Mammals of Africa Volume IV: Hedgehogs, Shrews and Bats 9781408122549, 9781472926944, 9781408189931

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mammals of africa volume IV

hedgehogs, shrews and bats

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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 Iv

hedgehogs, shrews and bats edited by meredith happold and 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) 978-1-4081-2254-9 ISBN (epdf) 978-1-4081-8993-1 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 IV: Happold, M. & Happold, D. C. D. (eds) 2013. Mammals of Africa.Volume IV: Hedgehogs, Shrews and Bats. Bloomsbury Publishing, London. Chapter/species profile: e.g. Yalden, D. W. & Happold, M. 2013. Otomops martiensseni Large-eared Giant Mastiff Bat; pp 554–556 in Happold, M. & Happold, D. C. D. (eds) 2013. Mammals of Africa:Volume IV. 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 Acknowledgements15 Acknowledgements for Volume IV16 Mammals of Africa: An Introduction and Guide – David Happold, Michael Hoffmann, Thomas Butynski & Jonathan Kingdon

17

Order ERINACEOMORPHA Hedgehogs – D. C. D. Happold

27

Family ERINACEIDAE Hedgehogs – D. C. D. Happold

29

GENUS Atelerix Hedgehogs – D. C. D. Happold Atelerix albiventris White-bellied Hedgehog (Four-toed Hedgehog) – D. C. D. Happold Atelerix algirus Algerian Hedgehog – D. C. D. Happold Atelerix frontalis Southern African Hedgehog – N. J. Dippenaar & R. M. Baxter Atelerix sclateri Somali Hedgehog – D. C. D. Happold

30

GENUS Hemiechinus Long-eared Hedgehog – D. C. D. Happold Hemiechinus auritus Long-eared Hedgehog – D. C. D. Happold

37

GENUS Paraechinus Desert Hedgehog – D. C. D. Happold Paraechinus aethiopicus Desert Hedgehog (Ethiopian Hedgehog) – D. C. D. Happold

39

31 33 34 36

37

39

Order SORICOMORPHA Shrews, Moles, Shrew Moles, Desmans and Solenodons – S. Churchfield

42

Family SORICIDAE Shrews – S. Churchfield

43

GENUS Congosorex Congo Shrews – R. Hutterer & W. T. Stanley Congosorex phillipsorum Phillips’s Congo Shrew – W. T. Stanley Congosorex polli Greater Congo Shrew – R. Hutterer Congosorex verheyeni Lesser Congo Shrew – P. Barrière & R. Hutterer

50 51 52

GENUS Crocidura Shrews (White-toothed Shrews) – D. C. D. Happold Crocidura aleksandrisi Cyrenaica Shrew – R. Hutterer Crocidura allex East African Highland Shrew – R. Hutterer Crocidura ansellorum Ansells’s Shrew – R. Hutterer Crocidura attila Hun Shrew (Cameroon Shrew) – P. D. Jenkins & S. Churchfield Crocidura baileyi Bailey’s Shrew (Simien Shrew) – L. A. Lavrenchenko Crocidura batesi Bates’s Shrew – J. C. Ray & R. Hutterer

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53 54 56 57 58 58 59 60

Crocidura bottegi Bottego’s Shrew – R. Hutterer Crocidura bottegoides Bale Shrew (Tricoloured Pygmy Shrew) – R. Hutterer Crocidura buettikoferi Büttikofer’s Shrew – D. C. D. Happold Crocidura caliginea African Dusky Shrew (Dusky Whitetoothed Shrew) – S. Churchfield, R. Hutterer & A. Dudu Crocidura cinderella Cinderella Shrew – R. Hutterer Crocidura congobelgica Congo Shrew (Congo Whitetoothed Shrew) – S. Churchfield, R. Hutterer & A. Dudu Crocidura crenata Jumping Shrew – R. Hutterer & P. Barrière Crocidura crossei Crosse’s Shrew – S. Churchfield & P. D. Jenkins Crocidura cyanea Reddish-grey Shrew – R. M. Baxter & N. J. Dippenaar Crocidura denti Dent’s Shrew (Dent’s White-toothed Shrew) – J. C. Ray & R. Hutterer Crocidura desperata Desperate Shrew – R. Hutterer Crocidura dolichura Long-tailed Shrew (Long-tailed Musk Shrew) – J. C. Ray & R. Hutterer Crocidura douceti Doucet’s Shrew (Doucet’s Musk Shrew) – R. Hutterer & D. C. D. Happold Crocidura eisentrauti Eisentraut’s Shrew – R. Hutterer Crocidura elgonius Elgon Shrew – W. T. Stanley Crocidura erica Heather Shrew (Angolan White-toothed Shrew) – P. D. Jenkins & S. Churchfield Crocidura fischeri Fischer’s Shrew – N. Oguge Crocidura flavescens Greater Red Shrew (Greater Red Musk Shrew) – R. M. Baxter & N. J. Dippenaar Crocidura floweri Flower’s Shrew – P. D. Jenkins & S. Churchfield Crocidura foxi Fox’s Shrew – J.-M. Duplantier & L. Granjon Crocidura fulvastra Savanna Shrew – S. Churchfield & P. D. Jenkins Crocidura fumosa Smoky Mountain Shrew (Smoky Whitetoothed Shrew) – P. D. Jenkins & S. Churchfield Crocidura fuscomurina Bicoloured Shrew (Bicoloured Musk Shrew, Tiny Musk Shrew) – N. J. Dippenaar & R. M. Baxter Crocidura glassi Glass’s Shrew (Ethiopian Mountain Shrew) – L. A. Lavrenchenko Crocidura goliath Goliath Shrew – R. Hutterer & D. C. D. Happold Crocidura gracilipes Short-footed Shrew (Peters’s Musk Shrew) – P. D. Jenkins & S. Churchfield Crocidura grandiceps Large-headed Shrew – R. Hutterer Crocidura grassei Grassé’s Shrew – R. Hutterer Crocidura greenwoodi Greenwood’s Shrew – P. D. Jenkins & S. Churchfield

61 62 62 63 64 65 66 67 68 69 70 71 72 73 74 75 75 76 78 78 79 80 81 82 83 84 85 85 86

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Contents

Crocidura harenna Harenna Shrew – R. Hutterer & D. W.Yalden Crocidura hildegardeae Hildegarde’s Shrew – W. T. Stanley Crocidura hirta Lesser Red Shrew (Lesser Red Musk Shrew) – R. M. Baxter & N. J. Dippenaar Crocidura jacksoni Jackson’s Shrew – N. Oguge Crocidura jouvenetae Jouvenet’s Shrew – D. C. D. Happold Crocidura kivuana Kivu Shrew – F. Dieterlen Crocidura lamottei Lamotte’s Shrew – P. D. Jenkins & S. Churchfield Crocidura lanosa Kivu Long-haired Shrew – F. Dieterlen Crocidura latona Latona’s Shrew – S. Churchfield, R. Hutterer & A. Dudu Crocidura littoralis Naked-tailed Shrew – J. C. Ray & R. Hutterer – Crocidura longipes Long-footed Shrew (Savanna Swamp Shrew) D. C. D. Happold Crocidura lucina Lucina’s Shrew – L. A. Lavrenchenko Crocidura ludia Ludia’s Shrew – R. Hutterer Crocidura luna Moonshine Shrew (Grey-brown Musk Shrew) – R. M. Baxter & N. J. Dippenaar Crocidura lusitania Mauritanian Shrew – L. Granjon & J.-M. Duplantier Crocidura macarthuri MacArthur’s Shrew – N. Oguge Crocidura macmillani Macmillan’s Shrew – L. A. Lavrenchenko Crocidura macowi Nyiro Shrew – S. Churchfield & P. D. Jenkins Crocidura manengubae Manenguba Shrew – R. Hutterer Crocidura maquassiensis Makwassie Shrew – R. M. Baxter & N. J. Dippenaar Crocidura mariquensis Swamp Shrew – R. M. Baxter & N. J. Dippenaar Crocidura maurisca Gracile Naked-tailed Shrew (Dark Shrew) – J. C. Kerbis Peterhans & S. O. Bober Crocidura monax Kilimanjaro Shrew (Rombo Shrew) – W. T. Stanley Crocidura montis Montane Shrew (Montane White-toothed Shrew) – R. Hutterer Crocidura muricauda West African Long-tailed Shrew – R. Hutterer Crocidura mutesae Ugandan Shrew (Ugandan Musk Shrew) – J. C. Ray & R. Hutterer Crocidura nana Somali Dwarf Shrew – S. Churchfield & P. D. Jenkins Crocidura nanilla Savanna Dwarf Shrew (Tiny Whitetoothed Shrew) – D. C. D. Happold Crocidura nigeriae Nigerian Shrew (Nigerian Musk Shrew) – S. Churchfield & P. D. Jenkins Crocidura nigricans Blackish Shrew (Blackish Whitetoothed Shrew) – P. D. Jenkins & S. Churchfield Crocidura nigrofusca African Black Shrew – R. Hutterer Crocidura nimbae Nimba Shrew – R. Hutterer Crocidura niobe Niobe’s Shrew – S. O. Bober & J. C. Kerbis Peterhans Crocidura obscurior West African Pygmy Shrew – R. Hutterer

87 88 89 90 91 92 93 94 95 96 97 97 98 99 100 101 101 102 103 104 105 106 107 108 109 110 111 112 112 113 114 115 116 117

