliest fossils, from the early Miocene (ca. 20 mya), are ochotonids from Kenya, Uganda, Namibia, Morocco and Libya (Erbajeva 1994, Winkler et al. 2005). It is assumed that these ochotonids arrived in Africa from Eurasia. The leporids are first known from Africa in the late Miocene (oldest is 6.5–6.6 mya) of Kenya (Winkler 2002, 2003; Mein & Pickford 2003) and Ethiopia (Haile-Selassie et al. 2004). From this time, leporids have radiated extensively (perhaps due to the expansion of grassland). Ochotonids are last known from Africa in the middle Miocene (14–15 mya; Mein & Pickford 2003, Winkler 2003) and now survive only in Eurasia and western North America. The single family of the Lagomorpha in Africa is the Leporidae. Further details are given in the family profile below. D. C. D. Happold

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Family LEPORIDAE

Family LEPORIDAE HARES, ROCK-HARES AND RABBITS Leporidae Fischer, 1817. Mém. Soc. Imp. Nat. Moscow, 5: 372.

Riverine Rabbit Hares Rabbit Bunyoro Rabbit Rock-hares

Bunolagus (1 species) Lepus (6 species) Oryctolagus (1 species) Poelagus (1 species) Pronolagus (4 species)

a

The family Leporidae occurs widely in the Palaearctic, Oriental and Ethiopian Regions, and certain species have been introduced into parts of South America, Australia and New Zealand, and many oceanic islands. Representatives of the family live in arctic, temperate semitropical, tropical, semi-arid and arid habitats.There are 11 genera and about 61 spp. in the family of which five genera and 13 species occur in Africa (see above); three of these genera are endemic to Africa (Bunolagus, Poelagus, Pronolagus). See Flux & Angermann (1990), Hoffmann (1993) and Hoffmann & Smith (2005) for further details. Species in the Leporidae in Africa are easily recognized by their relatively large size for a ‘small mammal’ (mean HB: 300–600 mm, mean WT: 1–3 kg, according to species), which is larger than the majority of rodents, long (or very long) narrow ears, which project upwards from the head, large eyes, small fluffy tail and thick woolly pelage. Other distinguishing characters of the family include two patches of different-textured pelage – the nuchal patch on the back of the neck and the gular patch on the throat and anterior part of the chest, relatively long limbs (hindlimbs usually longer than forelimbs), four or five digits on each foot, and thick dense hairs on the soles of the feet. The skull is lightly built and arched with moderate restriction between the orbits, prominent supraorbital processes, well-developed thick zygomatic arches, maxilla bone with numerous

p. 696 p. 698 p. 708 p. 710 p. 712

b

c

d e

pf

1

Figure 114. Selected characters of the skull of Leporidae. (a) upper front incisor teeth (frontal view) with deep groove filled with cement, (b) upper front incisor teeth (frontal view) with shallow groove not filled with cement, (c) cross-section of (a) above with the smaller second incisor posteriorly, (d) cross-section of (b) above with the smaller second incisor posteriorly, (e) ventral view of part of skull showing ‘minimum length of hard palate’ and ‘width of mesopterygoid space’. pf = palatal foramen, hp = hard palate, ms = mesopterygoid space, 1 = minimum length of hard palate, 2 = width of mesopterygoid space.

hp

ms

2

Table 48. Genera of Lagomorpha in Africa. Arranged in order of increasing mean length of ear. (n.d. = no data.) Genus Poelagus (1 sp.) Oryctolagus (1 sp.) Pronolagus (4 spp.) a Bunolagus (1 sp.) Lepus (6 spp.) a a b

E (mean) (mm) [E/HF as %]

HB (mean) (mm)

T (mean) (mm)

HF (mean) (mm)

E (mean) (mm) [E/GLS as %]

Width mesoptyergoid space (mm)

Minimum length hard palate (mm)

65 [67%]

415

56

97

64.7 [81%]

6.71

7.95

73 [87%]

368

69

84

73 [98%]

5.32

5.86

74 [81%]–94 [102%]

447–508

65–97

91–100

74 [81%]–94 [116%]

4.8–5.8

6.9–9.5

116 [111%]

429

92

104

116 [146%]

7.54

5.12

88 [92%]–140 [101%] b

452–561

68–126

95–138

88.2 [101%]–140 [139%]

7.5–10

6.0–7.5

For genera with more than one species, values for the smallest and largest species are given. Only L. fagani has a mean ear length of less than 100 mm, and the smallest values for T and HF.

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Family LEPORIDAE

fenestrae; a small hard palate (= bony palate, palatal bridge); and wide internal choanae (= mesopterygoid space). The skull is also notable for having two pairs of upper incisors, the secondary pair being very small and located behind the first principal pair where they have no cutting function; incisor teeth, which grow throughout life and are rooted in the premaxilla bone; no canine teeth; and well-developed diastema (as in rodents and artiodactyls). There are six high-crowned cheekteeth (P1, P2, P3, M1, M2, M3) on each side of the skull and five on each side of the maxilla.The first tooth (P1) of the upper cheekrow is smaller than the four succeeding teeth (P2, P3, M1, M2), and the last tooth (M3) is very small and sometimes missing (Figure 114). Dental formula: I 2/1, C 0/0, P 3/2, M 3/3 = 28. There is no baculum in the penis, and the testes are situated anteriorly to the penis. Hares, rock-hares and rabbits are terrestrial, living in open, rocky, grassland or bushy habitats. They are primarily nocturnal, but may be active close to dawn and dusk on cool cloudy days. During the day, hares rest in ‘forms’ (small open nests in the grass), rock-hares rest in rocky crevices or under boulders, and rabbits hide in complex underground burrows (or ‘warrens’), which they dig themselves. All species are noted for their fast quadripedal running and manoeuvrability, and rock-hares are capable of jumping from rock to rock and running up steep rock faces. All leporids are herbivorous, grazing on short fresh grass and herbs. Digestion of plant tissue is notoriously difficult; in leporids, efficiency of digestion is enhanced by a very large caecum in the hindgut and by coprophagy (a process whereby faecal pellets are eaten and food passes twice through the digestive system). Hares and rock-hares are solitary, and only occasionally seen in groups of up to three or four. In contrast, rabbits (particularly Oryctolagus cuniculus) are gregarious; several individuals share a burrow and may feed in small groups. Most species of hares and rock-hares have small litters, usually 1–3 young/litter; young are precocial at birth, fully furred with the eyes open, and capable of walking and running within a few hours. Rabbits have larger litters, up to 10–12/litter; young are altricial at birth, naked with the eyes closed, and they remain in the nest until 2–3 weeks of age. The terms ‘hare’ and ‘rabbit’ are not clearly defined. However, ‘hares’ are usually larger than ‘rabbits’, have comparatively longer hindlimbs, are solitary, run with a fast loping gait, have small litters

and do not dig burrows. ‘Rabbits’ exhibit a converse set of characters. Some genera do not fit precisely into either ‘hare’ or ‘rabbit’ categories, and show a mixture of characteristics. Here, the term ‘hare’ is used for species that live in non-rocky habitats, do not dig burrows, have small litters and precocial young; ‘rabbit’ for species that live in nonrocky habitats, dig burrows, usually have large litters and altricial young; and ‘rock-hares’ for species that live in rocky habitats, do not dig burrows, have small litters and (probably) altricial young. One genus, Poelagus, shows a mixture of hare and rabbit characteristics. Taxonomic relationships within the family are uncertain, especially for the genus Lepus. Many specific names have been given to the hares of Africa, mainly because some have large geographic ranges and show great variation in colour, size and length of ear in different parts of their ranges. Historically each ‘new’ form was described as a new species. Current taxonomic methods have reduced the number of species of Lepus in Africa to six, some with many synonyms; however, there may be many ‘cryptic species’ within the species currently allocated to Lepus. Molecular analysis confirms that leporids are a monophyletic group (Robinson & Matthee 2005) and speciation into the major clades occurred 3–6 millions years ago. Within Africa, the alternating periods of wet and dry climates and the concomitant expansion and contraction of forests and savannas has been an important factor in leporid evolution. The genera are distinguished by selected body and skull characters, chromosome number and geographic distribution (see Table 48). African hares and rabbits range in HB size from 368–432 mm (small), 433–496 mm (medium-sized) to 497–561 mm (large). Tail length ranges from 56–79 mm (short), 80–103mm (medium-sized) to 104–127 mm (long). Ear length ranges from 65–90 mm (short), 91–115 mm (medium-sized) to 116–140 mm (long). Ear length relative to GLS ranges from 77–100% (relatively short), 101–123% (medium relative length) to 124–146% (relatively long). The ratio of the mean width of the mesopterygoid space to the mean minimum length of the hard palate (abbreviated to MS/HP) ranges from 52– 84% (low), 85–117% (medium) to 118–149% (high). All of the above categories are based on means. D. C. D. Happold

MS/HP (as %)c

Chromosome number

Incisor teeth

Notes

80

n.d.

Deep groove, no cement

Central Africa

90

2n = 44

Deep groove, no cement

North Africa (islands off South Africa)

30–80

2n = 42

Deep groove, no cement

Southern and eastern Africa

150

2n = 44

No cement

South Africa only. Dark stripe from mouth to base of ear

100–150

2n = 48

Deep groove, with cement (except L. starki)

Throughout Africa

c

For MS = width of mesopterygoid space, HP = minimum length of hard palate (see Figure 114e).

695

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Family LEPORIDAE

GENUS Bunolagus Riverine Rabbit Bunolagus Thomas, 1929. Proc. Zool. Soc. Lond. 1929: 109. Type species: Lepus monticularis Thomas, 1903.

A monotypic genus occurring only in South Africa in bushy habitats close to streams and rivers. The genus is characterized by long ears (longer than in Pronolagus), a dark stripe on the lower jaw from near mouth to base of ear, soft silky hairs on soles of feet, uniformly coloured tail, and hard palate shorter than width of mesopterygoid space. Bunolagus exhibits significant difference in karyotype which separates it clearly from Lepus (Robinson & Skinner 1983): there are fewer chromosomes (2n = 44) than in Lepus (2n = 48). Of the species of Lagomorpha examined, this species has the most derived karyotype, differing from the hypothesized ancestor by seven fusions and five fissions (Robinson et al. 2002). Further details are given in the species profile. D. C. D. Happold

Bunolagus monticularis.

Bunolagus monticularis RIVERINE RABBIT Fr. Lapin des Boschimans; Ger. Flusskaninchen (Buschmannhase) Bunolagus monticularis (Thomas, 1903). Ann. Mag. Nat. Hist., ser. 7, 11: 78. Deelfontain, Cape Colony, South Africa.

Taxonomy Originally described in the genus Lepus. Synonyms: none. No subspecies. Chromosome number: 2n = 44 (Robinson & Skinner 1983).

Similar Species Lepus capensis. Dorsal pelage grizzled, greyish, less fluffy; tail black above, white below; no black stripe on lower jaw; groove on each upper incisor filled with cement; grassy habitats.

Description Small dark rabbit with long ears. Pelage soft and fluffy. Dorsal pelage grizzled (agouti) blackish-brown, without rufous patch on rump; hairs grey at base, with white subterminal band and black tip. Flanks similar to dorsal pelage, becoming rufous on lower flanks. Ventral pelage white or pale rufous, usually confined to narrow band on mid-ventral line. Nuchal patch rich rufous. Head similar to dorsal pelage, with conspicuous white or pale buff eye-ring; white or buff colouration may extend anteriorly to nasal region. Thin brown or black stripe along lower jaw to base of ear. Ears comparatively and relatively long, broad, inner margin lined with white hairs, tips rounded and bordered by short black hairs on outer surface. Forelimbs similar colour to flanks; soles of forefeet with thick dense pale rufous hairs. Hindlimbs similar colour to flanks; hindfoot medium brown above, pale rufous-brown below; soles of forefeet with thick dense pale rufous hairs. Tail mediumsized, dark brown; hairs long and fluffy, slightly grizzled, without any white hairs. Skull characteristics include: GLS comparatively short; minimum length of hard palate comparatively short; MS/HP ratio high (ca. 147%); lacks antero-external shoulders on zygoma (cf. other African lagomorph genera); single groove on each principal upper incisor tooth not filled with cement (Figure 115, see also Table 48). Nipples: not known. Geographic Variation None recorded. Figure 115. Skull and mandible of Bunolagus monticularis (BMNH 2.12.1.26).

696

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Bunolagus monticularis

of the Mesembryanthemaceae. Grasses are eaten only when available in the wet season (Duthie 1989, in Duthie & Robinson 1990). Social and Reproductive Behaviour Solitary with a polygamous mating system. Mean home-range for !! is 20.9 ha and for "" is 12.9 ha; home-range of a ! can overlap with that of several "" (Duthie 1989, in Duthie & Robinson 1990).Young born in nest lined with fur and grass in a burrow, rather similar to that of the European Rabbit. Reproduction and Population Structure Reproductive season extends from Aug to May during the warmer seasons of the year. Litter-size: one (occasionally two). Females may have a postpartum oestrus. At birth, young weigh ca. 40 g. Like other species of rabbits, the young are born altricial, blind and helpless at birth, and will only leave the burrow when old enough to look after themselves. Predators, Parasites and Diseases Little information. Predators include African Wild Cats Felis silvestris, domestic dogs and Cape Eagle-owls Bubo capensis. Bunolagus monticularis

Pronolagus spp. Pelage dense and harsh; tail uniformly rufous or rufous-black; ears relatively short or medium-sized; no black stripe on lower jaw; rocky habitats. Distribution Endemic to Africa. South-West Arid (Karoo) BZ. Restricted to a small area in Northern and Western Cape Provinces, South Africa (Districts of Victoria West, Beaufort West Sutherland, Calvinia, Touws River and Frazerburg; additional details in Duthie et al. 1989). Habitat Thick riverine bushland along seasonal rivers, especially where Sasola glabrescens and Lycium spp. predominate. In this habitat, grasses are uncommon and represent only about one-fifth of the cover provided by dicotyledonous plants (Duthie et al. 1989). The habitat is shared with Cape Hares (Lepus capensis). Abundance Uncertain, but rare with a very small geographic range. Two censuses suggested densities of 0.064–0.166 individuals/ha, i.e. about one individual/6–15 ha. Extrapolation of these figures suggests that, in 1989, the remaining suitable habitat could not support more than ca. 1500 individuals (Duthie 1989, in Duthie & Robinson 1990), although the current population size is estimated to be less than 250 mature individuals (IUCN Red List 2004). Adaptations Terrestrial and nocturnal. Constructs burrows with length of 200–300 mm, and entrance of 90–115 mm wide; nest chamber (120–170 mm wide) is formed at end of burrow. Burrow entrance plugged with soil and twigs when not in use. Locomotion rather slow compared with other leporids (Robinson 1981b). Foraging and Food Herbivorous. Forages by browsing on flowers and leaves of dicotyledons, particularly Pteronia erythrocaetha, Kochia pubescens, Salsola glabrescens, Rosenia humilis and several species

Conservation IUCN Category: Critically Endangered. The rarest of all African lagomorphs, and the only African lagomorph placed in this IUCN category. Before 1948 was seen commonly, but in recent years has become increasingly rare (Robinson 1981b, Duthie et al. 1989). Much of the former range (never large) is now used for cultivation (Duthie & Robinson 1990). Rarity is presumed to be due mainly to habitat changes and a reduction in the area of suitable habitat; in addition, reduction in the numbers of jackals has resulted in an increase in the numbers of Wild Cats Felis libyca and Caracals Felis caracal, which prey on Riverine Rabbits (Robinson 1981b). The species is now the focus of various conservation programmes. Measurements Bunolagus monticularis HB: 429 (337–470) mm, n = 14 T: 92 (70–108) mm, n = 13 HF: 104 (90–120) mm, n = 15 E: 116 (107–124) mm, n = 15 WT: n. d. (ca. 1.0–1.5 kg) GLS: 79.7 (78.5–81.5) mm, n = 6 GWS: 36.3 (35.8–37.9) mm, n = 6 P2–M3: 11.62 (11.2–11.9) mm, n = 6 Mesopterygoid space (width): 7.5 (7.2–7.8) mm, n = 6 Hard palate (minimum length): 5.1 (4.4–5.7) mm, n = 6 Upper principal incisor width: 2.0 (1.9–2.0) mm, n = 5 Bulla width: 9.0 (8.3–9.6) mm, n = 5 South Africa Body measurements: Smithers 1983 Skull measurements: TM Key References Duthie & Robinson 1990; Duthie et al. 1989; Robinson & Skinner 1983; Thomas 1903. D. C. D. Happold 697

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Family LEPORIDAE

GENUS Lepus Hares Lepus Linnaeus, 1758. Syst. Nat., 10th edn, 1: 57. Type species: Lepus timidus Linnaeus, 1758.

Lepus saxatilis.

The genus Lepus contains the largest number of species of any genus in the order Lagomorpha: about 32 spp. worldwide (Hoffmann & Smith 2005), six of these occurring in Africa. Members of the genus occur widely in arid, semi-arid and savanna habitats throughout the African continent. Two species (Lepus capensis, L. victoriae) have particularly large geographic ranges. Usually only one or two species occur in a single region (here often separated by habitat considerations), but three species are sympatric or syntopic on the Ethiopian Plateau and the Horn of Africa. Species in the genus are characterized by their long limbs (especially the hindlimbs), their fast movement (the fastest of all

Figure 116. Skull and mandible of Lepus victoriae (RMCA 92-149-M-0010, as Lepus crawshayi).

Lepus capensis.

698

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Lepus capensis

the lagomorphs) and their comparatively and relatively mediumsized to long ears (except comparatively short in L. fageni). Skull characteristics include: mean GLS > 87 mm (longer than other African lagomorphs except Pronolagus crassicaudatus and P. randensis); minimum length of hard palate short or medium-sized; MS/HP ratio high (except in L. fageni) as in Bunolagus; antero-external shoulders present on zygoma; single groove on each principal upper incisor tooth filled with cement (except L. starki, cf. all other African lagomorph genera) (Figure 116, Table 48). Species of Lepus live in open grassland and bushland habitats, and are not associated with rocks. They do not dig burrows. During the day they rest in ‘forms’ in the open, and they remain motionless to avoid detection. Young are precocial at birth, fully furred, the eyes are open, and they are capable of running within a few hours of birth. Hares are solitary, and associate with other hares only for courtship and mating, and when several congregate at highly favoured feeding areas. The taxonomy of the genus is controversial (see also order profile). Species of the genus that have large geographic ranges show great variation in overall size, pelage colour and length of ear. The two particularly widespread species in Africa, L. capensis and L. victoriae, each have many synonyms, testimony to the large variation within each

species. Taxonomic uncertainties are exemplified by (a) some forms (now synonyms) that may be valid species, (b) currently recognized species that may in fact not be valid species, e.g. L. habessinicus may be a subspecies of L. capensis (Azzaroli-Puccetti 1987a, Flux & Angermann 1990), and (c) different viewpoints on what is the correct name for a species, e.g. L. victoriae (see Hoffmann 1993) is considered to be L. microtis by Hoffmann & Smith (2005) – although the name microtis is considered to be nomina dubia by Petter (1972c) because the holotype is a young animal. The genus is in need of revision. It is not possible to identify hares (or indeed lagomorphs in general) using only one or two characters; a combination of many characters, and ratios between selected measurements, are required for precise identification (Azzaroli-Puccetti 1987a, b). Chromosome numbers do not vary across the world range, but DNA analyses seem likely to resolve many taxonomic problems in the future (Alves et al. 2003). The six species in the genus in Africa are distinguished by body size, ratio of width of mesopterygoid space to minimum length of hard palate, amount of black colouration on tip of ear, shape of groove on principal incisor tooth and presence/absence of cement in that groove. D. C. D. Happold

Lepus capensis CAPE HARE Fr. Lièvre du Cap; Ger. Kap-Hase Lepus capensis Linnaeus, 1758. Syst. Nat., 10th edn, 1: 58. ‘ad Cap. b. Spei’ (Cape of Good Hope, South Africa).

Taxonomy Specimens of this species from Kenya are almost identical to those of L. victoriae where the two species are parapatric or sympatric (Flux & Flux 1983, as L. crawshayi). A similar situation occurs in Somalia where some specimens have been difficult to identify and appear to exhibit characters intermediate between the two species (Azzaroli-Puccetti 1987a). Because of its widespread distribution and inter-population variation, many forms of L. capensis have been described; these were originally given species rank but are now considered to be synonyms even though some of them may yet prove to be valid species (Flux & Angermann 1990). Synonyms: 38 African synonyms are listed by Hoffmann & Smith (2005), of which the following are considered by them to be subspecies: aegyptius, aquilo, carpi, granti, hawkeri, isabellinus, sinaiticus.The taxonomic limits of this species, and its relationships with L. victoriae, are uncertain, and require detailed investigation. Subspecies: none recognized here. Chromosome number: probably 2n = 48 (Robinson 1981a). Description Medium-sized. Pelage soft, not as ‘fluffy’ as in Pronolagus spp. and Bunolagus. Dorsal pelage silvery-grey, grizzled (agouti) with black; hairs white at base with wide black subterminal band, whitish terminal band, and black or white tip. Underfur white or greyish-white. Flanks similar to dorsal pelage, becoming very pale buff on lower flanks. Ventral pelage pure white; long. Head similar in colour to dorsal pelage. Lateral profile of head (from forehead to nasal region) distinctly angular (i.e. with obtuse bend downwards above the eye (cf. smoothly convex in L. victoriae) (Flux & Flux 1983). Eye-ring white, often with rufous markings above and below eye-ring. Cheeks greyish-brown. Upper lips pale rufous. Chin and throat white. Gular collar buffy-white or buffy. Ears relatively long (ca. 142% of GLS),

finely covered with buffy hairs; inner margin fringed with long white hairs; outer margin fringed with very short white hairs; tip of ear rounded, fringed with short black hairs especially on outer surface. Nuchal patch brownish-pink; rather inconspicuous. Forelimbs pale rufous above, white below. Hindlimbs pale rufous. Soles of all feet with buffy-brown hairs. Tail comparatively long, fluffy, black above, white laterally and below. MS/HP ratio high (ca. 140%). Each principal incisor tooth with small groove filled with cement. Geographic Variation Pelage colour varies through range (and hence many synonyms). Individuals from arid and semi-arid habitats are paler in colour (dorsal pelage beige, oatmeal, with only a small amount of black speckling) than those from more mesic habitats (see Description above). Length of ear and hindfoot increase with increasing aridity of the habitat (see table below; BMNH): Form (n)

Locality

HF (mm)

E (mm)

Ratio E/HF %

aethiopicus (5) hawkeri (5) sefranus (3)

E Sudan W Sudan Algeria Niger, Algeria

80–102 99–104 99–110

110–118 93–106 100–112

114–132 93–102 91–113

95–114

117–135

114–127

whitakeri (7)

Similar Species L. victoriae. Dorsal pelage brown, grizzled with black; lateral profile of head smoothly convex; nuchal patch orange/brownish-orange; scrub, bush and grassland habitats. 699

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Family LEPORIDAE

into cover as do L. victoriae) (Flux & Flux 1983). Cape Hares may be affected by larger mammals in the same way as L. victoriae, but detailed information is lacking. Cape Hares may assist dispersal of seeds for those species that have seeds with hooks or barbs. Agnew & Flux (1970) list 17 species of plants (mainly grasses) that have been found attached to the pelage of hares. The commonest seed was Tragus berteronianus (76% of seeds), followed by Achyanthes aspera, Pupalia lapacea and Boerhavia repens (all 50% shrub cover (Frame & Wagner 1981). Numbers vary according to season of the year and habitat: in Queen Elizabeth N. P., Uganda, the number of hares seen during night counts ranged from 6.6 to 9.1 hares/km (annual mean 7.8 hares/km) in short grass/sparse thickets, to 0.6–2.3 hares/km (annual mean 1.4 hares/km) in short grass/dense thickets (Ogen-Odoi & Dilworth 1987). Hares are attracted to areas where grasses are sprouting after burning. Most estimates of density and biomass should be taken with caution because hares are difficult to census. The numbers of hares remains relatively constant during each season, probably because they breed throughout the year (see below). In this respect, they differ markedly from some temperate species in the genus that exhibit marked annual and multi-annual fluctuations in numbers. Adaptations Terrestrial and nocturnal. Normally run for cover when disturbed (cf. L. capensis) (Flux & Flux 1983). The number of hares may be partly determined by other species of mammals. Light to moderate grazing of grasslands by Common Hippopotami

Reproduction and Population Structure Reproduction occurs throughout the year in Kenya on the Equator, with all sampled "" being pregnant except in May and Nov when 80% of sampled "" were pregnant (Flux 1981a). Mean litter-size: 1.6. Weight of young at birth: ca. 100 g. Number of litters/year: 6–8. Mean number of young/year: 13.9 (equivalent to 68% of adult female weight – a very high percentage by lagomorph standards). This hare, like L. capensis in Kenya, is a good example of the reproductive strategy in a tropical hare near the Equator (i.e. long reproductive season, many litters/year, small number of young/litter and a high reproductive effort); such a strategy is possible because the environment (primarily rainfall and food resources) enables reproductive activity throughout the year (Flux 1981a). No information available from other localities. Predators, Parasites and Diseases No information. Conservation

IUCN Category: Least Concern (as L. microtis).

Measurements Lepus victoriae (as L. crawshayi) HB: 495 (415–575) mm, n = 13 T: 92 (68–121) mm, n = 13 HF: 113 (103–127) mm, n = 13 E: 102 (93–119) mm, n = 13 WT: 2.31 (1.36–3.17) kg, n = 5 GLS: 89.2 (84.9–93.6) mm, n = 5 GWS: 41.4 (39.4–43.2) mm, n = 5 P2–M3: 13.5 (12.9–14.3) mm, n = 5 Mesopterygoid space (width): 8.3 (7.4–9.1) mm, n = 5 Hard palate (minimum length): 6.7 (6.2–8.0) mm, n = 5 Upper principal incisor width: 3.0 (2.8–3.3) mm, n = 5 Bulla width: 11.0 (9.6–3.3) mm, n = 5 Kenya (BMNH) Key Reference Flux & Angermann 1990. D. C. D. Happold 707

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Family LEPORIDAE

GENUS Oryctolagus European Rabbit Oryctolagus Lilljeborg, 1873. Sverig. Og Norges Ryggradsdjur 1: 417. Type species: Lepus cuniculus Linnaeus, 1758.

Oryctolagus cuniculus.

A monotypic genus, widespread in Europe but confined in Africa to the extreme north-west of the continent and some islands off southern Africa. The genus is distinguished by its comparatively small body size, small hindfeet and short ears. Skull characteristics include: GLS short (shortest of all African lagomorphs); mean minimum length of hard palate short; MS/HP ratio medium (90%); antero-external shoulders present on zygoma; single groove on each principal upper incisor tooth not filled with cement (Figure 117, Table 48). The single species, O. cuniculus, is unique among African lagomorphs because it is a social species, constructs large subterranean burrows and has large litters. Other characteristics of the genus are given in the species profile.

Figure 117. Skull and mandible of Oryctolagus cuniculus (BMNH 19.17.7.2533).

D. C. D. Happold

Oryctolagus cuniculus EUROPEAN RABBIT Fr. Lapin de garenne; Ger. Europäisches Wildkaninchen Oryctolagus cuniculus (Linnaeus, 1758). Syst. Nat., 10th edn, 1: 58. ‘in Europa australis’ (= Germany).

Taxonomy Originally described in the genus Lepus.The European Rabbit has been rarely studied in Africa. Populations in Morocco and Algeria were probably introduced in the eighteenth century (Kowalski & Rzebik-Kowalska 1991). Loche (1867, in Kowalski & Rzebik-Kowalska 1991) referred to the Algerian Rabbit as Cuniculus algirus; more recent authors place algirus as a subspecies of Oryctolagus cuniculus on the basis of its smaller size. Populations on islands off the coast of South Africa (see below) are descended from domesticated strains of O. cuniculus first released in the 1650s (Smithers 1983). Synonyms: nine (all extralimital to Africa; Hoffmann 1993). Subspecies (Africa only): algirus. Chromosome number: 2n = 44 (Schroder & Van der Loo 1979). Description Oryctolagus cuniculus algirus is a small greyish-brown lagomorph, smaller than all other lagomorphs in Africa. Dorsal pelage pale brown, slightly flecked with black and buff; hairs ginger-

brown at base, buff terminally, sometimes with black tip. Underfur grey. Ventral pelage white tinged with pale ginger-buff; hairs white, some with ginger-buff tip. Head similar in colour to dorsal pelage. Chin and throat white. Eye-ring absent. Ears comparatively and relatively short (ca. 98% of GLS), dark brown, darker than dorsal pelage. Nuchal patch pale rufous-brown. Gular patch ginger-buff tinged with orange. Fore- and hindlimbs short (cf. Lepus spp.), pale brown. Hindfeet white above, soles thickly covered with pale brown hair. Tail short, same colour as dorsal pelage above, brown or white laterally, white below. Skull: see family and genus profile. Geographic Variation Extralimitally, pelage colour varies geographically. Specimens from North Africa are paler than those from Spain. Domestic rabbits (and feral descendants) vary in colour from white to brown and black, with or without different colour patches.

708

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Oryctolagus cuniculus

Similar Species Lepus capensis. Much larger in all measurements, ears comparatively and relatively much longer; each principal incisor tooth with groove filled with cement; widespread. Distribution Mediterranean Coastal BZ. Recorded from N Morocco and N Algeria (coastal regions and Tell Atlas), but not further eastwards to Libya; also on Habibas Is (west of Oran, Algeria). Introduced in the 1600s on to several islands near the coast of South Africa and Namibia: currently present on Jutten, Schaapen, Vondeling, Dassen and Robben Islands near Cape Town, Bird I. near Port Elizabeth, and Possession I. near Luderitz (Smithers 1983, Lever 1985, Flux et al. 1990). (Distribution on these islands not shown on map.) Introduced to several other smaller islands near the South African coast, but now locally extinct. Extralimitally widespread throughout continental Europe (natural populations); introduced into Britain (probably in eleventh century), Australia, New Zealand and several South American countries, and many oceanic islands (Flux & Fullager 1983, Flux et al. 1990, Lever 1985). Habitat High mountains, dense bushy regions, and arid habitats in Morocco (Aulagnier & Thévenot 1986). Tends to avoid forested habitats and open areas in Algeria (Kowalski & Rzebik-Kowalska 1991). Sometimes found in cultivated areas. Abundance Common in W Algeria, less common in E Algeria (Kowalski & Rzebik-Kowalska 1991). Populations in Algeria were reduced when the disease myxomatosis was introduced. Adaptations Mainly nocturnal, but active at dawn and dusk when conditions are suitable. Dig extensive underground burrows (‘warrens’) for resting in during the day, and for protection when threatened. Considered a pest in cultivated crops when population numbers are high. As for European Rabbits elsewhere, very adaptable

and prolific. Apart from North Africa, European Rabbits have not colonized Africa (as they have in other continents where they have been released); it seems that competition and predation have prevented individuals (whether introduced purposely, or escaped from captivity) from establishing permanent populations. Foraging and Food Herbivorous. Graze on grasses. No detailed information for African populations. Social and Reproductive Behaviour No information for North Africa, but likely to show similar behaviour to populations elsewhere. Social and territorial. A dominant ! associates with several "" and their young. When population numbers are low, groups are small; when high, social groups may defend territories but may join other groups to feed at night. Reproduction and Population Structure No detailed information for North Africa. Elsewhere very prolific, with an ability to breed opportunistically when conditions are favourable. In general, the reproductive season is shorter at higher latitudes and longer at lower latitudes, and more opportunistic in drier arid habitats than in wetter temperate climates. No data on reproductive season in North Africa, but probably occurs during summer months (Apr–Sep) in Atlas Mts, and in spring and autumn (or even winter) on the semiarid Mediterranean coast and opportunistic reproduction in summer when conditions are favourable. In North Africa, it is probable that reproductive data are similar to elsewhere in southern Europe (as given below). Young born in burrow, in nest of fur made by mother. Gestation: 28–30 days. Litter-size: 3–9, 4–6 or 3–4, depending on season and environmental conditions. Females have postpartum oestrus. At birth, young are altricial, naked, with eyes closed.Weaned ca. Day 20, when first leave nest. Maturity: 3–4 months. Population numbers fluctuate, often greatly. High productivity of young is matched by heavy predation by predators. Population numbers increase during the reproductive season and decline at other times of the year (Gibb 1990). The reproductive strategy of the European Rabbit contrasts greatly with that of hares (genus Lepus). Predators, Parasites and Diseases Occasionally preyed on by owls, and hunted by humans in Algeria (Kowalski & RzebikKowalska 1991). Probably has numerous other predators.The disease myxomatosis (caused by the virus Myxoma) has reduced population numbers in many parts of the world. Conservation IUCN Category: Least Concern (worldwide). Although common and widespread (and regarded as pests at times) in some continents, status in North Africa is not known.