Crocidura olivieri African Giant Shrew (Mann’s Musk Shrew, Euchareena’s Musk Shrew) – S. Churchfield & R. Hutterer Crocidura parvipes Small-footed Shrew – R. Hutterer Crocidura pasha Sahelian Tiny Shrew – S. Churchfield & P. D. Jenkins Crocidura phaeura Guramba Shrew – D. C. D. Happold & D. W.Yalden Crocidura picea Cameroon Shrew (Assumbo Shrew) – R. Hutterer Crocidura pitmani Pitman’s Shrew – S. Churchfield & P. D. Jenkins Crocidura planiceps Flat-headed Shrew – S. Churchfield & P. D. Jenkins Crocidura poensis Fraser’s Shrew (Fraser’s Musk Shrew) – S. Churchfield & R. Hutterer Crocidura polia Polia’s Shrew – R. Hutterer Crocidura raineyi Rainey’s Shrew – R. Hutterer Crocidura religiosa Egyptian Pygmy Shrew – D. C. D. Happold Crocidura roosevelti Roosevelt’s Shrew – D. C. D. Happold Crocidura russula Greater Shrew (Greater White-toothed Shrew) – S. Aulagnier & P. Vogel Crocidura selina Ugandan Lowland Shrew – R. Hutterer Crocidura silacea Lesser Grey-brown Shrew (Lesser Greybrown Musk Shrew) – R. M. Baxter & N. J. Dippenaar Crocidura smithii Desert Shrew (Desert Musk Shrew) – D. C. D. Happold Crocidura somalica Somali Shrew – R. Hutterer Crocidura stenocephala Kahuzi Swamp Shrew – F. Dieterlen Crocidura tansaniana Tanzanian Shrew (Amani Musk Shrew) – W. T. Stanley Crocidura tarella Tarella Shrew – P. D. Jenkins & S. Churchfield Crocidura tarfayensis Saharan Shrew (Tarfaya’s Shrew) – S. Aulagnier Crocidura telfordi Telford’s Shrew – W. T. Stanley Crocidura thalia Thalia’s Shrew – L. A. Lavrenchenko Crocidura theresae Therese’s Shrew – J.-M. Duplantier & L. Granjon Crocidura turba Turbo Shrew – N. Oguge Crocidura ultima Ultimate Shrew – S. Churchfield & P. D. Jenkins Crocidura usambarae Usambara Shrew – W. T. Stanley Crocidura viaria Savanna Path Shrew – R. Hutterer Crocidura virgata Mamfe Shrew – D. C. D. Happold & R. Hutterer Crocidura voi Voi Shrew – D. C. D. Happold Crocidura whitakeri Whitaker’s Shrew (Lesser Maghrebi Shrew) – S. Aulagnier Crocidura wimmeri Wimmer’s Shrew – S. Churchfield & P. D. Jenkins Crocidura xantippe Xanthippe’s Shrew (Yellow-footed Shrew) – W. T. Stanley Crocidura yankariensis Yankari Shrew – R. Hutterer Crocidura zaphiri Zaphir’s Shrew – S. Churchfield & P. D. Jenkins

118 120 121 121 122 123 124 125 126 127 127 128 129 130 131 132 133 134 135 135 136 137 138 139 139 140 141 142 143 144 144 145 146 147 148

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Contents

Crocidura zimmeri Upemba Shrew – R. Hutterer

148

GENUS Myosorex Mouse Shrews – R. Hutterer Myosorex babaulti Babault’s Mouse Shrew (Kivu Mouse Shrew) – F. Dieterlen Myosorex blarina Rwenzori Mouse Shrew (Mountain Mouse Shrew) – S. O. Bober & J. C. Kerbis Peterhans Myosorex cafer Dark-footed Mouse Shrew – R. M. Baxter & N. J. Dippenaar Myosorex eisentrauti Eisentraut’s Mouse Shrew – R. Hutterer Myosorex geata Geata Mouse Shrew – W. T. Stanley Myosorex kihaulei Kihaule’s Mouse Shrew – W. T. Stanley Myosorex longicaudatus Long-tailed Mouse Shrew – N. J. Dippenaar & R. M. Baxter Myosorex okuensis Oku Mouse Shrew – R. Hutterer Myosorex rumpii Rumpi Mouse Shrew – R. Hutterer Myosorex schalleri Schaller’s Mouse Shrew – R. Hutterer Myosorex sclateri Sclater’s Mouse Shrew – P. D. Jenkins & S. Churchfield Myosorex tenuis Thin Mouse Shrew (Transvaal Forest Shrew) – P. D. Jenkins & S. Churchfield Myosorex varius South African Mouse Shrew – R. M. Baxter & N. J. Dippenaar Myosorex zinki Kilimanjaro Mouse Shrew – R. Hutterer

149 150 151 152 153 154 155 156 157 158 158 159 160 161 163

GENUS Paracrocidura Large-headed Shrews – R. Hutterer164 Paracrocidura graueri Grauer’s Large-headed Shrew – R. Hutterer 164 Paracrocidura maxima Greater Large-headed Shrew – J. C. Kerbis Peterhans 165 Paracrocidura schoutedeni Schouteden’s Large-headed Shrew (Lesser Large-headed Shrew) – J. C. Ray & R. Hutterer 166 GENUS Ruwenzorisorex Rwenzori Shrew – R. Hutterer & J. C. Kerbis Peterhans Ruwenzorisorex suncoides Rwenzori Shrew – J. C. Kerbis Peterhans GENUS Scutisorex Armoured Shrew (Hero Shrew) – F. Dieterlen Scutisorex somereni Armoured Shrew (Hero Shrew) – F. Dieterlen & D. C. D. Happold GENUS Suncus Dwarf Shrews – D. C. D. Happold Suncus aequatorius Taita Dwarf Shrew – N. Oguge & R. Hutterer Suncus etruscus Etruscan Dwarf Shrew (Pygmy Whitetoothed Shrew) – S. Aulagnier & R. Fons Suncus hututsi Hutu-Tutsi Dwarf Shrew – J. C. Kerbis Peterhans Suncus infinitesimus Least Dwarf Shrew – R. M. Baxter & N. J. Dippenaar Suncus lixus Greater Dwarf Shrew – R. M. Baxter & N. J. Dippenaar Suncus megalura Climbing Dwarf Shrew – R. M Baxter & N. J. Dippenaar

Suncus murinus Asian House Shrew – J.-M. Duplantier Suncus remyi Remy’s Dwarf Shrew (Remy’s Pygmy Shrew) – D. C. D. Happold Suncus varilla Lesser Dwarf Shrew – R. M. Baxter & N. J. Dippenaar GENUS Surdisorex Mole-shrews – D. C. D. Happold & R. Hutterer Surdisorex norae Aberdare Mole-shrew – D. C. D. Happold Surdisorex polulus Mount Kenya Mole-shrew – D. C. D. Happold

179 181 181 183 183 184

GENUS Sylvisorex Forest Shrews – R. Hutterer 186 Sylvisorex camerunensis Cameroonian Forest Shrew – R. Hutterer186 Sylvisorex granti Grant’s Forest Shrew – F. Dieterlen 187 Sylvisorex howelli Howell’s Forest Shrew (Uluguru Forest Shrew) – W. T. Stanley 188 Sylvisorex isabellae Isabella Forest Shrew (Bioko Forest Shrew) – R. Hutterer 189 Sylvisorex johnstoni Johnston’s Forest Shrew (Pygmy Forest Shrew) – J. C. Ray & R. Hutterer 190 Sylvisorex konganensis Kongana Forest Shrew – J. C. Ray & R. Hutterer 191 Sylvisorex lunaris Moon Forest Shrew – D. C. D. Happold & F. Dieterlen 192 Sylvisorex morio Mount Cameroon Forest Shrew – D. C. D. Happold & R. Hutterer 193 Sylvisorex ollula Greater Forest Shrew – J. C. Ray & R. Hutterer 194 Sylvisorex oriundus Lesser Forest Shrew – R. Hutterer 195 Sylvisorex pluvialis Rainforest Shrew – D. C. D. Happold 196 Sylvisorex vulcanorum Volcano Forest Shrew (Volcano Shrew) – R. Hutterer 197 Order CHIROPTERA Bats – M. Happold

198

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Family PTEROPODIDAE Fruit Bats (Old World Fruit Bats) – M. Happold

223

169

GENUS Casinycteris Short-palated Fruit Bat – M. Happold Casinycteris argynnis Short-palated Fruit Bat – M. Happold

229 230

GENUS Eidolon Straw-coloured Fruit Bats – M. Happold Eidolon helvum African Straw-coloured Fruit Bat – D. Thomas & M. Henry

231

GENUS Epomophorus Epauletted Fruit Bats – M. Happold Epomophorus angolensis Angolan Epauletted Fruit Bat – P. J. Taylor Epomophorus anselli Ansell’s Epauletted Fruit Bat – W. Bergmans Epomophorus crypturus Peters’s Epauletted Fruit Bat – M. Happold Epomophorus gambianus Gambian Epauletted Fruit Bat – M. Happold Epomophorus grandis Sanborn’s Epauletted Fruit Bat – M. Happold

234

167

169 172 172 174 175 176 177 178

232

237 238 240 242 244 9

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Epomophorus labiatus Little Epauletted Fruit Bat – M. Happold Epomophorus minimus Least Epauletted Fruit Bat – M. Happold Epomophorus wahlbergi Wahlberg’s Epauletted Fruit Bat – M. Happold GENUS Epomops Singing Epauletted Fruit Bats – M. Happold Epomops buettikoferi Büttikofer’s Epauletted Fruit Bat – D. Thomas & M. Henry  Epomops dobsonii Dobson’s Epauletted Fruit Bat – M. Happold Epomops franqueti Franquet’s Epauletted Fruit Bat – M. Happold GENUS Hypsignathus Hammer-headed Fruit Bat – M. Happold Hypsignathus monstrosus Hammer-headed Fruit Bat – M. Happold GENUS Lissonycteris Angolan Soft-furred Fruit Bat – M. Happold Lissonycteris angolensis Angolan Soft-furred Fruit Bat – M. Happold GENUS Megaloglossus Woermann’s Long-tongued Fruit Bat – D. C. D. Happold & M. Happold Megaloglossus woermanni Woermann’s Long-tongued Fruit Bat – D. C. D. Happold GENUS Micropteropus Lesser Epauletted Fruit Bats – M. Happold Micropteropus intermedius Hayman’s Lesser Epauletted Fruit Bat – M. Happold Micropteropus pusillus Peters’s Lesser Epauletted Fruit Bat – D. Thomas & M. Henry GENUS Myonycteris Collared Fruit Bats – M. Happold Myonycteris relicta Bergmans’s Collared Fruit Bat – P. J. Taylor Myonycteris torquata Little Collared Fruit Bat – D. Thomas & M. Henry GENUS Nanonycteris Veldkamp’s Dwarf Epauletted Fruit Bat – J. Fahr Nanonycteris veldkampii Veldkamp’s Dwarf Epauletted Fruit Bat – J. Fahr GENUS Plerotes Anchieta’s Broad-faced Fruit Bat – M. Happold Plerotes anchietae Anchieta’s Broad-faced Fruit Bat (Benguela Fruit Bat) – M. Happold GENUS Pteropus Flying-foxes – M. Happold Pteropus seychellensis Seychelles Flying-fox – M. Happold Pteropus voeltzkowi Pemba Flying-fox – M. Happold & D. C. D. Happold