Oryctolagus cuniculus

Measurements Oryctolagus cuniculus HB: 368 (355–380) mm, n = 5 T: 69.0 (65–70) mm, n = 5 HF: 84.4 (80–89) mm, n = 5 E: 73 (70–76) mm, n = 4 WT: n. d. GLS: 74.5 (72.7–77.2) mm, n = 5 GWS 35.9 (34.6–36.7) mm, n = 5 709

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P2–M3: 11.8 (10.0–12.5) mm, n = 5* Mesopterygoid space (width): 5.3 (5.0–5.7) mm, n = 5 Hard palate (minimum length): 5.9 (5.5–6.1) mm, n = 5 Upper principal incisor width: n. d. Bulla width: n. d. Morocco (Petter & Saint-Girons 1972)

*Algeria (Kowalski & Rzebik-Kowalska 1991) Key References (Africa only) Flux et al. 1990; Kowalski & Rzebik-Kowalska 1991; Smithers 1983. D. C. D. Happold

GENUS Poelagus Bunyoro Rabbit Poelagus St Leger, 1932. Proc. Zool. Soc. Lond. 1932 (1): 119. Type species: Lepus marjorita St Leger, 1929.

Poelagus marjorita.

A monotypic genus distributed in savanna habitats in eastern and central Africa to the north of the Rainforest BZ. The genus lacks unique features (Corbet 1983). Like most Lepus, the body is mediumsized, individuals are mostly solitary, do not live in extensive underground burrows, and litters are small. As in Oryctolagus, the skeleton is not built for fast movement, the ears are comparatively and relatively short, and the young are altricial at birth. Skull characteristics include: GLS of medium length; minimum length of hard palate medium; MS/HP ratio low (ca. 84%) as in Pronolagus; antero-external shoulders present on zygoma; single groove on each principal upper incisor tooth not filled with cement (Figure 118, Table 48). The mixture of characteristics suggest that the single species does not fit into any other genus of Lagomorpha, and that a separate genus, Poelagus, is warranted. Other characteristics of the genus are given in the species profile.

Figure 118. Skull and mandible of Poelagus marjorita (RMCA no number).

D. C. D. Happold

Poelagus marjorita BUNYORO RABBIT (UGANDA GRASS HARE) Fr. Lapin sauvage d’Afrique centrale; Ger. Bunyoro-Buschkaninchen Poelagus marjorita (St Leger, 1929). Ann. Mag. Nat. Hist., ser. 10, 4: 292. Near Masindi, Bunyoro, Uganda.

Taxonomy Originally described in the genus Lepus. Synonyms: larkeni, oweni. Subspecies: none. Chromosome number: not known. Description

Medium-sized lagomorph with comparatively

short ears. Dorsal pelage buffy-brown, grizzled (agouti) with black hairs; hairs whitish-grey at base, with wide black subterminal band, pale brown to buff terminal band, and black tip. Underfur greyishwhite. Flanks paler, mainly buffy-brown; most hairs without black

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Poelagus marjorita

tips. Ventral pelage yellowish-buff; hairs whitish-grey on basal half, yellowish-buff on terminal half. White mid-ventral stripe from chest (ca. 20–30 mm wide) to lower abdomen (ca. 40–50 mm wide), extending posteriorly on to inner surface of hindlimbs; hairs pure white. Ventral underfur pure white. Head similar in colour to dorsal pelage; chin and throat white. Ears comparatively and relatively short (ca. 77% of GLS), similar in colour to dorsal pelage, usually without fringe of white hair on ear margins; brown hairs on inner surface; no black on tip. Nuchal patch rufous, not extending on to sides of neck. Fore- and hindlimbs brownish-buff, soles of feet with dense rufous or blackish hairs. Tail short; same colour as dorsal pelage above and on sides, paler (often with some white hairs) below. Both sexes have glandular slits on either side of the genitalia. Juveniles with deep rufous nuchal patch; hairs on soles of feet whitish or grey. Skull: see family and genus profiles. Nipples: not known. Geographic Variation None recorded. Similar Species L. victoriae. Tail longer; ear comparatively and relatively longer (ca. 114% of GLS); each principal incisor tooth with wide deep groove filled with cement; more widespread. L. capensis. Dorsal pelage silvery-grey, grizzled with black; tail and hindfoot longer; ear comparatively and relatively longer (ca. 142% of GLS); each principal incisor tooth with wide deep groove filled with cement; more widespread. Distribution Endemic to Africa. Eastern Rainforest–Savanna Mosaic and Guinea Savanna BZs. Recorded from C and W Uganda, S Sudan, NE DR Congo and NE Central African Republic. There is no evidence for the species in Ruanda, Burundi, Kenya, S Chad, S DR Congo and N Angola (contra Duthie & Robinson 1990 and Kingdon 1997) (see Happold & Wendelen 2006). Habitat Primarily woodland savanna; also stony habitats and hills with short grass (Hatt 1940a). May also occur in forests (e.g. in S Sudan; Setzer 1956). Abundance Uncertain. Reported to be common in open savanna scrub in S Sudan (Setzer 1956), and in Garamba N. P., DR Congo (Verheyen & Verschuren 1966). In Uganda, reported in 1928 as ‘Abundant in certain localities at night grazing on grassy tracks and roads’, and in 1958 as ‘Very common grazing by roads at night’ (labels, BMNH). Adaptations Terrestrial and primarily nocturnal. During the day, rests alone in a form in thick vegetation. Locomotion is more similar to that of a rabbit than of a hare (Kingdon 1974). This is probably because the skeleton is rabbit-like, e.g. scapula is long and narrow (broad and ‘shovel-like’ in hares), ulna is sturdy (reduced in hares), transverse processes of lumbar vertebrae are narrow (expanded in hares) and cervical vertebrae are short (elongated in hares) (St Leger 1932); however, in some specimens, these characters are not so pronounced and are more similar to those of Lepus (Hatt 1940a). Foraging and Food Forages at night on flowers and sprouting grasses. Tends to prefer pastures that have been heavily grazed by

Poelagus marjorita

larger mammals, newly mown fields and burnt areas where the grasses are sprouting (Kingdon 1974). Quantitative data on diet is not available. Social and Reproductive Behaviour Probably solitary when resting in a form; feeds at night in small groups (pairs, or "" with young). May be found on rocky habitats with Rock Hyraxes (Kingdon 1974). Reproduction and Population Structure Newborn young recorded in Jan, Feb, Mar, Jun, Aug and Oct in Garamba N. P., NE DR Congo (Hatt 1940a, Verheyen & Verschuren 1966), and juveniles (WT: 185–200 g) recorded in Jan, Feb, May and Aug (labels, RMCA) suggest that reproduction occurs in most (if not all) months of the year. One large embryo recorded in mid-Aug (Faradje, DR Congo; Hatt 1940a). Gestation thought to be about five weeks (Kingdon 1974). Litter-size: 1–2. Young born in short burrow, the entrance concealed by grass and soil (Kingdon 1974). At birth, young are blind and helpless, with sparse covering of short hair (as in Oryctolagus cuniculus). Predators, Parasites and Diseases No detailed information, but likely to include Servals Felis serval, Genets Genetta spp., hawks and owls (Kingdon 1974). In Uganda, hunted with nets and dogs. Conservation

IUCN Category: Least Concern.

Measurements Poelagus marjorita HB: 451.9 (400–605) mm, n = 17 T: 55.6 (38–70) mm, n = 17 HF: 97.5 (65–108) mm, n = 17 E: 64.7 (61–70) mm, n = 5 WT: 2.68 (2.26–3.17) kg, n = 5* 711

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Poelagus marjorita.

GLS: 83.8 (78.0–89.6) mm, n = 20 GWS: 40.3 (38.8–42.6) mm, n = 20 P2–M3: 13.6 (13.1–14.4) mm, n = 5* Mesopterygoid space (width): 6.7 (6.1–7.6) mm, n = 9* Hard palate (minimum length): 8.0 (7.3–8.7) mm, n = 9* Upper principal incisor width: 3.0 (2.9–3.3) mm, n = 9* Bulla width: 9.3 (8.4–10.2) mm, n = 9*

Uganda (BMNH) and DR Congo (Hatt 1940) *BMNH only Key References Duthie & Robinson 1990; Happold & Wendelen 2006; Kingdon 1974. D. C. D. Happold

GENUS Pronolagus Rock-hares Pronolagus Lyon, 1904. Smithson. Misc. Coll. 45:416. Type species: Lepus crassicaudatus I. Geoffroy, 1832.

that are only slightly longer than forelimbs (cf. Lepus), ears short to medium-sized, uniformly coloured rufous or rufous-black tail, and a considerable amount of reddish or rufous colouration on the limbs and ventral surface. Skull characters include: GLS short to mediumsized; minimum length of hard palate medium to long (depending on species); MS/HP ratio low; antero-external shoulders present on zygoma; and a single groove on each principal upper incisor tooth not filled with cement (Figure 119, Table 48) (cf. most Lepus spp.). Rock-hares are unique in their ability to run and jump over rocks and boulders; they live in small colonies (perhaps because of restrictions in the extent of their habitat) and litter-size is small. Young are altricial at birth (like rabbits) and are born in a nest lined with fur. Traditionally three species have been recognized (P. crassicaudatus, Pronolagus rupestris. P. randensis, P. rupestris) (Hoffmann & Smith 2005). Here, the form saundersiae (listed as a synonym of P. rupestris by Hoffmann & Smith The genus Pronolagus – the Rock-hares – contains three or four 2005) is also considered to be a valid species (Whiteford 1995, species that occur mainly in southern Africa. One species, P. Matthee & Robinson 1996). rupestris, is also represented in eastern Africa, but its taxonomic The species are mainly distinguished by the size of body, hindfoot status is uncertain. Rock-hares (unlike other lagomorphs) are and ear, geographic range and by selected ratios (see profiles). always associated with rocky habitats. Diagnostic characters of the genus are medium to large size (as in most Lepus spp.), hindlimbs D. C. D. Happold 712

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Pronolagus crassicaudatus

Figure 119. Skull and mandible of Pronolagus crassicaudatus (RMCA RG2624).

Pronolagus crassicaudatus NATAL RED ROCK-HARE Fr. Lièvre Roux du Natal; Ger. Natal-Rotkaninchen Pronolagus crassicaudatus (I. Geoffroy, 1832). Mag. Zool. Paris 2: cl. 1, pl. 9. ‘Port Natal’ (= Durban South Africa).

Taxonomy Originally described in the genus Lepus. Formerly treated as a subspecies of P. randensis but now considered to be a valid species. Synonyms: kariegae, lebombo, lebomboensis (lapsus), ruddi. Subspecies: possibly five (see below). Chromosome number: 2n = 42 (as in all Pronolagus spp.). Description Large lagomorph with reddish-coloured limbs. Pelage rather dense and harsh. Dorsal pelage brown, grizzled (agouti) and flecked with black, becoming bright rufous on rump; hairs pale rufous at base, with cream subterminal band, black at tip. Underfur grey. Flanks paler than dorsal pelage, with fewer black-tipped hairs. Ventral pelage pale rufous, with irregular white patches and streaks; hairs mostly with white tips. Head greyish-brown, slightly grizzled. Chin, lower cheeks and throat grey or greyish-white; with greyish-white band extending laterally along edge of jaw to nuchal patch. Nuchal patch brown to grey. Gular patch brownish-rufous (contrasting with colour on throat and chest). Ears relatively short (ca. 81% of GLS) and sparsely furred; whitish-grey on outer surface, grey (similar to cheek) on inner surface. Fore- and hindlimbs dull rufous; soles of all feet rufous-brown. Tail short, bright rufous above and below (but not as dark or as black as in other Pronolagus spp.). MS/HP low (ca. 61%). Each principal incisor tooth with groove (close to inner margin of tooth) not filled with cement.

(80–105 mm); tail on average longer (mean 86 mm); sympatric in part of range. Distribution Endemic to Africa. Coastal Forest Mosaic, Highveld and parts of Afromontane–Afroalpine BZs in SE South Africa. Recorded from eastern South Africa (primarily KwaZulu–Natal and Mpumalanga Provinces), Lesotho, Swaziland and extreme S Mozambique (Meester et al. 1986, Duthie & Robinson 1990). Occurs from sea level to 1550 m. Syntopic in northern and eastern part of range with P. saundersiae. Distribution is patchy (see below). Two records from Mozambique (Smithers & Lobão Tello 1976) are not assignable.

Geographic Variation Petter (1972c), Meester et al. (1986) and Flux & Angermann (1990) list five subspecies, but their status, characteristics and geographic limits are uncertain and their validity is doubtful. Listed here without comment: P. c. crassicaudatus, P. c. ruddi, P. c. karigae, P. c. bowkeri (considered to be a subspecies of P. rupestris by Smithers 1983, Hoffmann & Smith 2005) and P. c. lebombo. Subspecies distinguished partly by colour of nuchal patch (see Taylor 1998). Similar Species P. saundersiae. HB on average shorter (mean 447 mm; ear shorter

Pronolagus crassicaudatus

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Family LEPORIDAE

Habitat Rocky kopjes, rocky hills and ravines where grasses and shrubs grow among the rocks or at base of rocks. Abundance Relatively common and widespread in KwaZulu– Natal (Taylor, 1998). No detailed information from elsewhere. Adaptations Nocturnal. During the day, hides in crevices in rocks and under boulders, or in forms in dense grass. Relies heavily on rocks and boulders for cover, and rarely moves far away from rocks when foraging. The skeleton is ‘rabbit-like’ since the hindlimbs are only slightly longer than the forelimbs. Droppings are deposited in middens, which are often far from resting sites (Duthie 1997). Foraging and Food Herbivorous. No detailed information. Social and Reproductive Behaviour Lives in small colonies consisting of a few individuals (Duthie 1997). Reproduction and Population Structure In KwaZuluNatal, pregnant "" have been recorded in Jun and Aug, and lactating "" in Aug, Oct and Feb (Taylor, 1998). These data suggest that reproduction occurs throughout much of the year. Embryo number: 1 (n = 1) or 2 (n = 3).

Conservation

IUCN Category: Least Concern.

Measurements Pronolagus crassicaudatus HB: 508 (460–560) mm, n = 28 T: 65 (35–110) mm, n = 26 HF: 112 (100–125) mm, n = 12 E: 74 (60–80) mm, n = 17 WT: 2.6 (2.4–3.05) kg, n = 13 GLS: 91.3 (85.3–94.8) mm, n = 27 GWS: 39.6 (36.0–41.8) mm, n = 25 P2–M3: 15.5 (15.2–18.2) mm, n = 28 Mesopterygoid space (width): 5.8 (4.6–6.8) mm, n = 27 Hard palate (minimum length): 9.5 (8.4–11.2) mm, n = 28 Upper principal incisor width: 3.2 (2.9–3.6) mm, n = 24 Bulla width: 6.1 (5.1–7.0) mm, n = 27 South Africa Robinson & Dippenaar 1983a *Smithers 1983 Key References Taylor, 1998.

Duthie & Robinson 1990; Smithers 1993; D. C. D. Happold

Predators, Parasites and Diseases No information.

Pronolagus randensis JAMESON’S RED ROCK-HARE Fr. Lièvre Roux de Jameson; Ger. Jamesons Rotkaninchen (Rand-Wollschwanzhase) Pronolagus randensis Jameson, 1907. Ann. Mag. Nat. Hist., ser. 7, 20: 404. ‘Observatory kopje’, Johannesburg, South Africa. 5900 ft (1798 m).

Taxonomy Formerly included P. crassicaudatus as a subspecies. Synonyms: capricornis, caucinus, ekmani, kaokoensis, kobosensis, makapani, powelli, waterbergensis, whitei. Subspecies: none. Chromosome number: 2n = 42 (as in all Pronolagus spp.). Description Medium-sized; brownish, with rufous limbs and rump. Pelage dense, woolly with silky texture. Dorsal pelage brown, grizzled (agouti), pale rufous on rump and flanks; hairs pale cinnamon at base, with white or cream subterminal band and often black tip. Usually darker on upper back due to many black-tipped hairs. Underfur rufous-brown. Flanks paler. Ventral pelage pale cinnamon, sometimes with white patches; hairs cinnamon with white tip. Head grizzled brownish-grey (without any rufous). Lower cheeks and throat whitish-grey. Ears short (ca. 93% of GLS); brownish-grey with white hairs at tip. Nuchal patch rufous. Gular patch brownish-rufous; hairs with white tips. Fore- and hindlimbs pale rufous, similar to flanks; soles of feet dark brown. Tail medium-sized; blackish-rufous, with black-tipped hairs especially towards tip. MS/HP ratio low (ca. 52%). Each principal incisor tooth with groove (close to inner margin of tooth) not filled with cement. Geographic Variation A preliminary study of mtDNA collected from specimens from six South African localities reveals no geographic variation among populations representing four previously

recognized subspecies (P. r. randensis, P. r. powelli, P. r makapani and P. r. capricorni) (Matthee 1993). Thus there is little geographic variation within the western P. randensis group as suggested by Meester et al. (1986), who recognized only two subspecies: P. r. caucinus (Namibia) and P. r. randensis (South Africa, Zimbabwe and Botswana). Similar Species P. crassicaudatus. Ear shorter (60–80 mm); tail on average shorter (mean 65 mm); eastern South Africa; allopatric. P. rupestris. HB: on average shorter (mean 447 mm), but with ear on average longer (mean 94 mm) and tail on average shorter (mean 86 mm); South Africa, and Malawi northwards to Kenya; allopatric. P. saundersiae. Ear on average longer (mean 94 mm); tail on average shorter (mean 86 mm); marginally sympatric in part of range. Distribution Endemic to Africa. South-West Arid (Namib) and Zambezian Woodland BZs. Two disjunct populations: (1) NE South Africa, E Botswana and Zimbabwe; (2) C and NW Namibia, and perhaps extreme SW Angola. Distribution is patchy because of specialized habitat requirements (see below). Habitat Rocky kopjes, gorges and cliffs, and rocky hills with boulders. Rock crevices and boulders are an essential component of the habitat. In Botswana, occurs on isolated kopjes up to 22 km from

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Pronolagus rupestris

Foraging and Food Herbivorous. Grazes on grasses amongst rocks or at base of rocky hills. Congregates on recently burnt areas to feed on newly sprouting grass. No detailed information on diet. Social and Reproductive Behaviour Generally solitary; sometimes seen in small groups of " and young, or adult " with one or two !!. Several individuals may congregate when grazing (Smithers 1983). In Matopos Hills, Zimbabwe, most nocturnal observations were of single animals, and only 15% were of pairs (Peddie 1975, in Duthie & Robinson 1990). Reproduction and Population Structure Probably breeds throughout the year in Zimbabwe (Peddie 1975, in Duthie & Robinson 1990). Pregnancies recorded in Jan, Jul and Aug; and lactating "" in Jun, Jul and Aug in Zimbabwe (no data for Feb, Mar, Apr, May, Nov, Dec; Smithers & Wilson 1979, as P. crassicaudatus). Litter-size: 1.1 (1–2), n = 8 (Smithers 1983). No information on rate of growth or age at maturation. Predators, Parasites and Diseases No information. Conservation

Pronolagus randensis

nearest kopjes (and from other populations) so, when necessary, individuals have to disperse across intervening non-rocky habitat. Where sympatric with P. saundersiae in hilly mountainous areas, P. randensis tends to be found on the drier low-lying mountain slopes where there are many jumbled boulders and rock crevices, whereas P. saundersiae is found at higher altitudes with fewer boulders and crevices, and higher rainfall. Abundance Very common in the granite hills of Matopos Hills in Zimbabwe and sandstone formations of E Botswana (Smithers 1983). Adaptations Mainly nocturnal but may feed and sunbathe in the late afternoon. During the day, rests in rock crevices, under boulders, or in thick grass close to rocks. If disturbed, stays under cover until the last moment and then disappears behind nearby rocks. Like all rock-hares, can leap from rock to rock, and run up steep rock faces to reach crevices. Characteristic flattened pellet-like droppings are deposited in middens (as in other Pronolagus spp.). Unlike hyraxes, which also live in rocky habitats, Jameson’s Red Rock-hares do not expose themselves on observation boulders (Smithers 1983).

IUCN Category: Least Concern.

Measurements Pronolagus randensis HB: 463 (420–500) mm, n = 12 T: 97 (60–135) mm, n = 13 HF: 100 (87–110), n = 13 E: 84 (80–100) mm, n = 13 WT: 2.3 (1.82–2.95) kg, n = 43* GLS: 90.0 (86.1–92.9) mm, n = 14 GWS: 40.6 (38.6–42.6) mm, n = 14 P2–M3: 15.5 (14.7–16.3) mm, n = 14 Mesopterygoid space (width): 4.8 (3.9–5.3) mm, n = 14 Hard palate (minimum length): 9.3 (8.1–10.1) mm, n = 14 Upper principal incisor width: 3.0 (2.6–3.4) mm, n = 14 Bulla width: 6.3 (5.5–7.0) mm, n = 14 South Africa, Zimbabwe (Robinson & Dippenaar 1983a) *Smithers 1983 Key References

Duthie & Robinson 1990; Smithers 1983. D. C. D. Happold

Pronolagus rupestris SMITH’S RED ROCK-HARE Fr. Lièvre Roux de Smith; Ger. Smiths Rotkaninchen Pronolagus rupestris (A. Smith, 1834). S. Afr. Quart. J. 2: 174. ‘Rocky situations, South Africa’ (probably Rhynsdorp District, South Africa).

Taxonomy Originally described in the genus Lepus. Formerly included in P. crassicaudatus. The taxonomic status of the East African P. rupestris is uncertain and requires investigation, and is here treated as conspecific with the southern African P. rupestris. Synonyms: curryi, fitzsimonsi, melanurus, mülleri, nyikae, vallicola. Subspecies: none. Chromosome number: 2n = 42 (as in all Pronolagus spp.).

Description Medium-sized. Pelage thick and dense, woolly and frequently characterized by a reddish undertone. Dorsal pelage grizzled (agouti) brown anteriorly; hairs pale cinnamon at base, with subterminal white band and black tip; dorsal pelage rufous posteriorly and bright rufous on rump; hairs rufous with white tip. Flanks paler; hairs mostly with white tip and fewer with black tip. Ventral pelage pale rufous to whitish-rufous. Head greyish715

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Family LEPORIDAE

brown. Cheeks greyish-white. Ears ca. 116% of GLS; similar in colour to head with many small off-white or brown hairs. Gular patch brownish. Nuchal patch rufous. Forelimbs bright rufous, contrasting with body colour. Hindlimbs pale rufous (not as bright as forelimbs); hairs whitish-rufous at tip. Soles of all feet very densely covered with dark grey hair. Tail medium-sized, usually black, or black and dark red. MS/HP ratio low (ca. 77%). Each principal incisor tooth with groove (close to inner margin of tooth) not filled with cement. Length of frontal bone almost equal to length of muzzle (cf. P saundersiae). Geographic Variation Pelage colour varies geographically (Smithers 1983). Similar Species P. randensis. Ear on average shorter (mean 84 mm); tail on average longer (mean 97 mm); allopatric. P. saundersiae. Length of frontal bone shorter than length of muzzle; marginally sympatric in part of range. Distribution Endemic to Africa. South-West Arid and Zambezian Woodland BZs; also some areas of Afroalpine–Afromontane BZ. Two disjunct ranges: (1) NW South Africa; (2) SW Kenya, C Tanzania, Malawi and E Zambia. The two ranges are separated by ca. 1200 km. Not recorded from Botswana (Smithers 1971), Zimbabwe (Smithers & Wilson 1979), Swaziland (Monadjem 1998a) and Mozambique (Smithers & Lobão Tello 1976). Habitat Rocky kopjes and rocky hillsides with boulders. Rock crevices and boulders are essential components of the habitat. Habitat very similar to other species of Pronolagus. In South Africa, generally found at lower elevations than P. saundersiae (Matthee & Robinson 1996). Abundance In South Africa, fairly abundant throughout range; expected total population size exceeds 10,000 mature individuals (T. J. Robinson, unpubl.). No information on abundance for East African populations. Adaptations Nocturnal. Emits a wide range of vocalizations: an alarm ‘tu ... tu’ when approached at night, and a grunt when disturbed before sunrise. Young individuals produce a scream when handled, and a ‘churring sound’ when disturbed under a rock (Duthie 1997). Large disc-shaped faecal pellets are deposited in middens (Duthie 1997). Adaptations likely to be similar to those of other rock-hares. Foraging and Food Herbivorous. Forages near rocks, grazing mainly on grasses. In the Ngong Hills, Kenya, plant items in faecal pellets were grass epidermis (mainly Ischaemum afrum 33.5%) and stem fibres (30.6%); small amounts of other grass were present, and a very small amount of dicotyledon epidermis (Stewart 1971b).

Pronolagus rupestris

Reproduction and Population Structure Females give birth to young from spring to summer (Sep–Feb) in South Africa. Gestation: 35–45 days. Litter-size: one or two (Duthie 1997). At birth young are likely to be altricial, with sparse covering of hair and eyes closed (Smithers 1983). Predators, Parasites and Diseases Conservation

IUCN Category: Least Concern.

Measurements Pronolagus rupestris HB: 447 (380–535) mm, n = 15 T: 86 (50–115) mm, n = 15 HF: 92 (85–100) mm, n = 15 E: 94 (80–105) mm, n = 15 WT: 1.62 (1.35–2.05) kg, n = 18 GLS: 80.7 (75.1–85.3) mm, n = 67* GWS: 36.7 (34.5–39.4) mm, n = 67* P2–M3: 14.1 (13.2–15.6) mm, n = 15 Mesopterygoid space (width): 5.3 (4.3–6.8) mm, n = 81* Hard palate (minimum length): 6.9 (5.4–8.5) mm, n = 81* Upper principal incisor width: 2.5 (2.1–3.0) mm, n = 78* Bulla width: 7.2 (6.0–8.5) mm, n = 72* South Africa Body measurements: Robinson & Dippenaar 1983a Weight: Smithers 1983 Skull measurements: TM; *Whiteford 1995 Key References

Social and Reproductive Behaviour Female prepares nest of fur from her body, suggesting that young remain for some time in a nest (Smithers 1983).

No information.

Duthie 1997; Smithers 1983. D. C. D. Happold

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Pronolagus saundersiae

Pronolagus saundersiae HEWITT’S RED ROCK-HARE Fr. Lièvre Roux de Hewitt; Ger. Hewitts Rotkaninchen Pronolagus saundersiae Hewitt, 1927. S. Afr. Quart. J. 2: 174. ‘Rocky situations, South Africa’ (probably Albany district, South Africa).

Taxonomy Originally described as Pronolagus crassicaudatus saundersiae. Referred to as a subspecies of P. rupestris by Meester et al. (1986) and Hoffmann & Smith (2005) but here, following Whiteford (1995) and Matthee & Robinson (1996), it is considered to be a valid species. Synonyms: australis, barretti. Subspecies: none. Chromosome number: 2n = 42 (as in all Pronolagus spp.). Description Medium-sized. Pelage thick and dense, woolly. Dorsal pelage grizzled (agouti) brown anteriorly; hairs pale cinnamon at base, with subterminal white band and black tip; dorsal pelage rufous posteriorly and bright rufous on rump; hairs rufous with white tip. Flanks paler; hairs mostly with white tip and some with black tip. Ventral pelage pale rufous to whitish-rufous. Head greyish-brown. Cheeks greyish-white. Ears ca. 116% of GLS; similar in colour to head with many small off-white or brown hairs. Gular patch brownish. Nuchal patch rufous. Forelimbs bright rufous, contrasting with body colour. Hindlimbs pale rufous (not as bright as forelimbs); hairs whitish-rufous at tip. Soles of all feet very densely covered with dark grey hair. Tail usually red or pale sandy colour. MS/HP ratio low (ca. 77%). Each principal incisor tooth with groove (close to inner margin of tooth) not filled with cement. Length of frontal bone shorter than length of muzzle (cf. P. rupestris). Geographic Variation (Smithers 1983).

Pelage colour varies geographically

Similar Species P. crassicaudatus. HB on average longer; ear shorter (60–80 mm); tail on average shorter (mean 65 mm); sympatric in part of range. P. randensis. Ear on average shorter (mean 84 mm); tail on average longer (mean 97 mm); marginally sympatric in part of range. P. rupestris. Length of frontal bone almost equal to length of muzzle; marginally sympatric in part of range. Distribution Endemic to Africa. South-West Arid (Karoo), South-West Cape and Highveld BZs. Recorded from Western Cape, Eastern Cape, KwaZulu–Natal and Mpumalanga Provinces, South Africa, along the Great Escarpment of South Africa. Habitat Rocky kopjes and rocky hillsides with boulders. Rock crevices and boulders are an essential component of the habitat. Habitat very similar to that of other species of Pronolagus. Generally found at higher elevations than P. rupestris (Matthee & Robinson 1996). Abundance Fairly abundant throughout the range and it is expected that their total population size exceeds 10,000 mature individuals (T. J. Robinson unpubl.). Remarks Since this species was formerly included within P. rupestris, its biology is likely to be similar to that of P. rupestris. Comparative studies are required. No other information available. Conservation

IUCN Category: Least Concern.

Measurements Pronolagus saundersiae HB: 447 (380–535) mm, n = 15 T: 86 (50–115) mm, n = 15 HF: 92 (85–100) mm, n = 15 E: 94 (80–105) mm, n = 15 WT: 1.62 (1.35–2.05) kg, n = 18 GLS: 81.3 (70.6–91.9) mm, n = 109 GWS: 36.72 (32.7–41.4) mm, n = 89 P2–M3: n. d. Mesopterygoid space (width): 5.4 (4.1–8.5) mm, n = 105 Hard palate (minimum length): 6.9 (5.0–8.8) mm, n = 106 Upper principal incisor width: 2.5 (2.0–3.3) mm, n = 99 Bulla width: 6.6 (5.6–7.8) mm, n = 3 South Africa Body measurements: Robinson & Dippenaar 1983a (not differentiated from P. rupestris) Weight: Smithers 1983 Skull measurements: Whiteford 1995 Pronolagus saundersiae

Key References

Duthie 1997; Smithers 1983. D. C. D. Happold 717

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Appendix: New Taxa 2005–2010 Gliridae Graphiurus walterveheyeni Holden and Levine, 2009. Bull. Amer. Mus. Nat. Hist. 331: 341. Distribution: riparian equatorial tropical rainforest, central Congo Basin, Democratic Republic of the Congo. Known altitude 398–431 m.

Muridae Dendromus ruppi Dieterlen, 2009. Bonn. zool. Beitr. 56: 190. Distribution: Gilo, Imatong Mts, East Equatoria, South Sudan; altitude ca. 1800–1900 m. Grammomys brevirostris Kryštufek, 2008. Acta Zool. Lituanica 18: 222. Distribution: Lemesikio, Loliondo, Loita Plains, Kenya (01° 30' S, 35° 09' E). Hylomyscus anselli Carleton and Stanley, 2005. Proc. Biol. Soc. Washington 118: 636. Distribution: highlands in northern Zambia and westernmost Tanzania (Ufipa Plateau); known altitude 1220– 2300 m. Remarks: Described as a subspecies of Praomys denniae; elevated to species by Carleton & Stanley (2005). Hylomyscus arcimontensis Carleton and Stanley, 2005. Proc. Biol. Soc. Washington 118: 629. Distribution: Forested highlands from the Misuku Mts, northern Malawi; to Mt Rungwe and contiguous highlands, south-western Tanzania; eastwards through the Eastern Arc Mountain chain to the South Pare Mts, north-eastern Tanzania; known altitude 900–2410 m. Hylomyscus endorobae (Heller, 1910: 3). Distribution: Highlands of west-central Kenya, including Mt Kenya, the Aberdare Mts and Mau Escarpment; … known altitude 2135–3260 m. Remarks: Long considered a synonym of H. denniae; elevated to species by Carleton et al. 2006. Hylomyscus pamfi Nicolas, Olayemi, Wendelen & Colyn, 2010. Zootaxa 2579: 38. Distribution: type locality Lalama forest, Benin (06° 57' N, 02° 09' E); also recorded from Lougba and Gotcha (Benin), Palimé (Togo), and Asijire and Osogbo (Nigeria). Hylomyscus vulcanorum (Lonnberg and Gyldenstolpe, 1925: 4). Distribution: Mountains bordering the central Albertine Rift Valley, from south-western Uganda and east-central Democratic Republic of Congo, through Rwanda, to southern Burundi; known altitude 1670–3100 m. Remarks: Described as, and conventionally recognized as, a subspecies of H. denniae; elevated to species by Carleton et al. 2006. Hylomyscus walterverheyeni Nicolas, Wendelen, Barriere, Dudu and Colyn, 2008. J. Mammal. 89: 225. Distribution: Doudou Mounts, Ogooue-Maritime Province, SW Gabon (02° 09' S, 10° 30' E); 110 m. Lophuromys chercherensis Lavrenchenko, W. Verheyen, E. Verheyen, Hulselmans and Leirs, 2007. Bull. Inst. R. Sci. Nat. Belg. Biol. 77:

102. Distribution: 22 km north-east Hirna (near road Hirna–Deder), Chercher Mts, eastern Ethiopia (09° 19' N, 41° 15' E, 2700 m). Lophuromys kilonzoi W.Verheyen, Hulselmans, Dierckx, Mulungu, Leirs, Corti and E. Verheyen, 2007. Bull. Inst. R. Sci. Nat. Belg. Biol. 77: 33. Distribution: Magamba (04° 45' S, 38° 17' E; altitude 1550 m) (Tanzania). Lophuromys machangui W. Verheyen, Hulselmans, Dierckx, Mulungu, Leirs, Corti and E. Verheyen, 2007. Bull. Inst. R. Sci. Nat. Belg. Biol. 77: 34. Distribution: Mt Rungwe (09° 10' S, 33° 39' E; altitude 2300 m), forest (Tanzania). Lophuromys makundii W.Verheyen, Hulselmans, Dierckx, Mulungu, Leirs, Corti and E. Verheyen, 2007. Bull. Inst. R. Sci. Nat. Belg. Biol. 77: 38. Distribution: ‘Gerodom’ (foot of Mt Hanang along a brook (04° 28' S, 35° 23' E; altitude ca 2000 m) (Tanzania). Lophuromys menageshae Lavrenchenko, W. Verheyen, E. Verheyen, Hulselmans and Leirs, 2007. Bull. Inst. R. Sci. Nat. Belg. Biol. 77: 99. Distribution: Suba Forest Station, Menagesha Forest, Central Ethiopia (08° 57' N, 38° 33' E, 2600 m). Lophuromys pseudosikapusi Lavrenchenko,W.Verheyen, E.Verheyen, Hulselmans and Leirs, 2007. Bull. Inst. R. Sci. Nat. Belg. Biol. 77: 106. Distribution: Sheko Forest, south-west Ethiopia (07° 04' N, 35° 30' E, 1930 m). Remarks: The exact place of capture was in disturbed humid afromontane forest situated ca. 800 m northwards from the local agricultural office of the Sheko settlement. Lophuromys sabunii W. Verheyen, Hulselmans, Dierckx, Mulungu, Leirs, Corti and E.Verheyen, 2007. Bull. Inst. R. Sci. Nat. Belg. Biol. 77: 36. Distribution: Mbizi on the Ufipa Plateau,Tanzania (07.42° S, 31.40° E; altitude ±1750 m). Remarks: found on forest rim. Lophuromys stanleyi W.Verheyen, Hulselmans, Dierckx, Mulungu, Leirs, Corti and E. Verheyen, 2007. Bull. Inst. R. Sci. Nat. Belg. Biol. 77: 31. Distribution: Mt Rwenzori-Bujuku, Uganda (00.22° N, 29.58° E; altitude 3700 m). Praomys coetzeei Van der Straeten, 2008. Stuttgart. Beitr. Naturk. A, Neu. Ser. 1: 124. Distribution: Duque de Bragança (25 km N – 15 km E), Angola. Otomys dollmani Heller, 1912: 5. Distribution: Known only from Mt Gargues, Mathews Range, central Kenya. Remarks: Long considered a subspecies of O. irroratus or O. tropicalis; specific validity clarified by Carleton & Byrne (2006). Otomys orestes Thomas, 1900: 175. Distribution: Discontinuous in alpine habitats, ca. 2700–4200 m, from western and central Kenya. Remarks:Variously considered as a synonym of O. irroratus, O. tropicalis, or O. typus; specific validity clarified by Carleton & Byrne (2006). Otomys uzungwensis Lawrence and Loveridge, 1953: 61. Distribution: Mountain ranges in west central Tanzania to the Nyika Plateau, northern Malawi; altitude 1800–2750 m. Remarks: Conventionally considered a synonym of O. typus; specific validity clarified by Carleton & Byrne (2006).

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Glossary abbrev. = abbreviation adj. = adjective cf. = confer, compare with; as opposed to Lat. = Latin pl. = plural q.v. = quod vide, ‘which see’ acrocentric: describes a chromosome that has the centromere very near one end and 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: (abbrev.) 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 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: the alternation of pale and dark bands of colour on a hair resulting in the pelage having a grizzled, speckled or ‘pepper and salt’ appearance. Albertine Rift Valley: see Rift Valley (q.v.). alisphenoid: bone in the skull. 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). 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.). 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. precocial. 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 ventrally to the coronoid process (q.v.). anteorbital: in front of the orbit (q.v.). anterior palatal foramen (pl. foramina): foramen (q.v.) in the premaxilla and/or maxilla bone on the ventral surface of the skull

situated in rodents between the incisor teeth and the cheekteeth; foramina always in pairs and elongated in an anterior–posterior direction; sometimes referred to as the anterior incisive foramen. See also posterior palatal foramen. anteroloph: a low transverse enamel ridge that forms part of the anterior cingulum located on the anterior rim of the upper molars of many rodents. anterolophid: a low transverse enamel ridge that forms part of the anterior cingulum located on the anterior rim of the lower molars of many rodents. anthropophilic: living or thriving with humans; inhabiting domiciles in man-made structures and buildings; thriving in habitats substantially modified by humans (e.g. towns, farmlands). cf. lithophilic (q.v.), phytophylic (q.v.). apomorphy (adj. apomorphic): a character state that distinguishes a group of biological organisms from others descended from a common ancestor. cf. plesiomorphy (q.v.). arboreal: living above the ground (in trees and shrubs). cf. scansorial (q.v.); terrestrial (q.v.). auditory bulla (pl. bullae): bony structure encapsulating the middle and inner ear, situated on the ventral surface of the skull. Often greatly inflated in some taxa of arid zone rodents (e.g. Gerbillinae, Dipodidae). Composed of several separate bones, which vary in size and inflation in different genera. Sometimes referred to as tympanic bulla. See also ectotympanic 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. autosomal: pertaining to any chromosome other than the sex chromosomes. baculum (pl. bacula, adj. bacular): the os penis, or penis bone, which supports the penis in some mammals. 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. basicranial axis: a line drawn in the lateral view of the skull indicating the position of the floor of the braincase, in the median line (Harrison & Bates 1991). 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. bicuspid: having two points or cusps (particularly of teeth). bifid: divided by a shallow or deep notch. bipedal: body supported by the two hindlimbs; movement not using the forelimbs. biserial: arranged in pairs (as in the cusps of molar teeth in some mammals, e.g. some rodents). 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. 719

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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.). 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) sometimes added as a suffix to the hindfoot measurement (HF) to emphasize that the measurement includes the claw. cf. s.u. (q.v.). canine: the tooth situated immediately posterior to the incisors or, if incisors are absent, the most anterior tooth. Tall and pointed in most mammals, but absent in rodents and lagomorphs, and in some other orders. By definition, situated on the maxilla bone. See also diastema (q.v.), dental formula (q.v.). cauda epididymides: the ducts of the epididymides at the posterior end of the testes, which 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. CbL: (abbrev.) see condylobasal length. cement: bone-like material that covers part of tooth; material that anchors tooth into its socket. In lagomorphs, may be present in groove on front surface of incisor tooth. 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. cf. (in general usage): compare or compare with. In the context of descriptions, implies a difference or contrast: e.g. Upper incisors do not have a longitudinal groove (cf. Meriones). 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; the chewing surface for rodents and lagomorphs. 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 that 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). cingulum (pl. cingula): ridge around the base of the crown of a tooth. 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. CNL: (abbrev.) 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’. comparatively: used in the context of describing the size of one character compared with the size of the same character in a different species. Sizes described as small, medium or large (if range is divided into three) or very small, small, medium, large, very large (if range is divided into five). cf. relatively (q.v.). competitive exclusion: the principle that two different species cannot indefinitely occupy the same ecological niche. 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 of the skull. condyle: a rounded process on a bone, which articulates with a socket-like concavity in another bone. condylobasal length (CbL): the length of the skull from the most posterior point of one occipital condyle to the most anterior point of the premaxilla on the same side. congeneric: belonging to the same genus. conspecific: belonging the same species. 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 (adj. coprophagous): condition in which an individual reingests its own faeces; any animal that feeds on faeces. 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 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 (that part of the skull that surrounds the brain) when viewed from the side. craniodental: pertaining to the skull and teeth.

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cranium: that part of the skull housing the brain. Also called braincase. crepuscular: pertaining to the twilight, or active during twilight, when light intensity is higher than at night but lower than during the day. cf. diurnal (q.v.); nocturnal (q.v.). 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). cursorial: pertaining to running. cusp (adj. cuspidate): a prominence or sharp point, such as on the occlusal surface of some teeth. See also t. 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. 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 theWest African 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. 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 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 toothrow; superscript numbers indicate upper jaw, subscript numbers indicate lower jaw (mandible), e.g. P4 (upper fourth premolar), M2 (lower second molar). Dental formula, with respect to the presence and number of each of the four types of teeth, varies greatly between orders, families and genera of mammals. See also incisor (q.v.), canine (q.v.), premolar (q.v.) and molar (q.v.).

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. 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.). distichous: arranged in two rows; e.g. long hairs of the tail in some anomalurids and some dormice (as opposed to the hairs being evenly spread all around the tail). diurnal: pertaining to the daytime, or active during daytime, when light intensity is high. cf. crepuscular (q.v.); nocturnal (q.v.). 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. 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. dorsoventral (adj. dorsoventrally): from dorsal to ventral surface; from back to belly of an animal. E: length of external (outer) ear (= pinna), measured from tip of ear to the posterior point of the ear conch. For all mammals, length (and shape) is often affected by preservation. East Africa: Kenya, Uganda, Rwanda, Burundi and Tanzania. eastern Africa: SE Sudan, Ethiopia, Eritrea, Djibouti, Somalia, Kenya, Uganda, Rwanda, Burundi, 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.). ectotympanic bone: sheathing bone, typically hemiglobular in shape, that encloses the middle ear chamber and auditory ossicles in rodents and supports the tympanic membrane. See also auditory bulla. 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.). endemic: restricted to, peculiar to, or prevailing in, a specified country or region. entoconid: principal cusp located at the posterior lingual (inner) side of a lower molar. Eocene: geological epoch (within the Tertiary period), 38–55 mya. epiphysis (pl. epiphyses): any part of a long bone that is formed from a different centre of ossification and which later fuses with the bone to form its terminal part. 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 sufficient water is unavailable. 721

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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 vertebra. extant: living at the present time. cf. extinct. extrabuccal: condition in which incisor teeth extend anteriorly to the lips and are visible externally. 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.). fenestra (pl. fenestrae): opening on a bone, or between two bones, e.g. on maxilla. flank: the side of the body of a mammal. FN: (abbrev.) see fundamental number. 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. forest island: see relict forest. form: in taxonomy, a taxonomic unit (usually named) whose status as either a species or a subspecies is uncertain; one of the varieties found in a polymorphic species. fossorial: adapted for digging; burrowing. cf. subterranean (q.v.). 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. 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; subterranean mammals show many adaptations for life underground, e.g. short

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limbs, thickset shoulder blades and forelimbs, reduced eyes, reduced ability to see, reduction (or absence) of ear pinnae, large extra-buccal incisors, sensory hairs over all the body, feet fringed with hairs, extensive subterranean burrows, etc. (e.g. species of Bathyergidae and Spalacidae). 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. supraorbital: above (dorsal to) the orbit. supraoccipital crest: crest or ridge of bone, oriented 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 process: bony process on outer edge of frontal bone extending outwards above the orbit in lagomorphs. supraorbital ridge: ridge of bone along upper rim of orbit (eyesocket); can be well developed, low or absent. 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.). 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. 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), which is no longer considered to be 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.) (= tubercle) as used to describe and number the cusps on premolar and molar teeth, e.g. t1, t3, t5. The presence/ absence, position and size of longitudinal ridges between cusps are diagnostic for many taxa of rodents. T: (abbrev.) length of tail, measured from the anterior end 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.).

talonid: heel at the posterior end of a lower molar tooth. tapetum lucidum: light-reflecting layer in the retina of the eyes of some vertebrates. 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: 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 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. 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. The boundary of a territory is a line across which the status of the territory holder changes from dominant to subordinate. Tertiary: geological period, 2–65 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. 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. 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 and T. toothrow: the teeth situated posterior to the diastema in rodents and lagomorphs (and some other orders of herbivorous mammals). The upper and lower toothrows are comprised of premolar and molar teeth, or only molar teeth. Although small cusps are present on the surface of each tooth in a toothrow in young animals, these cusps wear with age to form a smooth grinding surface; the pattern of enamel and dentine of the grinding surface may be used to assess the age of the individual. A toothrow may contain three teeth (all molars, e.g. most murid rodents), four teeth (one premolar, three molars, e.g. some squirrels, porcupines, springhares), five teeth (two premolar, three molars, e.g. some squirrels) or six teeth (three premolars, three molars, e.g. lagomorphs). 729

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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 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). transverse: in a direction across the body from side to side. cf. longitudinal (q.v.). tricuspid: having three points or cusps (particularly of teeth). tubercle: a small rounded protuberance, e.g. a cusp of a tooth. 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 or neotype was collected. Also called topotypical locality. 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). 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. 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). Wirch: Ethiopian word for the alpine agricultural/economic altitudinal zone, above about 3000 m, too cold for most agriculture. Woina Dega: Ethiopian word for the warm-temperate agricultural/ economic altitudinal zone, about 1500–2300 m, warm enough for most agriculture. WT: (abbrev.) weight of an individual, usually expressed in grams (g) or kilograms (kg). 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 plate: expanded and flattened lower part of the maxillary process on the outer side of the infraorbital foramen (q.v.); variations in size and shape are useful for identification of some rodents. 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 III Stéphane Aulagnier INRA- Comportement et Ecologie de la Faune Sauvage BP 52627 F-31326 Castanet-Tolosan cedex France email: [email protected]

Hynek Burda Abt. Allgemeine Zoologie FB 9- Biologie Universitaet Essen D-45117 Essen Germany email: [email protected]

Khalilou Bâ Institut de Recherche pour le Développement (UMR 022 CBGP) Bel-Air, BP 1386 Dakar CP 18524 Senegal

Thomas M. 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 email: [email protected]

Afework Bekele Department of Biology Addis Ababa University PO Box 1176 Addis Ababa Ethiopia email: [email protected] Nigel C. Bennett Department of Zoology and Entomology University of Pretoria Pretoria 0002 South Africa email: [email protected] Sean O. Bober Division of Mammals Field Museum of Natural History 1400 South Lake Shore Drive Chicago Illinois 60605 USA email: [email protected] David Brugière University of Rennes Station Biologique 35 380 Paimpont France email: [email protected]

Michael D. Carleton Division of Mammals National Museum of Natural History Smithsonian Institution Washington DC 20560 USA email: [email protected] Christian T. Chimimba Mammal Research Institute Department of Zoology and Entomology University of Pretoria Pretoria 0002 South Africa email: [email protected] Viola Clausnitzer Heinzelstr. 3 02826 Görlitz Germany email: [email protected]

Edith R. Dempster School of Education and Development University of KwaZulu-Natal Scottsville 3209 South Africa email: [email protected] Christiane Denys Laboratoire Mammifères et Oiseaux Museum National d’Histoire Naturelle 55 rue de Buffon F-75005 Paris France email: [email protected] Fritz Dieterlen Staatliches Mueum für Naturkunde Stuttgart Rosenstein 1 D70191 Stuttgart Germany email: [email protected] Gauthier Dobigny Institut de Recherche pour le Développement Campus international de Baillarguet CS 30016 F-34988 Montferrier-sur-Lez cedex France email: [email protected] Jean-François Ducroz Mansfelder Strasse 48a 06108 Halle Germany Jean-Marc Duplantier Institut de Recherche pour le Développement Campus international de Baillarguet CS 30016 F-34988 Montferrier-sur-Lez cedex France email: [email protected]

C. G. Coetzee PO Box 3957 Vineta Swakopmund Namibia email: [email protected] 774

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Authors of Volume III

Louise H. Emmons Division of Mammals National Museum of Natural History Smithsonian Institution Washington DC 20560 USA email: [email protected]

Rainer Hutterer Zoologisches Forschungsmuseum und Museum Alexander Koenig Adenauerallee 160 D-53113 Bonn Germany email: [email protected]

Jonathan Kingdon Department of Zoology University of Oxford WildCRU, Tubney House Abingdon Road Tubney OX13 5QL UK

Elisabeth Fichet-Calvet Bernhard-Nocht Institute of Tropical Medicine Bernhard-Nocht Strasse 74 20359 Hamburg Germany email: [email protected]

Tim P. Jackson Department of Zoology and Entomology University of Pretoria Pretoria 0002 South Africa email: [email protected]

Leonid A. Lavrenchenko A. N. Severtsov Institute of Ecology and Evolution Russian Academy of Sciences Leninsky Pr. 33 117071 Moscow Russia email: [email protected], [email protected]