245

GENUS Rousettus Rousettes – M. Happold Rousettus aegyptiacus Egyptian Rousette – M. Happold Rousettus lanosus Long-haired Rousette – M. Happold

288 289 292

GENUS Scotonycteris Tear-drop Fruit Bats – J. Fahr Scotonycteris ophiodon Pohle’s Fruit Bat (Snake-toothed Fruit Bat) – J. Fahr Scotonycteris zenkeri Zenker’s Fruit Bat – J. Fahr

294

248 249 252 253

Family RHINOLOPHIDAE Horseshoe Bats – M. Happold & F. P. D. Cotterill

295 297 300

255 256 259 260 262 263 266 266 268 269 270 272 273 275 277 278 280 281 282 284 286

GENUS Rhinolophus Horseshoe Bats – M. Happold Rhinolophus adami Adam’s Horseshoe Bat – M. Happold Rhinolophus alcyone Halcyon Horseshoe Bat – M. Happold Rhinolophus blasii Blasius’s Horseshoe Bat (Peak-saddle Horseshoe Bat) – M. Happold Rhinolophus capensis Cape Horseshoe Bat – R. T. F. Bernard Rhinolophus clivosus Geoffroy’s Horseshoe Bat (Cretzschmar’s Horseshoe Bat) – R. T. F. Bernard & M. Happold Rhinolophus darlingi Darling’s Horseshoe Bat – F. P. D. Cotterill & M. Happold Rhinolophus deckenii Decken’s Horseshoe Bat – M. Happold Rhinolophus denti Dent’s Horseshoe Bat – F. P. D. Cotterill Rhinolophus eloquens Eloquent Horseshoe Bat – F. P. D. Cotterill Rhinolophus euryale Mediterranean Horseshoe Bat – J. Gaisler Rhinolophus ferrumequinum Greater Horseshoe Bat – J. Gaisler Rhinolophus fumigatus Rüppell’s Horseshoe Bat – F. P. D. Cotterill & M. Happold Rhinolophus guineensis Guinean Horseshoe Bat – J. Fahr Rhinolophus hildebrandtii Hildebrandt’s Horseshoe Bat – F. P. D. Cotterill & M. Happold Rhinolophus hilli Hill’s Horseshoe Bat – J. Fahr Rhinolophus hillorum Upland Horseshoe Bat – J. Fahr Rhinolophus hipposideros Lesser Horseshoe Bat – J. Gaisler Rhinolophus landeri Lander’s Horseshoe Bat – M. Happold Rhinolophus maclaudi Maclaud’s Horseshoe Bat – J. Fahr Rhinolophus maendeleo Maendeleo Horseshoe Bat – M. Happold Rhinolophus mehelyi Méhely’s Horseshoe Bat – J. Gaisler Rhinolophus ruwenzorii Rwenzori Horseshoe Bat – J. Fahr Rhinolophus sakejiensis Sakeji Horseshoe Bat – F. P. D. Cotterill Rhinolophus silvestris Forest Horseshoe Bat – F. P. D. Cotterill Rhinolophus simulator Bushveld Horseshoe Bat – F. P. D. Cotterill & M. Happold Rhinolophus swinnyi Swinny’s Horseshoe Bat – F. P. D. Cotterill Rhinolophus ziama Ziama Horseshoe Bat – J. Fahr Family HIPPOSIDERIDAE Old World Leaf-nosed Bats – M. Happold

303 309 311 312 314 316 318 320 322 323 325 327 329 331 332 334 336 338 340 342 343 345 347 348 350 351 353 355 357

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Contents

GENUS Asellia Trident Leaf-nosed Bats – S. Aulagnier Asellia patrizii Patrizi’s Trident Leaf-nosed Bat – S. Aulagnier Asellia tridens Geoffroy’s Trident Leaf-nosed Bat – S. Aulagnier

360

GENUS Cloeotis Percival’s Trident Bat – M. Happold Cloeotis percivali Percival’s Trident Bat (Short-eared Trident Bat) – D. S. Jacobs

364

GENUS Hipposideros Old World Leaf-nosed Bats – M. Happold Hipposideros abae Aba Leaf-nosed Bat – M. Happold Hipposideros beatus Benito Leaf-nosed Bat – M. Happold Hipposideros caffer Sundevall’s Leaf-nosed Bat – R. T. F. Bernard & M. Happold Hipposideros camerunensis Cameroon Leaf-nosed Bat – M. Happold Hipposideros curtus Short-tailed Leaf-nosed Bat – M. Happold Hipposideros cyclops Cyclops Leaf-nosed Bat – J. Fahr Hipposideros fuliginosus Sooty Leaf-nosed Bat (Temminck’s Leaf-nosed Bat) – J. Fahr Hipposideros gigas Giant Leaf-nosed Bat – M. Happold Hipposideros jonesi Jones’s Leaf-nosed Bat – J. Fahr Hipposideros lamottei Lamotte’s Leaf-nosed Bat – J. Fahr Hipposideros marisae Aellen’s Leaf-nosed Bat – J. Fahr Hipposideros megalotis Large-eared Leaf-nosed Bat – M. Happold Hipposideros ruber Noack’s Leaf-nosed Bat – M. Happold Hipposideros vittatus Striped Leaf-nosed Bat – M. Happold

362

367 372 373 375 378 379 380 383 385 387 389 391 392 393 395

Family MEGADERMATIDAE False Vampire Bats – M. Happold

401

GENUS Cardioderma Heart-nosed Bat – M. Happold Cardioderma cor Heart-nosed Bat (African False Vampire Bat) – M. Happold

403

GENUS Lavia Yellow-winged Bat – M. Happold Lavia frons Yellow-winged Bat – M. Happold

406 406

Family EMBALLONURIDAE Sheath-tailed Bats – M. Happold

GENUS Coleura African Sheath-tailed Bats – M. Happold Coleura afra African Sheath-tailed Bat – M. Happold

421 422

Subfamily Taphozoinae Pouched Bats and Tomb Bats – M. Happold

424

365

398 399

GENUS Rhinopoma Mouse-tailed Bats – S. Aulagnier Rhinopoma hardwickii Lesser Mouse-tailed Bat – S. Aulagnier Rhinopoma macinnesi MacInnes’s Mouse-tailed Bat – S. Aulagnier Rhinopoma microphyllum Greater Mouse-tailed Bat – S. Aulagnier

421

360

GENUS Triaenops Trident Bats – M. Happold Triaenops afer African Trident Bat – M. Happold

Family RHINOPOMATIDAE Mouse-tailed Bats – S. Aulagnier

Subfamily Emballonurinae Sheath-tailed Bats, Sacwinged Bats, Ghost Bats and others – M. Happold

404

409 410 412

GENUS Saccolaimus Pouched Bats – M. Happold Saccolaimus peli Pel’s Pouched Bat (Giant Pouched Bat, Black-hawk Bat) – J. Fahr

424

GENUS Taphozous Tomb Bats – M. Happold Taphozous hamiltoni Hamilton’s Tomb Bat – M. Happold Taphozous hildegardeae Hildegarde’s Tomb Bat – A. McWilliam & M. Happold Taphozous mauritianus Mauritian Tomb Bat – M. Happold Taphozous nudiventris Naked-rumped Tomb Bat – M. Happold Taphozous perforatus Egyptian Tomb Bat – P. J. Taylor

427 428

Family NYCTERIDAE Slit-faced Bats – M. Happold

438

GENUS Nycteris Slit-faced Bats – M. Happold Nycteris arge Bates’s Slit-faced Bat – J. Fahr Nycteris aurita Andersen’s Slit-faced Bat – V. Van Cakenberghe & M. Happold Nycteris gambiensis Gambian Slit-faced Bat – M. Happold Nycteris grandis Large Slit-faced Bat – M. Happold Nycteris hispida Hairy Slit-faced Bat – M. Happold Nycteris intermedia Intermediate Slit-faced Bat – J. Fahr Nycteris macrotis Large-eared Slit-faced Bat – F. P. D. Cotterill & M. Happold Nycteris major Dja Slit-faced Bat (Ja Slit-faced Bat) – J. Fahr Nycteris nana Dwarf Slit-faced bat – J. Fahr Nycteris parisii Parisi’s Slit-faced Bat – F. P. D. Cotterill Nycteris thebaica Egyptian Slit-faced Bat – R. T. F. Bernard & M. Happold Nycteris vinsoni Vinson’s Slit-faced Bat – M. Happold Nycteris woodi Wood’s Slit-faced Bat – F. P. D. Cotterill

440 442

Family MOLOSSIDAE Free-tailed Bats – M. Happold & F. P. D. Cotterill GENUS Mormopterus Little Mastiff Bats and others – M. Happold Mormopterus acetabulosus and M. francoismoutoui Mauritian Little Mastiff Bat and Réunion Little Mastiff Bat – M. Happold

425

429 431 434 436

444 445 446 448 450 451 453 455 456 457 460 461 464 472 473

414 415 418

GENUS Myopterus Winged-mouse Bats – J. Fahr Myopterus daubentonii Daubenton’s Winged-mouse Bat – J. Fahr Myopterus whitleyi Bini Winged-mouse Bat (Whitley’s Winged-mouse Bat) – J. Fahr