Patrick Gouat Laboratoire d’Ethologie Experimentale et Comparee UPRES-A 7025–Université Paris Nord F-93430 Villetaneuse France email: [email protected] Laurent Granjon Institut de Recherche pour le Développement Campus international de Baillarguet CS 30016 F-34988 Montferrier-sur-Lez cedex France email: [email protected], [email protected] Peter Grubb (deceased) 82 Drewstead Road London SW16 1AG UK email: [email protected] D. C. D. Happold Research School of Biology Australian National University Canberra ACT 0200 Australia email: [email protected] Mary Ellen Holden 494 Wallace Drive Charleston South Carolina 29412 USA email: [email protected]

J. U. M. Jarvis Department of Zoology University of Cape Town Rondebosch 7701 South Africa email: [email protected] Jan Kalina Soita Nyiro Conservancy PO Box 149 Nanyuki 10400 Kenya email: [email protected] F. Keesing Department of Biology Bard College PO Box 1266 Annandale-on-Hudson NY 12504 USA email: [email protected] Julian C. Kerbis Peterhans Division of Mammals Field Museum of Natural History 1400 South Lake Shore Drive Chicago Illinois 60605 USA email: [email protected] Michael H. Kesner Department of Biology Indiana University of Pennsylvania Indiana PA 15705 USA email: [email protected]

Herwig Leirs Evolutionary Biology Group University of Antwerp (RUCA) Groenenborgerlaan 171 B-2020 Antwerpen Belgium email: [email protected] Alicia V. Linzey Department of Biology Indiana University of Pennsylvania Indiana, PA 15705 USA email: [email protected] Jay R. Malcolm Faculty of Forestry University of Toronto Toronto Ontario M5S 3B3 Canada A. Monadjem Department of Biological Sciences University of Swaziland Kwaluseni Swaziland email: [email protected] Guy G. Musser 494 Wallace Drive Charleston South Carolina 29412 USA email: [email protected]

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Authors of Volume III

J. A. J. Nel Department of Zoology University of Stellenbosch Matieland 7602 South Africa email: [email protected]

Chad E. Schennum Division of Mammals National Museum of Natural History Smithsonian Institution Washington DC 20560 USA

Lindsay A. Pappas Division of Mammals Smithsonian Institution Washington DC 20560 USA

Bruno Sicard Institut de Recherche pour le Développement BP 2528 Bamako Mali email: [email protected]

Mike Perrin School of Botany and Zoology University of KwaZulu-Natal Scottsville 3209 Pietermaritzburg South Africa email: [email protected] Francis Petter (deceased) Laboratoire Mammifères et Oiseaux Museum National d’Histoire Naturelle 55 rue de Buffon 75005 Paris France Neville Pillay Dept of Animal, Plant and Environmental Sciences University of the Witwatersrand Witwatersrand 2050 South Africa email: [email protected] Justina C. Ray Wildlife Conservation Society Canada 720 Spadina Avenue Toronto Ontario M5S 2T9 Canada email: [email protected], [email protected]

Brian J. Stafford Division of Mammals National Museum of Natural History Smithsonian Institution Washington DC 20560 USA S. Takata Museum of Vertebrate Zoology University of California Berkeley CA 94720 USA P. J. Taylor Department of Resource Management University of Venda Thohoyandou 0950 South Africa email: [email protected] Michel Thévenot 353 chemin Mendrous 34170 Castelnau le Lez France email: [email protected]

Richard W. Thorington, Jr Division of Mammals National Museum of Natural History Smithsonian Institution Washington DC 20560 USA email: [email protected] Erik Van der Straeten Departement Biologie Evolutie et Biologie Universiteit Antwerpen Groenenborgerlaan 171 B-2020 Antwerpen Belgium email: [email protected] F. Veyrunes Institut des Sciences de l’Evolution Université Montpellier II Place E. Bataillon F-34095 Montpellier cedex 5 France email: [email protected] Jane M. Waterman Department of Biological Sciences University of Manitoba Winnipeg MB R3T 2N2 Canada email: [email protected] D. W. Yalden Faculty of Life Sciences University of Manchester Manchester M13 9PT UK email: [email protected]

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Indexes French names Ammodile 263 Anomalure de Beecroft 604 de Derby 606 de Pel 608 nain à grandes oreilles 612 nain de Zenker 614 pygmée 610 Athérure d’Afrique 672 Campagnol de Musters 211 Écureuil d’Aubinn 41 de brousse à ventre roux 84 de brousse d’Alexander 74 de brousse de Boehm 75 de brousse de Cooper 79 de brousse de Kershaw 87 de brousse de Smith 77 de brousse de Vincent 88 de brousse ocre 82 de brousse rayé 80 de brousse rouge et noir 81 des palmiers 45 pygmée 71 Écureuil foisseur de Barbarie 43 du Damara 99 du Cap 96 du Sahel 94 unicolore 100 Écureuil géant de Stanger 90 Écureuil gris 92 Funisciure à oreilles noires 47 à pattes rousses 58 d’Isabella 55 de Bocage 50 de Duchaillu 54 de Kintampo 60 de montagne 51 de Thomas 49 du Congo 52 rayé 56 Gerbille à queue courte 267 à queue grosse 341 à queue noire 281 à queue touffue 347 à ventre blanc 279 charmant 298 d’Anderson 299 d’Éthiopie 324 de Berber 297 de Boehm 272 de Botta 300 de Brockman 301

de Cosens 304 de Dunn 306 de Flower 307 de Gambie 274 de Gorongoza 277 de Guinée 276 de Harwood 309 de Henley 310 de Hoogstraal 312 de Jebel Marra 315 de Julian 313 de Kemp 278 de Khartoum 330 de Lataste 314 de Maroc 311 de Nubie 316 de Percival 322 de Phillips 283 de Rosalinde 327 de savanne 285 de Simon 328 de Somalie 330 de Tarabul 331 de Wagner 304 de Waters 332 des champs 302 des rochers 328 du Cap 270 du Maghreb 317 du Nigeriae 320 du Soudan 318 du Veld 273 naine 318 naine de Brauer 264 occidentale 321 pâle 323 pygmée à pieds velus 288 pygmée à queue touffue 293 pygmée de Setzer 290 pygmée de Somalie 340 pygmée des dunes 291 robuste 284 Gerboise Orientale 141 tétradactyle 136 Goundi de Thomas 632 de l’Atlas 630 du Félou 635 du Mzab 636 Grand Aulacode 688 Grande Gerbille d’Egypte 325 Graphiure à grosse queue 113 à tête plate 130 d’Angola 110 de Christy 112 de Johnston 114 de Lorrain 118 de Monard 123

de Nagtglas 126 de Noack 120 des rochers 131 du Cap 128 murin 124 nain 116 sourd 133 Héliosciure à pieds roux 66 de forêt 65 de Gambie 62 de Zanj 69 du Mont Rwenzori 68 variable 64 Hétérocéphale glabre 668 Lapin de garenne 708 des Boschimans 696 sauvage d’Afrique centrale 710 Lérot du Sud-Est Asiatique 105 Nord-africain 107 Lièvre d’Abyssinie 702 des buissons 703 des haut plateaux d’Ethiopie 705 des savanes 706 du Cap 699 éthiopien 701 Lièvre Roux de Hewitt 717 de Jameson 714 de Smith 715 du Natal 713 Lièvre Sauteur d’Afrique de l’Est 624 d’Afrique du Sud 619 Mérione à queue rougeâtre 336 de Shaw 338 de Sundevall 335 Mulot sylvestre 378 Pectinator de Speke 638 Petit Aulacode 687 Petit Écureuil de brousse 85 Petit Rat à abajoues 155 Petit souris adipeux 198 Petite Gerbille de Peters 325 Petite Gerbille des sables 308 Petite Gerboise d’Egypte 138 Porc-épic de l’Afrique du Nord 678 de l’Afrique du Sud 676 Ragondin 691 Rat à abajoues

à longue queue de Mearns 164 du Cap 162 petit Rat à collier 554 Rat à crinière 214 Rat à fourrure de Hopkins 517 des Iles Ssese 518 des marais 514 des ruisseaux 515 plus petite 519 Rat à manteau roux 237 Rat à museau roux commun 510 d’Afrique de l’Ouest 512 Rat à poil doux africain 437 d’Allen 431 d’Angola 433 de Baer 433 de montagne 434 de Stella 438 de Thomas 430 grand 436 Rat à queue courte 506 Rat aquatique d’Afrique 390 d’Ethiopie 508 Rat arboricole à queue noire 559 de Loring 558 de Shortridge 562 des Acacias 561 Rat cible de Defua 400 Rat d’Ethiopie à pieds blancs 549 à queue blanche 550 à queue grise 552 de Rupp 553 Rat de Brant 597 Rat de Harrington 402 Rat de Littledale 600 Rat de Yalden 403 Rat des fourrés charmant 566 de Kemp 564 de Schouteden 565 Rat des rochers de Bocage 364 de Grant 367 de Hinde 368 de Kaiser 370 de Namibie 681 de Nyika 373 de Tete 369 de Thomas 376 du Mont Selinda 374 du Namaqua 371 occidental 375

777

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Indexes

roux 365 Rat des sables obese 344 pâle 346 Rat du Mont Oku 440 Rat du Vlei d’Afrique du Sud 583 d’Anchiete 576 d’Angoni 577 d’Ethiopie 592 de Barbour 579 de Burton 580 de Cuanza 581 de Dent 582 de Karoo 594 de Saunders 588 de Sloggett 589 du lac 585 du tropique 591 laminé 586 occidental 587 Rat forestier à front plat 424 à trois bandes dorsales 425 bai 427 de Basilio 423 du Cameroun 422 du Rwenzori Rat fouisseur gris-argent 665 Rat Géant d’Emin 158 de Gambie 159 Rat hirsute d’Angola 397 Africain 395 de Fox 394 de montagne 396 roux 399 Rat noir 541 Rat palustre commun 458 de Cansdale 456 de Edwards 457 Rat rayé champêtre 545 Rat roussard d’Abyssinie 381 d’Ethiopie 383 de Neumann 386 du Kenya 384 du Nil 387 guinéen 388 soudanien 382 Rat-taupe d’Ansell 649 du Damara 651 de Bocage 650 de Darling 653

de Fox 654 de l’Est Africain 151 des dunes de Namaqua 644 des dunes du Cap 646 du Cap 663 du Kafue 658 du Togo 661 géant 659 géant d’Ethiopie 149 hottentot 655 ocre 660 Souris à crinière 472 Souris à grandes oreilles 186 Souris à mammelles multiples d’Awash 462 de Guinée 464 de Hubert 465 de Kollmannsperger 467 de Shortridge 471 du Natal 468 du sud 463 naine 470 Souris à queue blanche d’Afrique 201 Souris adipeuse commune 199 de Bocage 192 de Jackson 195 de Krebs 196 de Pousargues 197 du Nord-Ouest 193 gracile 194 Souris arboricole de bananier 177 de Bates 190 de Brazzaville 185 de Heuglin 178 de Lovat 173 de Nyika 181 de Vernay 183 des montagnes 171 du Cameroun 182 du Kivu 179 du Mont Kahuzi 172 grise 174 noisette 176 Souris d’Angola 503 Souris de Dybowski 500 Souris de Hildegarde 568 Souris de Rudd 259 Souris de Verreaux 505 Souris de Woosnam 569 Souris des montagnes Bale 188 Souris des rochers à queue courte 204F de Shortridge 208 des Monts Brukkaros 207

pygmée 205 Souris d’Ethiopie 501 Souris domestique 487 Souris épineuse de Heuglin 222 de Johan 224 de Kemp 225 de l’Aïr 219 de Louise 226 de Mullah 227 de Percival 228 de Wilson 234 dorée 230 du Caire 220 du Cap 233 petite 231 rougeâtre 223 Souris fuligineuse 504 Souris fumeuse 418 Souris hérissée à pieds d’or 242 à queue court 240 à queue moyenne 250 à tâche jaune 246 à ventre fauve 249 à ventre feu 252 de Dieterlen 244 de Eisentraut 245 de Hutterer 248 de l’Ouest 255 de Rahm 253 de Rosevear 254 de Woosnam 257 d’Ethiopie 251 gris 243 Souris naine à ventre gris 497 crapaud 477 d’Afrique australe 484 d’Afrique de l’Ouest 486 d’Orange 490 de Baoulé 476 de Callewaert 478 de l’Oubangui 491 de la Gounda 479 de Mahomet 482 de Matthey 483 de Neave 489 de Peters 492 de Setzer 493 de Thomas 494 delicate 496 du desert 481 Haussa 480 Souris palustre de Delany 166 Souris rayée d’Afrique 452

de Barbarie 443 de Bellier 444 de Griselda 445 de Hoogstraal 446 de Mittendorf 449 de Rosalie 449 de Rosevear 451 de Senegal 447 tachetée 448 zébrée 454 Souris sauvage 495 Souris sylvestre commune 410 d’Afrique de l’Ouest 536 d’Ethiopie 417 d’Afrique Est 413 de Bunting 407 de Büttner 359 de Dalton 522 de De Graaff 524 de De Roo 526 de fôret tropicale 414 de Hartwig 527 de Jackson 528 de Jebel Marra 406 de la rivière 533 de Lukolela 529 de MacMillan 416 de Misonne 531 de Mozambique 409 de Petter 535 de Tullberg 537 de Verschuren 538 delicate 525 du Kenya 408 du Mont Cameroun 532 du Mont Kenya 412 du Rwenzori 411 obscure 534 plus petite 530 Spalax d’Ehrenberg 145 Surmulot 540 Tatérille d’Emin 352 de Cuvier 356 de Petter 355 de Tranier 357 des sables 350 du Congo 351 du Lac Chad 354 gracile 353 Zenkerelle 616

German names Äthiopien-Maus Grauschwänzige 552 Rupps 553 Weißchwänzige 550 Weißfüßige 549 Bachratte

Angolanische 514 Hopkins 517 Kleine 519 Rillenzahn- 515 Ssese Inseln 518 Bandicootratte Kurzschwanz- 506

Baumhörnchen (see also Buschhörnchen, Dünnschwanzhörnchen, Palmenhörnchen, Sonnenhörnchen, Riesenhörnschen, Zwerghörnchen) Bocages 50 Carruthers 51 Duchaillu 54

Kintampo- 60 Kongo 52 Lady Burtons 55 Orangenköpf 57 Rotfüßiges 58 Rotloses 49 Streifiges 56

778

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German names

Baumratte Lorings 558 Schwarzschwänzige 559 Shortridges 562 Sundevalls 561 Bilch (see also Löffelbilch) Angolischer 110 Brillen- 128 Busch- 124 Christys 112 Dickschwanz- 113 Felsen- 131 Flachkopf- 130 Johnstons 114 Kellens 116 Kurzohr- 133 Lorraines 118 Monards 123 Nagtglas 126 Noacks 120 Bilchrennmaus 347 Blessmulle (see also Erdbohrer, Graumulle, Nacktmulle, Strandgräber) Kap 663 Blindmäuse Ehrenberg- 145 Breitkopfmaus Hildegardes 568 Woosnams 569 Bürstenfellmaus Rudds 259 Bürstenfellratte Braunbauchige 255 Dieterlens 244 Eisentrauts 245 Gelbbauch 249 Gelbgefleckte 246 Goldfüßige 242 Graue 243 Hutterers 248 Kurzschwanz- 240 Mittelschwänzige 250 Rahms 253 Rosevears 254 Rotbauchige 252 Schwarzkrallen- 251 Woosnams 257 Buschhörnchen (see also Baumhörnchen, Dunnschwanzhörnchen, Palmenhörnchen, Sonnenhörnchen, Riesenhörnschen, Zwerghörnchen) Alexanders 74 Boehms 75 Coopers 79 Gestreiftes 80 Grünes 85 Lushoto- 87 Ockerfarbiges 82 Rotbaüchiges 84 Smiths 77 Tanganjika- 81 Vincents 88 Buschkaninchen (see also Flusskaninchen, Hase, Rotkaninchen,Wildkaninchen) Bunyoro- 710 Buschmaus Buntings 407 Dryas- 411 Glanz- 414