475 476 478 11

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GENUS Otomops Giant Mastiff Bats – M. Happold Otomops martiensseni Large-eared Giant Mastiff Bat – D. W.Yalden & M. Happold

479

GENUS Platymops Peters’s Flat-headed Bat – M. Happold Platymops setiger Peters’s Flat-headed Bat – M. Happold

483 483

GENUS Sauromys Roberts’s Flat-headed Bat – M. Happold Sauromys petrophilus Roberts’s Flat-headed Bat – F. P. D. Cotterill

485

GENUS Tadarida Tadarine Free-tailed Bats – M. Happold Tadarida aegyptiaca Egyptian Free-tailed Bat – R. T. F. Bernard & M. Happold Tadarida aloysiisabaudiae Duke of Abruzzi’s Free-tailed Bat – J. Fahr Tadarida ansorgei Ansorge’s Free-tailed Bat – F. P. D. Cotterill Tadarida bemmeleni Gland-tailed Free-tailed Bat – J. Fahr Tadarida bivittata Spotted Free-tailed Bat – F. P. D. Cotterill Tadarida brachyptera Short-winged Free-tailed Bat – M. Happold Tadarida chapini Pale Free-tailed Bat (Chapin’s Free-tailed Bat, Long-crested Free-tailed Bat) – M. Happold & F. P. D. Cotterill Tadarida condylura Angolan Free-tailed Bat – M. Happold Tadarida congica Congo Free-tailed Bat – J. Fahr Tadarida demonstrator Mongalla Free-tailed Bat – J. Fahr Tadarida fulminans Madagascan Free-tailed Bat (Malagasy Free-tailed Bat) – F. P. D. Cotterill Tadarida gallagheri Gallagher’s Free-tailed Bat – F. P. D. Cotterill Tadarida lobata Big-eared Free-tailed Bat – F. P. D. Cotterill Tadarida major Lappet-eared Free-tailed Bat – M. Happold Tadarida midas Midas Free-tailed Bat – F. P. D. Cotterill & M. Happold Tadarida nanula Dwarf Free-tailed Bat – M. Happold Tadarida niangarae Niangara Free-tailed Bat – M. Happold Tadarida nigeriae Nigerian Free-tailed Bat – F. P. D. Cotterill & M. Happold Tadarida niveiventer White-bellied Free-tailed Bat – F. P. D. Cotterill Tadarida petersoni Peterson’s Free-tailed Bat – M. Happold Tadarida pumila Little Free-tailed Bat – M. Happold Tadarida russata Russet Free-tailed Bat – M. Happold Tadarida spurrelli Spurrell’s Free-tailed Bat – M. Happold Tadarida teniotis European Free-tailed Bat – C. Ibáñez & R. Arlettaz Tadarida thersites Railer Free-tailed Bat (Railer Bat) – M. Happold Tadarida trevori Trevor’s Free-tailed Bat – M. Happold Tadarida ventralis Giant Free-tailed Bat – F. P. D. Cotterill

487

Family VESPERTILIONIDAE Vesper Bats – M. Happold

541

480

486

490 493 495 497 499 501 503 505 507 509 511 513 515 516 518 520 522 523 525 526 528 530 532 533 535 537 539

Subfamily VESPERTILIONINAE Barbastelles, Serotines, Butterfly Bats, Long-eared Bats, Noctules, Pipistrelles, House Bats and others – M. Happold

545

GENUS Barbastella Barbastelles – M. Happold Barbastella barbastellus Western Barbastelle – A. Sierro Barbastella leucomelas Eastern Barbastelle – M. Happold

546 547 549

GENUS Eptesicus Serotines – V. Van Cakenberghe & M. Happold Eptesicus bottae Botta’s Serotine – V. Van Cakenberghe & M. Happold Eptesicus floweri Horn-skinned Serotine – V. Van Cakenberghe & M. Happold Eptesicus hottentotus Long-tailed Serotine – F. P. D. Cotterill & M. Happold Eptesicus platyops Lagos Serotine – V. Van Cakenberghe & M. Happold Eptesicus serotinus Common Serotine – S. Aulagnier

550 552 554 555 557 558

GENUS Glauconycteris Butterfly Bats – M. Happold Glauconycteris alboguttata Striped Butterfly Bat – M. Happold Glauconycteris argentata Common Butterfly Bat – M. Happold Glauconycteris beatrix Beatrix Butterfly Bat – M. Happold Glauconycteris curryae Curry’s Butterfly Bat – J. Eger Glauconycteris egeria Bibundi Butterfly Bat – M. Happold Glauconycteris gleni Glen’s Butterfly Bat – M. Happold Glauconycteris humeralis Spotted Butterfly Bat – M. Happold Glauconycteris kenyacola Kenyacola Butterfly Bat – M. Happold Glauconycteris machadoi Machado’s Butterfly Bat – M. Happold Glauconycteris poensis Abo Butterfly Bat – M. Happold Glauconycteris superba Pied Butterfly Bat (Superb Butterfly Bat) – J. Fahr Glauconycteris variegata Variegated Butterfly Bat – M. Happold

560

GENUS Laephotis African Long-eared Bats – T. Kearney Laephotis angolensis Angolan Long-eared Bat – T. Kearney Laephotis botswanae Botswanan Long-eared Bat – T. Kearney Laephotis namibensis Namibian Long-eared Bat – T. Kearney Laephotis wintoni de Winton’s Long-eared Bat – T. Kearney

578 580 581 583 584

GENUS Mimetillus Moloney’s Mimic Bat – M. Happold Mimetillus moloneyi Moloney’s Mimic Bat (Moloney’s Flatheaded Bat) – J. Fahr

585

563 564 566 567 568 569 570 572 573 574 575 576

586

GENUS Nyctalus Noctules – S. Aulagnier 589 Nyctalus lasiopterus Giant Noctule – C. Ibáñez 590 Nyctalus leisleri Leisler’s Noctule (Leisler’s Bat) – S. Aulagnier592 GENUS Nycticeinops Schlieffen’s Twilight Bat – M. Happold Nycticeinops schlieffeni Schlieffen’s Twilight Bat (Schlieffen’s Bat) – M. Happold

594 595

12

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GENUS Otonycteris Hemprich’s Desert Bat – M. Happold 597 Otonycteris hemprichii Hemprich’s Desert Bat – I. Horáček598

Plecotus gaisleri Gaisler’s Long-eared Bat – P. Benda & S. Aulagnier

GENUS Pipistrellus Pipistrelles – V. Van Cakenberghe & M. Happold 600 Pipistrellus aero Mt Gargues Pipistrelle – V. Van Cakenberghe & M. Happold 608 Pipistrellus anchietae Anchieta’s Pipistrelle – T. Kearney 610 Pipistrellus ariel Fairy Pipistrelle – V. Van Cakenberghe & M. Happold 611 Pipistrellus brunneus Dark-brown Pipistrelle – J. Fahr 613 Pipistrellus capensis Cape Pipistrelle – T. Kearney 614 Pipistrellus crassulus Broad-headed Pipistrelle – J. Fahr 617 Pipistrellus deserti Desert Pipistrelle – V. Van Cakenberghe & P. Benda 619 Pipistrellus eisentrauti Eisentraut’s Pipistrelle – V. Van Cakenberghe & M. Happold 621 Pipistrellus grandidieri Yellow Pipistrelle – V. Van Cakenberghe & M. Happold 623 Pipistrellus guineensis Guinean Pipistrelle – V. Van Cakenberghe & M. Happold 624 Pipistrellus hanaki Hanák’s Pipistrelle – S. Aulagnier & P. Benda 626 Pipistrellus cf. helios Samburu Pipistrelle – M. Happold & V. Van Cakenberghe 627 Pipistrellus hesperidus Dusk Pipistrelle – T. Kearney 629 Pipistrellus inexspectatus Aellen’s Pipistrelle – V. Van Cakenberghe & M. Happold 631 Pipistrellus kuhlii Kuhl’s Pipistrelle – V. Van Cakenberghe & P. Benda 633 Pipistrellus cf. melckorum Melcks’s Pipistrelle – T. Kearney 635 Pipistrellus musciculus Mouse-like Pipistrelle – 637 V. Van Cakenberghe & M. Happold Pipistrellus nanulus Tiny Pipistrelle – V. Van Cakenberghe & M. Happold 638 Pipistrellus nanus Banana Pipistrelle (Banana Bat) – M. Happold 639 Pipistrellus permixtus Dar-es-Salaam Pipistrelle – V. Van Cakenberghe & M. Happold 642 Pipistrellus pipistrellus Common Pipistrelle – S. Aulagnier 643 Pipistrellus rendalli Rendall’s Pipistrelle – V. Van Cakenberghe & M. Happold 645 Pipistrellus rueppellii Rüppell’s Pipistrelle – M. Happold 647 Pipistrellus rusticus Rustic Pipistrelle (Rusty Pipistrelle) – T. Kearney 649 Pipistrellus savii Savi’s Pipistrelle – S. Aulagnier 651 Pipistrellus somalicus Somali Pipistrelle (Somali Serotine) – V. Van Cakenberghe & M. Happold 653 Pipistrellus tenuipinnis White-winged Pipistrelle (Slenderwinged Pipistrelle) – J. Fahr 655 Pipistrellus zuluensis Zulu Pipistrelle (Aloe Bat) – M. Happold, V. Van Cakenberghe & T. Kearney 657

GENUS Scotoecus Lesser House Bats – M. Happold Scotoecus albofuscus Light-winged Lesser House Bat (Gambian Lesser House Bat) – M. Happold Scotoecus hirundo Dark-winged Lesser House Bat (Swallow-like Lesser House Bat) – M. Happold

GENUS Plecotus Long-eared Bats – S. Aulagnier Plecotus balensis Bale Long-eared Bat – L. A. Lavrenchenko Plecotus christii Christie’s Long-eared Bat – P. Benda & S. Aulagnier