Graukopf- 408 Große 412 Jebel Marra 406 Macmillans 416 Minna- 417 Mosambik- 409 Ostafrikanische 413 Wald- 410 Buschratte Harrington 402 Yaldens 403 Defua-Ratte 400 Deomys-Waldmaus 237 Dickichtratte Anmutige 566 Kemps 564 Schoutedens 565 Dornschwanzhörnchen Beecrofts 604 Derbys 606 Pels 608 Zwerg- 610 Dornschwanzbilche Kameruner 616 Dünnschwanzhörnchen 41 (see also Baumhörnchen, Buschhörnchen, Palmenhörnchen, Sonnenhörnchen, Riesenhörnschen, Zwerghörnchen) Erdbohrer (see also Blessmulle, Graumulle, Nacktmulle, Strandgräber) Silbergrauer 665 Erdhörnchen Damara- 99 Kap- 96 Gestreiftes 94 Nordafrikanisches 43 Streifenloses 100 Felsenmäuse Barbours 204 Brukkaros- 207 Shortridges 208 Zwerg 205 Felsenratte 681 Fettmäuse Bocages 192 Gemeine 199 Jacksons 195 Kleine 198 Krebs 196 Nordwestliche 193 Pousargues 197 Zierliche 194 Fettschwanzmaus 341 Flusskaninchen 696 (see also Buschkaninchen, Hase, Rotkaninchen,Wildkaninchen) Gleitbilche Großohr- 612 Zenkers 614 Grasmaus Belliers 444 Einstreifen- 449 Griselda- 445 Hoogstraals 446 Mehrstreifen- 443

Mittendorfs 449 Rosevears 451 Senegal- 447 Streifen- 452 Tüpfel- 448 Vierstreifen- 545 Zebra- 454 Grasratte Ansorges 382 Äthiopische 381 Blicks 383 Nairobi- 384 Neumanns- 386 Nil- 387 Rote 388 Grauhörnchen 92 Graumulle (see also Blessmulle, Erdbohrer, Nacktmulle, Strandgräber) Ansells 649 Bocages 650 Damaraland 651 Darlings 653 Foxs 654 Hottentotten- 655 Kafue 658 Ockerfarbige 660 Riesige 659 Togo- 661 Großohrmaus 186 Gundi Atlas- 630 Buschschwanz- 638 Felou- 635 Sahara- 636 Thomas 632 Hamsterratten Kap- 162 Langschwanz- 155 Mearns 164 Hase (see also Buschkaninchen, Flusskaninchen, Rotkaninchen, Wildkaninchen) Abessinischer 702 Afrikanischer Savannen 706 Äthiopischer 701 Äthiopischer Hochland 705 Busch- 703 Kap- 699 Hausratte 541 Klettermaus (see also Riesenklettermaus, Samt-Klettermaus) Bananen- 177 Bates 190 Brants 176 Gebirgs- 171 Graue 174 Kahusi- 172 Kamerun- 182 Kastanienbraune 178 Kivu- 179 Lovats 173 Nyika- 181 Vernays 183 Koboldrennmaus Peels 340 Kurzschwanz-Rennmäuse 267 Lamellenzahnratte

Äthiopische 592 Anchietas 576 Angonis- 577 Barbours 579 Burtons 580 Cuanza 581 Dents 582 Gemeine 583 Kap- 586 Karoo- 594 Saunders 588 Sloggetts 589 Tropische 591 Wasser- 585 Westliche 587 Lamotte-Maus Oku 440 Löffelbilch (see also Bilch) Großohr 105 Nordafrikanischer 107 Mähnenratte 214 Maulwurfsratte Afrikanische 151 Riesen- 149 Maus (see also Zwergmaus) Algerische 495 Haus- 487 Mühlenratte Dybowskis 500 Königs- 501 Nacktmulle 668 (see also Blessmulle, Erdbohrer, Graumulle, Strandgräber) Nacktsohlen-Rennmäuse Boehms 272 Brants 273 Fransenschwanz- 284 Gambische 274 Gorongoza- 277 Guinea- 276 Kap- 270 Kemps 278 Phillips 283 Savannen- 285 Schwarzschwanz- 281 Weißbauch- 279 Nutria 691 Palmenhörnchen (see also Baumhörnchen, Buschhörnchen, Dünnschwanzhörnchen, Sonnenhörnchen, Zwerghörnchen) Westliches 45 Pfeifratte Brants 597 Littledales 600 Quastenstachler (see also Stachelschweinartige) Afrikanischer 672 Rauchmaus Afrikanische 418 Rennmaus (see also NacktsohlenRennmaus, Zwergrennmaus) Andersons 299 Anmutige 298 Berbera- 297

779

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Indexes

Bottas 300 Brockmans 301 Cosens 304 Dunns 306 Felsen- 328 Flowers 307 Große Ägyptische 325 Harwoods 309 Helle 323 Henleys 310 Hoogstraals 312 Julians 313 Khartum- 330 Kleine 308 Kleine Kurzschwanz- 328 Kleines SandLatastes 314 Lowes 315 Mackillingins 316 Mahgreb- 317 Marokkanische 311 Nigerianische 320 Nordafrikanische 302 Percivals 322 Polster- 324 Rosalindes 327 Somalische 330 Sudan Tarabuls 331 Wagners 304 Waters 332 Westliche 321 Winzige 325 Zwerg- 318 Rennmäuschen Emins 352 Kongo- 351 Petters 355 Sahel- 350 Schlankes 353 Senegal- 356 Traniers 357 Tschadsee- 354 Rennratte Libysche 336 Shaws 338 Sundevalls 335 Riesenhamsterratte Emins 158 Gambia- 159 Riesenhörnschen (see also Baumhörnchen, Buschhörnchen, Dünnschwanzhörnchen, Palmenhörnchen, Sonnenhörnchen, Zwerghörnchen) Africanische 90 Riesenklettermaus 188 Rohrratte Große 688 Kleine 687

Rotkaninchen (see also Buschkaninchen, Flusskaninchen, Hase, Wildkaninchen) Hewitts 717 Jamesons 714 Natal- 713 Smiths 715 Rotnasenratte Gemeine 510 Westafrikanische- 512

Ruwenzori- 423 Sumpfklettermaus Delanys 166 Sumpfratte Cansdales 456 Edwards 457 Langfüßige 458

Samt-Klettermaus 185 Sandratte Dünne 346 Fette 344 Simien-Maus 472 Springhase Ostafrikanischer 624 Südafrikanischer 619 Springmäuse (see also Vierzehen-Jerboa) Kleine Ägyptische 138 Orientalische 141 Sonnenhörnchen (see also Baumhörnchen, Buschhörnchen, Dünnschwanzhörnchen, Palmenhörnchen, Riesenhörnschen, Zwerghörnchen) Gambisches 62 Geflecktes 65 Mutables 64 Rotbeiniges 66 Ruwenzori 68 Zanj 69 Stachelmaus Aïr- 219 Gold- 230 Heuglins 222 Johans 224 Kairo 220 Kap- 233 Kemps 225 Kleine 231 Louises 226 Mullah- 227 Percivals 228 Rote 223 Wilsons 234 Stachelschweine (see also Quastenstachler) Nordafrika- 678 Südafrikanische 676 Strandgräber (see also Blessmulle, Erdbohrer, Graumull, Nacktmulle) Kap- 646 Namaqua- 644 Streifenmaus Basilios 423 Drei- 425 Ein- 427 Kamerun- 422 Millers 424

Veld-Ratte Bocages 364 Grants 367 Hindes 368 Kaisers 370 Namaqua 371 Nyika 373 Rote 365 Selinda 374 Tete 369 Thomas 376 Zinn 375 Vielzitzenmaus Awash- 462 Guinea- 464 Huberts 465 Kollmannspergers 467 Natal- 468 Shortridges 471 Südliche 463 Zwerg- 470 Vierstreifengrasmaus 545 Vierzehen-Jerboa 136 (see also Springmaus)

Togomaus Buettners 359

Waldbachmaus Afrikanische 390 Waldmaus 378 Waldmaus Allens 431 Angolische 433 Baers 433 Gebirgs- 434 Große 436 Kleine 437 Perlen- 430 Stella- 438 Walo 263 Wanderratte 540 Wasserratte Äthiopische 508 Weichhaarmaus Daltons 522 De Graaffs 524 Deroos 526 Dunkle 534 Hartwigs 527 Jacksons 528 Kamerun- 532

Kleine 530 Lukolele- 529 Misonnes 531 Muton- 533 Petters 535 Reizende 525 Tullbergs 537 Verschurens 538 Wald- 536 Wiesenmaus Angolische 503 Rauchgrau 504 Verreauxs 505 Weißschwanzhamster 201 Wildkaninchen (see also Buschkaninchen, Flusskaninchen, Hase, Rotkaninchen) Europäisches 708 Wollhaarratte Afrikanische 395 Angolanische 397 Foxs 391 Montane 396 Rote 399 Wühlmaus Musters 211 Zielscheibenratte 554 Zwerghörnchen (see also Baumhörnchen, Buschhörnchen, Dünnschwanzhörnchen, Palmenhörnchen, Riesenhörnschen, Sonnenhörnchen) Afrikanisches 71 Zwergmaus (see also Maus) Baoule- 476 Callewaerts 478 Gounda- Fluss 479 Graubauch- 497 Haussa- 480 Kleine 484 Kröten- 477 Mattheys 483 Mohammed- 482 Neaves 489 Orange 490 Oubangui- 491 Peters 492 Setzers 493 Westafrikanische 486 Wüsten- 481 Zarte 496 Zentralafrikanische 494 Zwergrennmaus Brauers 264 Dünen- 291 Paeba 288 Pinselschwanz- 293 Setzers 290

English names Acacia Rat, Black-tailed 559 Loring’s 558 Shortridge’s 562 Sundevall’s 561 Acacia Rats 556–563

African Climbing Mice 168–200 African Climbing Mouse, Banana 177 Brants’s 176 Cameroon 182 Chestnut 178

Grey 174 Kahuzi 172 Kivu 179 Lovat’s 173 Montane 171

Nikolaus’s Nyika 181 Vernay’s 183 African Dormice 109–134 (see also Garden Dormouse)

780

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English names

African Dormouse, Angolan 110 Christy’s 112 Flat-headed 130 Forest 124 Johnston’s 114 Kellen’s 116 Large Savanna 120 Lorraine’s 118 Monard’s 123 Nagtglas’s 126 Rupicolous 131 Short-eared 133 Spectacled 128 Thick-tailed 113 Ammodile 262, 263 Anomalure, Beecroft’s 604 Cameroon 615, 616 Lesser 610 Lord Derby’s 606 Pel’s 608 Anomalures 602–617 (see also Pygmy Anomalure) Bandicoot Rat, Short-tailed 506 Bandicoot Rats 506–508 Blesmol (see Cape Mole-rat) Blind Mole-rat 145 (see also Mole-rat) Middle East 145 Broad-headed Mice 567–571 Broad-headed Mouse, Hildegarde’s 568 Woosnam’s 569 Brush-furred Mouse, Rudd’s 258, 259 Brush-furred Rat, Black-clawed 251 Buff-bellied 249 Dieterlen’s 244 Eisentraut’s 245 Fire-bellied 252 Golden-footed 242 Grey 243 Hutterer’s 248 Medium-tailed 250 Rahm’s 253 Rosevear’s 254 Rusty-bellied 255 Short-tailed 240 Woosnam’s 257 Yellow-spotted 246 Brush-furred Rats 238–258 Brush-tailed Porcupine, African 672 Brush-tailed Porcupines 672–674 (see also Porcupine) Bush Squirrel, Alexander’s 74 Black-and-red 81 Boehm’s 75 Cooper’s 79 Green 85 Ochre 82 Red 84 Smith’s 77 Striped 80 Swynnerton’s 87 Vincent’s 88 Bush Squirrels 72–89 Cane Rat, Lesser 687 Greater 688 Cane Rats 685–690 Climbing Mouse (see also African Climbing Mouse) Bates’s 189, 190

Giant 188 Velvet 184, 185 Coypu 691 Creek Rat, Angolan 514 East African 515 Hopkins’s 517 Least 519 Ssese Islands 518 Creek Rats 513–519 Crested Porcupines 674–679 Dassie Rat (see Noki) Defua Rat 400 Dormice 102–134 Dormouse (see African Dormouse, Garden Dormouse) Dune Mole-rat, Cape 646 Namaqua 644 Dune Mole-rats 644–648 (see also Molerat) Dwarf Gerbil, Brauer’s 264 Ethiopian Rat, Grey-tailed 552 Rupp’s 553 White-footed 549 White-tailed 550 Ethiopian Rats 547–554 Fat Mice 191–200 Fat Mouse, Bocage’s 192 Common 199 Dainty 194 Jackson’s 195 Krebs’s 196 North-western 193 Pousargues’s 197 Tiny 198 Field Mice 377–379 Field Mouse, Long-tailed 378 Flying Squirrel (see Anomalure) Forest Mice 420–429 Forest Mouse, Basilio’s 423 Büttner’s 358, 359 Cameroon 422 Liberian 424 One-striped 427 Rwenzori 423 Three-striped 425 Garden Dormice 104–108 (see also African Dormouse) Garden Dormouse, Large-eared 105 Maghreb 107 Gerbil, Anderson’s 299 Berbera 297 Black-tailed 281 Boehm’s 272 Botta’s 300 Brockman’s 301 Bushveld 279 Cape 270 Charming 298 Cosens’s 304 Cushioned 324 Dunn’s 306 Dwarf 318 Flower’s 307 Fringe-tailed 284 Gambian 274 Gorongoza 277

Greater Egyptian 325 Guinea 276 Harwood’s 309 Henley’s 310 Highveld 273 Hoogstraal’s 312 Julian’s 313 Kemp’s 278 Khartoum 330 Lataste’s 314 Least 325 Lesser Egyptian 308 Lowe’s 315 Mackilligin’s 316 Maghreb 317 Moroccan 311 Nigerian 320 North African 301 Occidental 321 Pale 323 Percival’s 322 Phillips’s 283 Rock 328 Rosalind’s 327 Savanna 285 Simon’s 328 Somalian 330 Sudan 318 Tarabul’s 331 Wagner’s 304 Waters’s 322 Gerbils 268–286, 295–333 (see also Ammodile, Hairy-footed Gerbil, Dwarf Gerbil, Jird, Pygmy Gerbil, Sand Rat, Short-tailed Gerbil, Tateril) Giant Squirrel 89, 90 Forest 90 Grass Mice 441–455 Grass Mouse, Barbary 443 Bellier’s 444 Buffoon 448 Four-striped 544, 545 Griselda’s 445 Hoogstraal’s 446 Mittendorf’s 449 Rosevear’s 451 Senegal 447 Single-striped 449 Striated 452 Zebra 454 Grass Rat, Ansorge’s 382 Blick’s 383 Ethiopian 381 Nairobi 384 Neumann’s 386 Nile 387 Rufous 388 Grass Rats 379–389 Three-toed 499–502 Ground Squirrel, Barbary 42, 43 Cape 96 Damara 99 Striped 94 Unstriped 100 Ground Squirrels 93–101 Gundi, Atlas 630 Felou 634, 635 Mzab 636 Thomas’s 632 Gundis 628–638 (see also Pectinator)

Hairy-footed Gerbil, Brush-tailed 293 Dune 291 Pygmy 288 Setzer’s 290 Hairy-footed Gerbils 287–295 (see also Gerbils) Hare, Abyssinian 702 African Savanna 706 Cape 699 Ethiopian 701 Ethiopian Highland 705 Scrub 703 Stark’s 705 Hares 693, 694, 698–707 (see also Rock-hares and Rabbits) Jerboa, Four-toed 136 Greater Egyptian 141 Lesser Egyptian 138 Jerboas 135–142 Jird, Bushy-tailed 347 Fat-tailed 341 Libyan 336 Shaw’s 338 Sundevall’s 335 Jirds 333–338, 341–343, 347–349 (see also Gerbils) Link Rat, Rusty 235, 237 Maned Rat 212, 213, 214 Meadow Mice 502–506 Meadow Mouse, Angolan 503 Brockman’s 504 Verreaux’s 505 Mill Rat, Dybowski’s 500 King 501 Mill Rats 499–502 Mole-rat, Ansell’s 649 Bocage’s 650 Cape 662, 663 Damaraland 651 Darling’s 653 Fox’s 654 Giant 659 Hottentot 655 Kafue 658 Naked 667, 668 Ochre 660 Silvery 664, 665 Togo 661 Mole-rats 144–147, 641–670 (see also Dune Mole-rat, Blind Mole-rat, Root-rat) Mountain Squirrel, Carruthers’s 51 Cooper’s 79 Mouse (see also Grass Mouse, Pygmy Mouse and many others) Algerian 495 Bale 188 House 487 Long-eared 186 Velvet 390 Multimammate Mice 460–472 Multimammate Mouse, Awash 462 Dwarf 470 Guinea 464 Hubert’s 465 Kollmannsperger’s 467 Natal 468