660 661 663

GENUS Scotophilus House Bats – V. Van Cakenberghe & M. Happold Scotophilus dinganii Yellow-bellied House Bat – M. Happold Scotophilus leucogaster White-bellied House Bat – V. Van Cakenberghe & M. Happold Scotophilus nigrita Giant House Bat – M. Happold Scotophilus nucella Robbins’s House Bat – V. Van Cakenberghe & M. Happold Scotophilus nux Nut-coloured House Bat – V. Van Cakenberghe & M. Happold Scotophilus viridis Green House Bat – V. Van Cakenberghe & M. Happold

664 666 667 669 672 674 676 678 680 681 682

Subfamily MYOTINAE Wing-gland Bats and Myotises – M. Happold

684

GENUS Cistugo Wing-gland Bats – T. Kearney Cistugo lesueuri Lesueur’s Wing-gland Bat – T. Kearney Cistugo seabrae Angolan Wing-gland Bat – T. Kearney

685 685 687

GENUS Myotis Myotises (Mouse-eared Bats, Hairy Bats) – M. Happold Myotis bocagii Rufous Myotis (Rufous Mouse-eared Bat) – M. Happold Myotis capaccinii Long-fingered Myotis – S. Aulagnier & E. Cosson Myotis dieteri Dieter’s Myotis – M. Happold Myotis emarginatus Geoffroy’s Myotis – S. Aulagnier Myotis morrisi Morris’s Myotis – D. W.Yalden Myotis mystacinus Whiskered Myotis (Whiskered Bat) – S. Aulagnier Myotis nattereri Natterer’s Myotis (Natterer’s Bat) – S. Aulagnier Myotis punicus Maghreb Myotis – M. Ruedi & R. Arlettaz Myotis scotti Scott’s Myotis – D. W.Yalden Myotis tricolor Temminck’s Myotis (Temminck’s Hairy Bat) – R. T. F. Bernard Myotis welwitschii Welwitsch’s Myotis – M. Happold Subfamily MINIOPTERINAE Long-fingered Bats – M. Happold GENUS Miniopterus Long-fingered Bats – M. Happold Miniopterus fraterculus Lesser Long-fingered Bat – R. T. F. Bernard & M. Happold Miniopterus inflatus Greater Long-fingered Bat – M. Happold Miniopterus minor Least Long-fingered Bat – M. Happold

688 692 694 696 697 699 700 702 703 705 706 708 710 711 712 714 716 13

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Contents

Miniopterus natalensis Natal Long-fingered Bat (Natal Clinging Bat) – R. T. F. Bernard & M. Happold Miniopterus schreibersii Schreibers’s Long-fingered Bat – J. Eger

718

GENUS Phoniscus Trumpet-eared Bats – M. Happold Phoniscus aerosa Dubious Trumpet-eared Bat – M. Happold

721

Appendix: New Taxa 2005–2010736

Subfamily KERIVOULINAE Woolly Bats – M. Happold

723

Glossary737

GENUS Kerivoula Woolly Bats – M. Happold Kerivoula africana Tanzanian Woolly Bat – M. Happold Kerivoula argentata Damara Woolly Bat – F. P. D. Cotterill Kerivoula cuprosa Copper Woolly Bat – J. Fahr Kerivoula eriophora Heuglin’s Woolly Bat – D. W.Yalden Kerivoula lanosa Lesser Woolly Bat – F. P. D. Cotterill Kerivoula phalaena Spurrell’s Woolly Bat – J. Fahr Kerivoula smithii Smith’s Woolly Bat – J. Fahr

724 725 726 727 729 730 731 733

Bibliography752

734 734

Authors of Volume IV

789

Indexes French names German names English names Scientific names

792 794 796 798

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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:

15

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Acknowledgements for Volume IV Meredith Happold and D. C. D. Happold

We owe a great debt of gratitude to the many people who have helped and encouraged us with our research on African small mammals for nearly 50 years and hence have contributed enormously to this volume. When we were beginning our research, the following colleagues were marvellous sources of inspiration: Frank Ansell†, Fritz Dieterlen, John Edwards Hill†, Dieter Kock, Karl Koopman†, Erwin Kulzer, Waldo Meester†, Francis Petter†, Don Rosevear†, Hank Setzer† and Reay Smithers†. We 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. We are also extremely grateful to our colleagues (many of whom are authors of profiles in this volume) who have given so 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, we have visited museums whenever the opportunity has occurred, and we are most grateful to the curators and their assistants who have helped during our visits: David Harrison, Paul Bates and Malcolm Pearch (Harrison Zoological Institute, Sevenoaks, England), Gabor Csorba (Hungarian Natural History Museum), Francis Petter† and Jacques Cuisin (Museum National d’Histoire Naturelle, Paris, France), Lydia Kigo (National Museum of Kenya, Nairobi, Kenya), Frederike Spitzenberger and Barbara Herzig (Naturhistorisches Museum, Vienna, Austria), Wim Wendelen (Royal Museum of Central Africa, Tervuren, Belgium), Judith Eger (Royal Ontario Museum, Toronto, Canada), Dieter Kock (Senkenberg Museum, Frankfurt, Germany), Fritz Dieterlen (Staatliches Museum für Naturkunde, Stuttgart, Germany), Teresa Kearney and Christian Chimimba (Transvaal Museum, Pretoria, South Africa) and Rainer Hutterer (Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany). We owe a special debt of gratitude to the Natural History Museum, London, England (formerly British Museum of Natural History), especially Gordon Corbet, John Edwards Hill†, Robert Hayman†, Paula Jenkins, Jean Ingles, Daphne Hills and Louise Tomsett who have allowed us (since 1964) to examine the collections and have assisted us in numerous ways. Particular thanks also are due to Michael and Mona Ensor and Sara Churchfield in London, the Ostermeyer and Schlöder families in Bonn, our friends in Corniche Verte, Brussels, and Mike and Anne-Marie Swift in Nairobi, who provided us with welcoming homes and sustenance after our busy days in museums.

We wish to thank the various universities, where one or both of us has been employed, for their support for our 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 our more recent work in Malawi. 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 them for their dedication and perseverance, and for accepting that many of the sections (especially the descriptions) needed heavy editing to ensure that the profiles became uniform in terminology and content, compatible and comparable, and hence functional and unambiguous. Our thanks are also due to the many reviewers (listed here in alphabetical order) who reviewed profiles for this volume: Stéphane Aulganier, Paul Bates, Rod Baxter, Petr Benda, Nigel Bennett, Wim Bergmans, Gary Bronner, Charles Calisher, Gabor Csorba, J. F. Dahl, Jan Decher, Nico Dippenaar, Njikoha Ebigbo, Judith Eger, Jakob Fahr, Roger Fons, Jiří Gaisler, Norberto Giannini, Werner Haberl, Rainer Hutterer, David Jacobs, Jenny Jarvis, Teresa Kearney, Julian Kerbis Peterhans, Kazimierz Kowalski, Barbara RzebikKowalska, Dieter Kock, Erwin Kulzer, Emile Lecompte, Anthony Maddock, Conrad Matthee, Françoise Poitevin, Nigel Reeve, Manuel Ruedi, Duane Schlitter, Nancy Simmons, Katja Soer, Sandy Sowler, Peter Taylor, Michel Thévenot, Erik Thorn, Don Thomas,Yves Tupinier, Victor Van Cakenberghe, Peter Vogel and Johan Watson. We also acknowledge, with deep gratitude, all who have provided specialist knowledge and advice: in particular, Paul Bates, Wim Bergmans, Patrick Boland, Gary Bronner, Charles Calisher, Mike Carleton, Laurent Granjon, Colin Groves, David Harrison, Tony Hutson, Dieter Kock, Ernest Seamark, Nancy Simmons, Victor Van Cakenberghe and Marianne Volleth. We also thank Stéphane Aulagnier who provided the French names of bats, Jakob Fahr and Cornelia Rumpp who provided the German names of bats, and Anke Hoffman who provided many of the German names of other small mammals. We are extremely grateful 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 Series Acknowledgements. We are also especially grateful to Zoe Cokeliss who digitized all the distribution maps of the small mammals.

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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 serve as a guide to identification. Africa has changed greatly in recent decades because of increases in human populations, exploitation of natural resources, agricultural development and urban expansion. Throughout the continent, extensive areas of forest have been destroyed and much of the forest that remains is degraded and fragmented. Savanna habitats have been altered by felling of trees and development for agriculture. Many of the drier areas are threatened with desertification. As a result, the abundance and geographic ranges of many species of mammals have declined – some marginally, some catastrophically, some to

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.

extinction. Hence, it seems appropriate that our knowledge of each 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: they also contain 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 17

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An Introduction and Guide

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

Contents

Number of species

Editors

I

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

49

II

Primates

93

III

Rodents, Hares and Rabbits Hedgehogs, Shrews and Bats

408

V

Carnivores, Pangolins, Equids and Rhinoceroses

93

VI

Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer and Bovids

93

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

IV

380

account of the species. They have been edited by six editors who distributed their work according to the orders with which they were most familiar. Each editor chose authors who had extensive knowledge of the species (or higher taxon) and, preferably, had experience with the species in the field. Each volume follows the same general format with respect to arrangement, subheadings and contents. Because Mammals of Africa has contributions from 356 authors (each with a different background and speciality), and 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 ‘super-species’ (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 (2005), although authors have employed alternative taxonomies when there were good reasons for doing so. Volume I differs from the other volumes in that it contains a number of introductory chapters about Africa and its environment, and about African mammals in general.

The continent of Africa For the purposes of this work, ‘Africa’ is defined as the continent of Africa (bounded by the Mediterranean Sea, the Atlantic Ocean, the Indian Ocean, the Red Sea and the Suez Canal) and the islands on the continental shelf that, at some time in their history, have been joined to the African continent. The largest of the ‘continental islands’ are Zanzibar (Unguja), Mafia and Bioko (Fernando Po). All ‘oceanic islands’, e.g. São Tomé, Principe, Annobón (Pagulu), Madagascar, Comoros, Seychelles, Mauritius, Socotra, Canaries, Madeira and Cape Verde are excluded, with the exception of Pemba, which is included because of its close proximity (ca. 50 km) to the mainland. The names of the countries of Africa are taken from the Times Atlas (2005). The Republic of Congo is referred to as ‘Congo’ and the Democratic Republic of Congo (formerly 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 Provinces 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 together with their previous names that are used in the older literature on African mammals (Table 3). Africa is the second largest continent in the world (after Asia), but it differs from other continents (except Australia and Antarctica) in being essentially an island. At various times in the past, Africa has been joined to other continents – a situation that has had a strong influence on the fauna and flora of the continent. Africa is a vast continent (29,000,000 km², 11,200,000 mi²) that straddles the Equator, with about two-thirds of its area in the northern hemisphere and one-third in the southern hemisphere. As a result, Africa has many varied climates (with seasons in each hemisphere being 6 months out of phase), many habitats (including deserts, savannas, woodlands, swamps, rivers, lakes, moist forests, monsoon forests, mountains and glaciers), and altitudes ranging from 155 m (509 ft) below sea level at L. Assal, Djibouti, in the Danakil (Afar) Depression, to 5895 m (19,341 ft) on Mt Kilimanjaro, Tanzania. Africa is comprised of 47 countries, some of which are very large (e.g. Sudan [2,506,000 km²; 967,000 mi²], Algeria (2,382,000 km², 920,000 mi²], and Democratic Republic of Congo [2,345,000 km², 905,000 mi²]), and others that are relatively small (e.g. Djibouti [23,200 km², 9,000 sq miles], Swaziland [17,400 km², 6,700 mi²] and The Gambia [11,300 km², 4,400 mi²]). The human population of each country also varies greatly, from about 346/km² in Rwanda to only about 2.5/km² in Namibia. With its great size and varied habitats, Africa supports a high biodiversity, including a large number of species of mammals. Likewise, most countries have a high diversity of mammals (especially when compared with temperate countries). Africa may also be divided into biotic zones (Figure 2). A biotic zone (BZ) 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 exploited by each species of mammal 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 be explained

18

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The continent of Africa

0°

10°

a

30°

M

c oro

10°

co

20°

Tunisia

30°

30°

Western Sahara

le Ni

Algeria Libya

20°

Egypt 40°

Mauritania

Niger

r Nige

Chad

Burkina Faso

Somaliland Ethiopia

ia

South Sudan

al

a

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

Uganda

Congo

Kenya

Co

ng

o

10°

Central African Republic

So

Liberia

10°

an

Côte d’Ivoire

Djibouti

Nigeria

Gh

GuineaGuinea Bissau Sierra Leone

500 1000 km

0°

Pemba Zanzibar

Tanzania

Mafia

10°

10°

Angola

10°

Malawi

Zambia

qu

e

i bez am

bi

Z

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

Zimbabwe

20°

Namibia

am

500

Rwanda Burundi

50°

oz

0

Democratic Republic of Congo

M

0

50°

Eritrea

Sudan

m

Senegal The Gambia 10°

20°

Mali

Botswana

20° 40°

Swaziland

c

30°

30°

South Africa

Lesotho 30°

20°

le Ni Awa sh

W hite Nile

Tana

Za

Shire

e en un

e

Limpopo

Gauteng

North West

a um Ruv Lake Malawi

opo mp Li

Or

b

Free State Northern Cape

Eastern Cape Western Cape 0

ang e

KwaZulu– Natal

zi be m

Lake Kariba Okavango Delta

C

Mpumalanga

Rufiji

Lake Mweru Lake Bangweulu

o ng ba Cu

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

Lualaba

ili Kw o ang Kw

1000 miles

1000 km

i Lomam Sankuru Kasai

é

oou

500 500

a

a Og

0 0

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

Sangh

Cross

e nu Be Mt Cameroon aga San Ivindo

Lu an gw a

Lake Volta

Lake Tana

Ouban gui

Black Volta

olta ite V Wh

Lake Chad

ile eN Blu

gal

e Sen r Nige

0

300 miles 300 km

19

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An Introduction and Guide

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

Area (km2) ’000

Area (miles2) ’000

Human population ’000 (2006)

People per km2

2,382 1,247 113 582 274 27.8 475

920.0 481.0 43.0 225.0 106.0 10.7 184.0

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

14.1 12.7 77.0 3.1 49.6 280.5 36.2

623 1,284 342 322 2,345

241.0 496.0 132.0 125.0 905.0

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

6.9 5.8 10.8 61.2 26.7

23.2 1,001 28.1

9.0 387.0 10.8

800 75,400 500

34.5 75.3 17.8

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

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

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

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

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

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

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

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

56.8 164 236 753 391 29,448

21.9 63.0 91.0 291.0 151.0 11,383

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

110.9 61.6 117.4 15.8 33.5 56.8

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

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Species profiles

The hedgehogs, shrews and bats of Africa

1

This volume is devoted to the orders Erinaceomorpha (hedgehogs), Soricomorpha (shrews) and Chiroptera (bats). These orders comprise 380 species (i.e. about 34% of all African mammals). Two of the orders contain many species – Soricomorpha (150 species) and Chiroptera (224 species) – and are the second and third most speciose orders after the Rodentia (395 species) (see Table 1). The seven orders that are considered to be the ‘small mammals of Africa’ (orders Afrosoricida and Macroscelidea [Volume I], Rodentia and Lagomorpha [Volume III], and Erinaceomorpha, Soricomorpha and Chiroptera [this volume]) collectively comprise 74.1% (827 of 1116) of all African mammalian species. Many species in the orders described in this volume have not been studied in detail because of their rarity and small geographic ranges; however, there is considerable knowledge about many of the species of bats. There are two editors for this volume: Meredith Happold (Chiroptera; 224 species) and David C. D. Happold (Erinaceomorpha and Soricomorpha; 156 species). The profiles for Volume IV were submitted to the editors between 2001 and 2005. It has not been possible to revise profiles since then; however, notes have sometimes been added to draw attention to important changes in taxonomy and distribution, the IUCN Categories of threat in the conservation sections have been updated, and citations of papers previously given as ‘in press’ have been completed. An Appendix has been added listing new taxa described during the period 2005–2010.

2

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

4

7

5 6a

8 6

6b

6c 10 9

11a 11b

12 11c 13

Figure 2. The biotic zones of Africa.

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 subregions that reflect the different biogeographical distributions of species within the zone, each region/subregion 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 subregions of the Rainforest 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 is as follows:

r

ge Ni

WEST CENTRAL

Niger Delta

Gabon

Eastern Nigeria ga

a San

Gabon

EAST CENTRAL

SOUTH CENTRAL

i

go

n Co

0

500

1000

1500

2000 km

East Central

laba Lua

am Lom

Figure 3. The Rainforest Biotic Zone showing the regions, subregions 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. Subregions are indicated in lower case letters. Refugia are indicated in lower case italics and yellow (after Happold 1996 and references therein; see also Happold & Lock, Volume I, Mammals of Africa).

Ben

Cross

Ghanaian

Ivory Coast

ue

Western Nigeria

Ubangi

a

ra

Western

lt Vo

Sassand

Liberian WESTERN

South Central

21

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An Introduction and Guide

Scientific name (genus and species)  The currently accepted name of the species. Vernacular names English, French and German names are given, as available. The first given English name is the preferred vernacular name for the species; alternative names are given in parentheses for some species. Wilson & Cole (2000) list proposed vernacular names for all the world’s mammals; most of these names were also given in the third edition of Mammal Species of the World (Wilson & Reeder 2005). Although these works have been consulted, the names used have not always been adopted in Mammals of Africa. For the names of bats, there was collaboration between the authors of profiles, other bat specialists and Nancy Simmons (Simmons 2005): consequently the bat names, with very few exceptions, are the same in both publications. French names were either provided by Stéphane Aulagnier (bats), other profile authors, or taken from Gunther (2002). Most of the German names were provided by Jakob Fahr and Cornelia Rumpp (bats) and by Anke Hoffman (other taxa). 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 where the species was described, and the type locality (i.e. where the holotype [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. Major synonyms are listed (without the taxonomic authority unless essential for clarity), and the number of subspecies (usually only in Africa) is given: most of this information is 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. Description  This section, together with the illustrations and relevant tables, includes sufficient information to identify the species as well as describing characters that are relevant to the habits and life-style of the species. The section begins with a brief overall description of the species, including an indication of size. (For the bats, the first sentence lists the most useful diagnostic characters of the species, ending with those that distinguish it from its most similar species.) This is followed by a more detailed description of the external characters and skulls (including the diagnostic characters); the information given covers all subspecies (if any). 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. The table-keys (referred to as tables in the text) function as keys to the species, and should be read from the left column to the right column, which gives the name of the species thereby identified. The number and arrangement of nipples in adult females (for taxa other than bats) 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. Geographic Variation  Variation within the species may be clinal (without subspecies) or subspecific. If the variation is clinal, there is a description of the character(s) that alters clinally across the geographic range of the species. If the variation is subspecific, each subspecies is listed together with its geographic range and the characters that distinguish it from the other subspecies.

ear crown dorsal spines

forehead eye area of face mask

flank spines

nostril

mouth

tail

ventral p

elage

Digit 1 Digit 2

forelimb

Digit 3 Digit 4

hallux (Digit 1)

hindfoot

Digit 5

External characters of a hypothetical hedgehog.

22

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Species profiles

vibrissa

eye

outer surface of ear

long hairs

muzzle

tail

dorsal region

rhinarium nostril lips

al

ventr

chest forefoot forelimb

region

hindlimb dorsal surface of hindfoot

position of lateral gland

External characters of a hypothetical shrew.