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Shortridge’s 471 Southern African 463 Noki 680, 681 Old World Mice 473 Palm Squirrel 44 Western 44, 45 Pectinator 628, 638 Speke’s 638 Porcupine, Cape Crested 676 North African Crested 678 Porcupines 670–679 (see also Brush-tailed Porcupine) Pouched Mice 161–165 Pouched Mouse 153, 161 Cape 162 Mearns’s 164 Pouched Rat 153 Emin’s Giant 158 Gambian Giant 159 Long-tailed 154, 155 Pouched Rats, Giant 157–160 Pygmy Anomalure, Long-eared 612 Zenker’s 614 Pygmy Anomalures 611–615 (see also Anomalures) Pygmy Gerbil, Peel’s 339, 340 Pygmy Mouse 473 Baoule 476 Callewaert’s 478 Delicate 496 Desert 481 Gounda River 479 Grey-bellied 497 Hausa 480 Mahomet 482 Matthey’s 483 Neave’s 489 Orange 490 Oubangui 492 Peters’s 492 Setzer’s 493 Sorella 494 Tiny 484 Toad 477 West African 486 Pygmy Squirrel 70, 71 African 71 Rabbit, Bunyoro 710 European 708 Riverine 696 Rabbits 693, 694, 708–710 Rat, Black 541 Brown 540 Mount Oku 439, 440 Norway 540 Rats 539–543 (see also Defua Rat, Brush-

furred Rat and many others) Rock-hare, Jameson’s Red 714 Natal Red 713 Hewitt’s Red 717 Smith’s Red 715 Rock-hares 693, 694, 712–717 Rock Mice, Pygmy 203–209 Rock Mouse, Barbour’s Pygmy 204 Brukkaros Pygmy 207 Pygmy 205 Shortridge’s Pygmy 208 Rodents 27–692 Root-rat, African 151 Giant 149 Root-rats, African 147–152 Rope Squirrel, Carruthers’s 51 Congo 52 du Chaillu’s 54 Fire-footed 58 Kintampo 60 Lady Burton’s 55 Lunda 50 Red-cheeked 57 Ribboned 56 Thomas’s 49 Rope Squirrels 46–61 Rufous-nosed Rat, Common 510 West African 512 Rufous-nosed Rats 509–513 Sand Rat, Fat 344 Pale 346 Sand Rats 343–346 (see also Gerbil) Scrub Rat, Harrington’s 402 Yalden’s 403 Scrub Rats 402–404 Shaggy Rat, Angolan 397 Common 395 Fox’s 394 Montane 396 Rufous 399 Shaggy Rats 392–400 Short-tailed Gerbil, Cape 266, 267 Smoky Mouse, African 418 Soft-furred Mice 519–539 Soft-furred Mouse, Cameroon 532 Dalton’s 522 De Graaff’s 523 Delicate 524 De Roo’s 526 Hartwig’s 527 Jackson’s 527 Least 530 Lukolela 529 Misonne’s 531 Obscure 534 Petter’s 5350 Riverine 533 Tullberg’s 537 Verschuren’s 538

West African 536 Spiny Mice 217–235 Spiny Mouse Aïr 219 Cairo 220 Cape 233 Fiery 223 Golden 230 Grey 222 Johan’s 224 Kemp’s 225 Least 231 Louise’s 226 Mullah 227 Percival’s 228 Wilson’s 234 Springhare, Southern African 619 East African 614 Springhares 618–627 Squirrel, Aubinn’s 41 Grey 92 Slender-tailed 40, 41 Squirrels 38–101 (see also Ground Squirrel, Giant Squirrel, Palm Squirrel, Rope Squirrel, Sun Squirrel, Pygmy Squirrel, Bush Squirrel) Scaly-tailed 603–611 Striped Mouse, Ethiopian 472 Sun Squirrel, Gambian 62 Mutable 64 Punctate 65 Red-legged 66 Rwenzori 68 Small 65 Zanj 69 Sun Squirrels 61–70 Swamp Mouse, Delany’s 165, 166 Swamp Rat, Cansdale’s 456 Edwards’s 457 Long-footed 458 Swamp Rats 455–460

Jebel Marra 406 Kemp’s 564 Macmillan’s 416 Mozambique 409 Schouteden’s 565 Shining 414 Woodland 410 Thicket Rats 404–418, 563–567 Togo Mouse 359 Büttner’s 359 Tree Rats 556–563

Target Rat 554 Tateril, Congo 351 Emin’s 352 Lake Chad 354 Petter’s 355 Sand 350 Senegal 356 Slender 353 Tranier’s 357 Taterils 349–358 (see also Gerbils) Thicket Rat, Albertine Rift 411 Bunting’s 407 Charming 566 East African 413 Ethiopian 417 Giant 412 Grey-headed 407

Wading Rat, African 390 Water Rat, African 389, 390 Ethiopian 508 Whistling Rat, Brant’s 597 Littledale’s 600 Whistling Rats 571, 596–601 White-tailed Rat 201–203 African 201 Wood Mice 429–439 Wood Mouse, Allen’s 431 Angolan 433 Baer’s 433 Beaded 430 Large 436 Lesser 437 Montane 434 Stella 438

Veld Rat, Bocage’s 364 Grant’s 367 Hinde’s 368 Kaiser’s 370 Namaqua 371 Nyika 373 Red 365 Selinda 374 Tete 368 Thomas’s 376 West African 375 Veld Rats 362–377 Vlei Rat, Anchieta’s 576 Angoni 577 Barbour’s 579 Burton’s 580 Cuanza 581 Dent’s 582 Ethiopian 592 Karoo 594 Lake 585 Laminate 586 Saunders’s 588 Sloggett’s 589 Southern African 583 Tropical 591 Western 587 Vlei Rats 571, 574–596 Vole, Musters’s 211 Voles 211–212

Scientific names Acomys 217–235 airensis 219 cahirinus 220 cineraceus 222 ignitus 223 johannis 224

kempi 225 louisae 226 mullah 227 percivali 228 russatus 230 spinosissimus 231

subspinosus 233 wilsoni 234 Aethomys 362–377 bocagei 364 chrysophilus 365 granti 367

hindei 368 ineptus 369 kaiseri 370 namaquensis 371 nyikae 373 silindensis 374

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stannarius 375 thomasi 376 Allactaga 136–137 tetradactyla 136 Allosciurus 40–41 aubinnii 41 Ammodillus 262–264 imbellis 263 Anomaluridae 602–617 Anomalurus 603–611 beecrofti 604 derbianus 606 pelii 608 pusillus 610 Apodemus 377–379 sylvaticus 378 Arvicanthis 379–389 abyssinicus 381 ansorgei 382 blicki 383 nairobae 384 neumanni 386 niloticus 387 rufinus 388 Arvicolinae 210–212 Atherurus 672–674 africanus 672 Atlantoxerus 42–44 getulus 43 Bathyergidae 641–670 Bathyergus 644–648 janetta 644 suillus 646 Beamys 154–157 hindei 155 Bunolagus 696–697 monticularis 696 Colomys 389–392 goslingi 390 Cricetidae 210–215 Cricetomyinae 153–165 Cricetomys 157–161 emini 158 gambianus 159 Cryptomys 648–662 anselli 649 bocagei 650 damarensis 651 darlingi 653 foxi 654 hottentotus 655 kafuensis 658 mechowi 659 ochraceocinereus 660 zechi 661 Ctenodactylidae 628–640 Ctenodactylus 629–634 gundi 630 vali 632 Dasymys 392–400 foxi 394 incomtus 395 montanus 396 nudipes 397 rufulus 399 Delanymyinae 165–167 Delanymys 166–167 brooksi 166

Dendromurinae 168–200 Dendromus 169–184 insignis 171 kahuziensis 172 kivu 179 lovati 173 melanotis 174 mesomelas 176 messorius 177 mystacalis 178 nyasae 179 nyikae 181 oreas 182 ruppi 718 vernayi 182 Dendroprionomys 184–186 rousseloti 185 Deomyinae 217–260 Deomys 235–238 ferrugineus 237 Dephomys 400–402 defua 400 Desmodilliscus 264–266 braueri 264 Desmodillus 266–268 auricularis 267 Desmomys 402–404 harringtoni 402 yaldeni 403 Dipodidae 135–142 Eliomys 104–108 melanurus 105 munbyanus 107 Epixerus 44–46 ebii 45 Felovia 634–636 vae 635 Funisciurus 46–61 anerythrus 49 bayonii 50 carruthersi 51 congicus 52 duchaillui 54 Isabella 55 lemniscatus 56 leucogenys 57 pyrropus 58 substriatus 60 Georychus 662–664 capensis 663 Gerbillinae 260–358 Gerbilliscus 268–286 afra 270 boehmi 272 brantsii 273 gambianus 274 guineae 276 inclusus 277 kempi 278 leucogaster 279 nigricaudus 281 phillipsi 283 robustus 284 validus 285 Gerbillurus 287–295 paeba 288 setzeri 290 tytonis 291

vallinus 293 Gerbillus 295–333 acticola 297 amoenus 298 andersoni 299 bottai 300 brockmani 301 campestris 302 cosensi 304 dasyurus 304 dunni 306 floweri 307 gerbillus 308 harwoodi 309 henleyi 310 hesperinus 311 hoogstraali 312 juliani 313 latastei 314 lowei 315 mackilligini 316 maghrebi 317 nancillus 318 nanus 318 nigeriae 320 occiduus 321 percivali 322 perpallidus 323 pulvinatus 324 pusillus 325 pyramidum 325 rosalinda 327 rupicola 328 simoni 328 somalicus 330 stigmonyx 330 tarabuli 331 watersi 332 Gliridae 102–134 Grammomys 404–418 aridulus 406 brevirostris 718 buntingi 407 caniceps 408 cometes 409 dolichurus 410 dryas 411 gigas 412 ibeanus 413 kuru 414 macmillani 416 minnae 417 Graphiurus 109–134 angolensis 110 christyi 112 crassicaudatus 113 johnstoni 114 kelleni 116 lorraineus 118 microtis 120 monardi 123 murinus 124 nagtglasii 126 ocularis 128 platyops 130 rupicola 131 surdus 133 Heimyscus 418–420 fumosus 418 Heliophobius 664–667

argenteocinereus 665 Heliosciurus 61–70 gambianus 62 mutabilis 64 punctatus 65 rufobrachium 66 ruwenzorii 68 undulatus 69 Heterocephalus 667–670 glaber 668 Hybomys 420–429 badius 422 basilii 423 lunaris 423 planifrons 424 trivirgatus 425 univittatus 427 Hylomyscus 429–439 aeta 430 alleni 431 anselli 718 arcimontensis 718 baeri 433 carillus 433 denniae 434 endorobae 718 grandis 436 pamfi 718 parvus 437 stella 438 vulcanorum 718 walterverheyeni 718 Hystricidae 671–679 Hystrix 674–679 africaeaustralis 676 cristata 678 Idiurus 611–615 macrotis 612 zenkeri 614 Jaculus 137–142 jaculus 138 orientalis 141 Lagomorpha 693–717 Lamottemys 439–441 okuensis 440 Leimacomyinae 358–360 Leimacomys 359–360 buettneri 359 Lemniscomys 441–455 barbarus 443 bellieri 444 griselda 445 hoogstraali 446 linulus 447 macculus 448 mittendorfi 449 rosalia 449 roseveari 451 striatus 452 zebra 454 Leporidae 694–717 Lepus 698–707 capensis 699 fagani 701 habessinicus 702 saxatilis 703 starcki 705 victoriae 706

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Lophiomyinae 212–215 Lophiomys 213–215 imhausi 214 Lophuromys 238–258 brevicaudus 240 chercherensis 718 chrysopus 242 cinereus 243 dieterleni 244 eisentrauti 245 flavopunctatus 246 huttereri 248 kilonzoi 718 luteogaster 249 machangui 718 makundii 718 medicaudatus 250 melanonyx 251 menagesha 718 nudicaudus 252 pseudosikapusi 718 rahmi 253 roseveari 254 sabunii 718 sikapusi 255 stanleyi 718 woosnami 257 Malacomys 455–460 cansdalei 456 edwardsi 457 longipes 458 Malacothrix 186–188 typica 186 Massoutiera 636–638 mzabi 636 Mastomys 460–472 awashensis 462 coucha 463 erythroleucus 464 huberti 465 kollmannspergeri 467 natalensis 468 pernanus 470 shortridgei 471 Megadendromus 188–189 nikolausi 188 Meriones 333–338 crassus 335 libycus 336 sacramenti 335 shawi 338 Microdillus 339–341 peeli 340 Microtus 211–212 mustersi 211 Muriculus 472–473 imberbis 472 Muridae 216–601 Murinae 261–571 Mus 473–499 baoulei 476 bufo 477 callewaerti 478 goundae 479 haussa 480 indutus 481 mahomet 482 mattheyi 483 minutoides 484

musculoides 486 musculus 487 neavei 489 orangiae 490 oubanguii 491 setulosus 492 setzeri 493 sorella 494 spretus 495 tenellus 496 triton 497 Mylomys 499–502 dybowskii 500 rex 501 Myocastoridae 691–692 Myocastor 691–692 coypus 691 Myomyscus 502–506 angolensis 503 brockmani 504 verreauxii 505 Myosciurus 70–72 pumilio 71 Mystromyinae 201–203 Mystromys 201–203 albicaudatus 201 Nesokia 506–508 indica 506 Nesomyidae 153–209 Nilopegamys 508–509 plumbeus 508 Oenomys 509–513 hypoxanthus 510 ornatus 512 Oryctolagus 708–710 cuniculus 708 Otomyinae 571–601 Otomys 574–596 anchietae 576 angoniensis 577 barbouri 579 burtoni 580 cuanzensis 581 denti 582 dollmani 718 irroratus 583 lacustris 585 laminatus 586 occidentalis 587 orestes 718 saundersiae 588 sloggetti 589 tropicalis 591 typus 592 unisulcatus 594 uzungwensis 718 Pachyuromys 341–342 duprasi 341 Parotomys 596–601 brantsii 597 littledalei 600 Paraxerus 72–89 alexandri 74 boehmi 75 cepapi 77 cooperi 79 flavovittis 80

lucifer 81 ochraceus 82 palliatus 84 poensis 85 vexillarius 87 vincenti 88 Pectinator 638–640 spekei 638 Pedetes 618–627 capensis 619 surdaster 624 Pedetidae 618–627 Pelomys 513–519 campanae 514 fallax 515 hopkinsi 517 isseli 518 minor 519 Petromuridae 680–684 Petromus 681–684 typicus 681 Petromyscinae 203–209 Petromyscus 204–209 barbouri 204 collinus 205 monticularis 207 shortridgei 208 Poelagus 710–712 marjorita 710 Praomys 519–539 coetzeei 718 daltoni 522 degraaffi 523 delectorum 524 derooi 526 hartwigi 527 jacksoni 527 lukolelae 529 minor 530 misonnei 531 morio 532 mutoni 533 obscurus 534 petteri 535 rostratus 536 tullbergi 537 verschureni 538 Prionomys 189–191 batesi 190 Pronolagus 712–717 crassicaudatus 713 randensis 714 rupestris 715 saundersiae 717 Protoxerus 89–91 stangeri 90 Psammomys 343–346 obesus 344 vexillaris 346 Rattus 539–543 norvegicus 540 rattus 541 Rhabdomys 544–547 pumilio 545 Rodentia 27–692 Saccostomus 161–165 campestris 162 mearnsi 164

Sciuridae 38–101 Sciurus 92–93 carolinensis 92 Sekeetamys 347–349 calurus 347 Spalacidae 143–152 Spalacinae 144–147 Spalax 145–7 ehrenbergi 145 Steatomys 191–200 bocagei 192 caurinus 193 cuppedius 194 jacksoni 195 krebsii 196 opimus 197 parvus 198 pratensis 199 Stenocephalemys 547–554 albipes 549 albocaudata 550 griseicauda 552 ruppi 553 Stochomys 554–556 longicaudatus 554 Tachyoryctinae 147–152 Tachyoryctes 148–152 macrocephalus 149 splendens 151 Tatera, see Gerbilliscus Taterillus 349–358 arenarius 350 congicus 351 emini 352 gracilis 353 lacustris 354 petteri 355 pygargus 356 tranieri 357 Thallomys 556–563 loringi 558 nigricauda 559 paedulcus 561 shortridgei 562 Thamnomys 563–567 kempi 564 schoutedeni 565 venustus 566 Thryonomyidae 685–690 Thryonomys 686–690 gregorianus 687 swinderianus 688 Uranomys 258–260 ruddi 259 Xerus 93–101 erythropus 94 inauris 96 princeps 99 rutilus 100 Zelotomys 567–571 hildegardeae 568 woosnami 569 Zenkerella 615–617 insignis 616

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