Similar Species  A list of similar species is given together with the diagnostic characters and/or measurements most useful to distinguish that particular species from the profiled species. If a character of a similar species is described as being ‘larger’ or ‘smaller’ than that of the profiled species, there is no known overlap in the ranges of measurements of this character. In contrast, if the ranges overlap but the means are different, a character is said to be ‘larger on average’ or ‘smaller on average’. For the Order Chiroptera, similar species are considered to be those that share a specified combination of characters, whether or not they live sympatrically. For other orders, with some exceptions, the lists of similar species are restricted to those that are sympatric or parapatric with the profiled species; this section is omitted for the genus Crocidura because of the very large number of species (but see Table 8). Distribution  The first sentence is often ‘Endemic to Africa’ indicating that the species is found (in the wild) only in Africa. Alternatively, the section begins with the distribution in Africa, and the extralimital distribution is given at the end of the section. The Biotic Zone (or Zones) in which the species has been recorded are listed because this information indicates the sorts of environments exploited by that species, 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. Next, the distribution in African countries, or parts of countries, is described, and altitudinal ranges may be given. As a general rule, descriptions of the ranges of 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) may be recorded. Other information may include average annual rainfall, altitudinal limits and seasonal variation in habitat characteristics. Abundance  This section attempts to indicate the comparative abundance of the species. For many species, quantitative data are unavailable but the species can be assessed as ‘abundant’, ‘common’, ‘rare’, ‘rarely seen but often heard’, ‘rarely collected’ etc. For some species, abundance is indicated by quantitative 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 captured’). 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, which show how the species uniquely interacts with its environment, with conspecifics, and with other animals. This section may also describe species-specific adaptations for locomotion, burrowing, mechanisms for orientation, production of sound, sensory mechanisms and activity patterns. It may also include descriptions of domiciles and population movements (such as migration). 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 organization (e.g. solitary, social or colonial), group 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 and social vocalizations. Reproduction and Population Structure  If data are available, this section describes the reproductive strategy of the species, this being determined by the litter-size and the timing of reproductive events (i.e. the reproductive chronology). Reproductive chronologies cover the times of year when spermatogenesis, ovulation, copulation, gestation, parturition and lactation occur, and consequently this indicates the duration of pregnancy and lactation, and the number of pregnancies each female may have in one year. Reproductive chronologies give data for both individuals and local populations. Special adaptations such as reproductive delays (e.g. delayed implantation) and postpartum oestrus are mentioned, and 23

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An Introduction and Guide

the timing of reproductive events in relation to climatic seasons, availability of food and other relevant events are discussed. This section may also include birth-weights and sizes of young, growthrates, age at weaning and sexual maturity, and longevity. Finally, if data are available, population structure is discussed. This includes sex ratios, adult/young ratios, the abundance of different cohorts in the population at different times of the year, and mortality rates. In general, information on reproduction is much better known for bats than for other species described in this volume.

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.

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’).

Measurements  A series of morphological measurements is provided. For each species in a particular order there is a standard set of measurements. The abbreviation (and definition) for each measurement are given in the Glossary. A measurement is cited as the mean value (with minimum value to maximum value in parentheses) and sample size. For some, the standard deviation (mean ± 1 S.D.) is given instead of, or in addition to, 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, the localities of the measured specimens and the source of the data are provided. Sources are either cited publications, or specimens in museums, or unpublished information from the authors or others. The acronyms for museums where specimens were examined 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 editors.

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 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

Key References  A select list of references provides more information on the species. Each reference is given in full in the Bibliography.

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

Description

Extinct (EX)

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

Extinct in the Wild (EW)

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

Not Evaluated (NE)

24

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Distribution maps

Table 5.  Acronyms for museum and private collections. Acronym

Museum name

Acronym

Museum name

AM

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)]. California Academy of Sciences, San Francisco, USA. Carnegie Museum of Natural History, Pittsburgh, USA. Centro de Zoologia, Lisboa, Portugal. Durban Natural Science Museum, Durban, South Africa. Estacición Biológica de Doñana, Seville, Spain. Fahr Collection, Ulm, Germany (private collection). Field Museum of Natural History, Chicago, USA. Happold Collection, Canberra, Australia (private collection). Hungarian Natural History Museum, Budapest, Hungary. Harrison Zoological Museum, Sevenoaks, Kent, UK. Instituto da Conservação da Natureza, Lisboa, Portugal. Instituto de Investigação Científica Tropical, Centro de Zoologia, Lisboa, Portugal. Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium. Kansas Museum of Natural History, Lawrence, USA. Los Angeles County Museum, Los Angeles, USA. Museum of Comparative Zoology, Harvard University, Cambridge, USA. Musée d’Histoire Naturelle, La Chaux-de-Fonds, Switzerland. Muséum d’Histoire Naturelle, Genève, Switzerland. Musée d’Histoire Naturelle, Strasbourg, France. Museums of Malawi, Blantyre, Malawi. Muséum National d’Histoire Naturelle, Paris, France. Museo Civico di Storia Naturale di Milano, Milan, Italy. Makerere University, Museum of Zoology, Kampala, Uganda. Museo Zoologico ‘La Specola’, Università di Firenze, Italy.

MZUT NAU

Museo di Zoologia, Università di Torino, Italy. Northern Arizona University Museum of Vertebrates, Flagstaff, Arizona, USA. Naturhistorisches Museum, Berlin, Germany. National Museum (Bloemfontein), South Africa. Naturhistorisches Museum, Bern, Switzerland. National Museums of Kenya, Nairobi, Kenya. Natal Mueum, Pietermartizburg, South Africa. Naturhistorisches Museum, Wien (Vienna), Austria. Natural History Museum of Zimbabwe, Bulawayo, Zimbabwe. Naturhistoriska Riksmuseet, Stockholm, Sweden. Oklahoma State University, Stillwater, USA. Royal Museum for Central Africa, Tervuren, Belgium. Nationaal Natuurhistorisch Museum, Leiden, the Netherlands. (formerly Rijksmuseum Natuurlijke Historie) Royal Ontario Museum, Toronto, Canada. South African Museum, Cape Town, South Africa. Station Biologique de Paimpont, Université de Rennes 1, F-35380 Paimpont, France. Senckenberg Museum, Frankfurt, Germany. Staatliches Museum für Naturkunde, Dresden, Germany. Staatliches Museum für Naturkunde, Stuttgart, Germany. Transvaal Museum, Pretoria, South Africa. United States National Museum of Natural History, Smithsonian Institution, Washington, USA. Yale Peabody Museum, New Haven, Conneticut, USA. Museum Alexander Koenig, Bonn, Germany. Zoologisch Museum, Amsterdam, the Netherlands. Museum für Naturkunde, Humboldt University, Berlin, Germany. Zoological Museum, Moscow University, Moscow, Russia. Zoologisk Museum Universitet, Kobenhavn, Denmark. Zoologisches Museum der Universitat, Zurich, Switzerland.

AMNH BMNH CAS CM CZL DM EBD FC FMNH HC HNHM HZM ICN IICT/CZ IRSN KU LACM MCZ MHNC MHNG MHNS MMB MNHN MSNM MUMZ MZUF

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 (sometimes in the form of table-keys) provide details of the main characteristics of these taxa and can be used as an aid to identification. The tables were prepared by the editors.

Higher order profiles The profiles for orders, families and genera are less structured than for the species profiles. Each profile usually begins with a listing of the taxa in the next lower taxon; for example, each family profile lists the genera in that family. An exception to this arrangement is where a taxon has only one lower taxon. Higher taxa profiles provide the characteristics common to all members of that taxon. Some of these characteristics 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 Crocidura olivieri should be consulted in association with the genus

NHMB NMB NMBE NMK NMP NMW NMZB NRM OSU RMCA RMNH ROM SAM SBPU1 SMF SMND SMNS TM USNM YPM ZFMK ZMA ZMB ZMMU ZMUC ZMUZ

Crocidura profile. For the Chiroptera, the higher taxon profiles end with information that enables readers to recognize and distinguish the next lowest taxa (e.g. family profiles end with the diagnostic characters of the genera in that family).

Distribution maps Each species profile, with a 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 25

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An Introduction and Guide

distribution of a species is indicated as:

Editors of Mammals of Africa

• red shading = current range(s). • × = individual localities when only a few localities are known, or isolated localities considered to be separate from the main geographic range(s). Some localities indicated by × may include two or more closely spaced localities. • ? = locality of uncertain validity; relevant information usually in text. • red arrow = recorded from the island indicated by the arrow.

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 Erinaceomorpha

Order ERINACEOMORPHA – Hedgehogs Erinaceomorpha Gregory, 1910. Erinaceidae (3 genera, 6 species)

Spiny Hedgehogs

p. 29

The order Erinaceomorpha contains one family, ten genera and 20 species, distributed throughout Eurasia and Africa (Corbet 1988, Reeve 1994). The single family contains two subfamilies: the Galericinae (the Gymnures or Hairy Hedgehogs) and the Erinaceinae (the Spiny Hedgehogs). The five genera and six species of Hairy Hedgehogs live in the temperate and tropical forests of Asia, and are not considered further in this account. The Spiny Hedgehogs – usually referred to as just ‘Hedgehogs’ – occur widely in temperate Eurasia and Africa; currently, they are classified into five genera and 14 species, of which six species (in two or three genera) live in Africa. The order (as family Erinaceidae) was previously included within the order Insectivora (Hutterer 2005a). Although the name Erinaceomorpha is not new – it was originally proposed by Gregory in 1910 – recent research has indicated that the Insectivora (hedgehogs and moonrats, shrews, moles, golden-moles, otter-shrews and tenrecs) are not a closely related group, and that the order Insectivora should be divided into three orders – Erinaceomorpha (hedgehogs and moonrats), Soricomorpha (shrews and moles) and Afrosoricida (golden-moles, otter-shrews, tenrecs). The following account deals only with the African hedgehogs – the sole representatives of the Erinaceomorpha in Africa. The two best known characteristics of hedgehogs are their short pointed spines, which densely cover all of the back and flanks, and their ability to curl up into a ball when disturbed. Each spine is thought to be the equivalent of several hairs, where the follicles have coalesced, rather than of a single hair. The spines grow in lines, each spine pointing outwards in a slightly different direction; this results in a dense mat of spines that point in all directions. The spines are shed and replaced, just like normal hairs but more slowly. Hedgehogs can curl into a ball so that the head, limbs, tail and ventral surface are protected by the spiny back and flanks. Under the spiny skin are two large muscles, the orbicularis and panniculus carnosus, which encircle the back, chest and flanks forming a hood-like structure over the animal. Other muscles run downwards from the orbicularis and overlie the forehead, shoulders and rump; when a hedgehog is disturbed, these muscles contract very rapidly pulling the orbicularis downwards. At the same time, the orbicularis itself contracts and the whole body becomes enveloped within the stretched panniculus muscle. This movement stretches the very flexible skin and causes the spines to erect, providing additional protection (Reeve 1994). Hedgehogs can remain in this state for hours on end; they also roll up (but not so tightly) when asleep or in torpor and hibernation. Hedgehogs are characteristically parasitized by many fleas, which may be observed frequently among the spines. Hedgehogs are small mammals, weighing on average 130–205 g as adults (African species). They are compact, rotund little animals because the neck, tail and limbs are short. The head is broad, the snout is slightly elongated and mobile, and the well-developed sense of smell is used for locating prey. The eyes are of moderate

size but sight is primarily monochrome and highly developed. The ears vary from being quite small to very large, and hearing is very acute; some desert species have enlarged auditory bullae (as do some desert rodents), which enhances detection of very quiet noises in open spaces. Olfaction and hearing are the dominant senses for hedgehogs. The skull is strongly built with wide zygomatic arches so that the head appears rather broad. The dental formula is I  3/2, C 1/1, P 3/2, M 3/3 = 36. The teeth are similar in structure and function to other small species of insectivores. One of the upper incisors is long and caniform, and points anteriorly. The canine teeth are small, as are the front premolars. The last premolar and the molars are large with well-developed pointed cusps that crush and slice through the exoskeletons of arthropod prey. Hedgehogs live in a very wide range of habitats from cold temperate steppes to hot tropical savannas. In the cooler parts of their range (where sub-zero temperatures, frosts and snow occur in winter), they enter hibernation (see below). In the hotter parts of their range, they may be active all year, but may enter torpor during the dry season when food is scarce. Unlike many mammals, they adapt to human-modified environments and may be common (but rarely seen) in towns, cities and gardens. They do not live in rainforests or in very dry deserts. Although sometimes considered as rather ‘primitive’ mammals with rather few species, their widespread distribution and adaptability show that, as a family, they are very successful. Hedgehogs are terrestrial, although some species are scansorial and can climb over logs and fences. They have short limbs, each with four or five digits ending in claws. They walk or trot on the soles of the feet, and can move surprisingly quickly for their size. They are nocturnal and active during most of the night, usually with peaks of activity before midnight and around 03:00h. During the day, they rest in a variety of habitats – under logs and piles of stones, in caves and rocky crevices, and in dense litter and hedgerows; some species dig burrows or rest in the burrows made by other animals. Surprisingly, most species can swim. Hedgehogs are omnivorous, but their preferred prey is arthropods (mainly insects), earthworms, snails, small reptiles and eggs. Some species eat fruits and fungi in season. They appear to be resistant to toxins produced by some of their prey (such as centipedes and bees). When food is abundant, hedgehogs store fat under the skin and around some organs, and increase in weight. Fat is utilized (and weight declines) when food is scarce during the colder and drier months of the year. Hedgehogs have the ability to change their metabolic rate in relation to environmental conditions. When the climate is cool or cold (and food may be limited), hedgehogs lower their metabolic rate and reduce their body temperature, and enter a state known as torpor or hibernation. The ability to hibernate is well understood and documented in temperate species, but it appears that African species enter torpor during the cold season of North Africa and South Africa, and during the dry season of tropical Africa. Torpor is physiologically different to hibernation and the metabolic rate 27

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Order Erinaceomorpha

does not decline to the same extent, but it does enable individuals to reduce their energy expenditure and heat loss when environmental conditions are unfavourable. African hedgehogs, like temperate ones, are mostly solitary. Very little is known about their behaviour and social organization in Africa. Likewise, little is known about their reproduction in the wild; the few observations that are available suggest that birth of young in most localities is seasonal. There are usually 2–6 young in a litter, and growth is rather slow. Most individuals probably do not breed until they are about a year old.

In Africa, fossil remains of hedgehogs are known from the early Miocene (Yates 1984). All the extant species of African hedgehogs are rather similar, but each of the six species has radiated into a different environment. The species are essentially allopatric with very limited overlap in their geographical ranges. African hedgehogs are found throughout the continent except in the Rainforest BZ and the driest parts of the Sahara and South-West Arid BZs. D. C. D. Happold

Table 6.  Characteristics of species in the family Erinaceidae in Africa. See also Figure 5. Species

Mean HB (mm)

Central parting of spines

Surface of spines

Face-mask

Digit 1 on hindfoot

Ears (length as % of HF)

Atelerix albiventris

167

Narrow

Smooth

Black; well defined, slight posterior extension on to lower cheek

Absent (or very rudimentary)

Small, rounded, shorter than adjacent spines (73%)

Ateleris algirus

ca. 235

Narrow

Smooth

Small

Small, rounded, shorter than adjacent spines (77%)

Atelerix frontalis

ca. 190

Narrow

Smooth

Small

Small, rounded, shorter than adjacent spines (76%)

Atelerix sclateri

225

Narrow

Smooth

Hemiechinus auritus

179

Absent

Papillate with fine longitudinal grooves

None

Large

Paraechinus aethiopicus

196

Wide

Papillate with fine longitudinal grooves

Black; well defined

Large

None; black ‘spotting’ on muzzle in some individuals Black or dark brown; well defined, extends ventrally to merge with black ventral pelage Black; well defined, extends slightly on to cheek

Small

Small, rounded, shorter than adjacent spines (88%) Large, considerably longer than adjacent spines (114%) Large, slightly pointed, protrude above adjacent spines (132%)

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Family ERINACEIDAE

Family ERINACEIDAE Hedgehogs Erinaceidae G. Fischer, 1817. Mem. Soc. Imp. Nat., Moscow, 5: 372. Hedgehogs Long-eared Hedgehog Ethiopian Hedgehog

Atelerix (4 species) Hemiechinus (1 species) Paraechinus (1 species)

p. 30 p. 37 p. 39

The family Erinaceidae occurs widely in temperate Eurasia and Africa; currently, they are classified into five genera and 14 species. Characteristics of the family are given in the order profile above. African hedgehogs are placed in three genera, Atelerix, Hemiechinus and Paraechinus (Figure 4). Atelerix has been considered as a subgenus of Erinaceus (Corbet 1974a, Yates 1984), which contains several species of European and temperate Asian species, but Robbins & Setzer (1985) showed that it warrants generic distinction (see also Hutterer 2005a). The genera are distinguished by many characters (Figure 5,Table 6), the most important being the presence or absence of a central parting between the spines of the scalp (and if present, its comparative width), the presence or absence of papillae on the spines, the size of the ears and their size relative to the length of the adjacent spines, the form of some of the teeth, the width and form of the palatal shelf, the form of the auditory bullae and (in ??)

the structure of the glans penis (Corbet 1988). Compared with the European Hedgehog Erinaceus europaeus, there are few detailed studies on African hedgehogs, especially in the wild. African hedgehogs range in size (mean HB) from 167 mm to ca. 235 mm. Species are considered as ‘small’ (mean HB of 201 mm). Ear length is considered as ‘small’ (30 mm) (see Table 6).

Hemiechinus auritus.

Atelerix albiventris.

Colour of ventral pelage and limbs

Number of roots on premolar

Notes Paraechinus aethiopicus.

Widespread in savanna and semi-arid. Senegal to Ethiopia; E Africa north of Zambezi R. Morocco to Libya north of Sahara

White; limbs pale

2

White; limbs pale

3

Grey to black; limbs grey to black

2 or 3

Southern Africa only

White or buffywhite; limbs dark

2 (barely divergent)

Somalia

White; limbs long, white

1, 2 (barely divergent), 3

Egypt and Libya

White, dark posteriorly; limbs long, dark brown

1

Sahara Desert and surrounding semi-arid regions

Figure 4. The three genera of African hedgehogs.

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Family ERINACEIDAE

The genera are distinguished by the presence/absence of the central parting on the forehead, whether the spines are papillate or smooth, and the width of the palatal shelf.

a

D. C. D. Happold b

c

Figure 5. External characters of African hedgehogs: Atelerix (left), Hemiechinus (centre) and Paraechinus (right). (a) Head showing parting and face-mask. (b) Left hindfoot. (c) Section of spine. After Corbet (1988).

Genus Atelerix Hedgehogs Atelerix Pomel, 1848. Arch. Sci. Phys. Nat. Geneve 9: 251. Type species: Erinaceus albiventris Wagner, 1841.

The genus Atelerix contains four species endemic to Africa, which occur throughout the savanna regions of the continent. All species in the genus have a narrow parting of the spines on the head, smooth spines (without papillae), rather small ears, broad palatal shelf and small auditory bullae (Figure 6). The hallux (Digit 1 of hindfoot) is

small or absent.The genus Atelerix has often not been recognized, and the four species have been included within the genus Erinaceus (which includes three species very widely distributed in the Palaearctic regions). However, multivariate analysis of cranial characteristics (Robbins & Setzer 1985) as well as other non-cranial characters

Atelerix albiventris.

Atelerix sp.

Figure 6. Skull and mandible of Atelerix frontalis (BMNH 34.10.10.213).

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Atelerix albiventris

Atelerix albiventris.

and distribution patterns, clearly warrant separation of the African species in a separate genus (Corbet 1988, Hutterer 2005a). The four species are allopatric; one species – A. albiventris – is very widespread in central, East and West Africa, and the other species have more limited geographic distributions.

The species are distinguished by presence/absence of face-mask, colour of ventral pelage, presence/absence of Digit 1 on hindfoot, body size and geographic distribution (see Table 6). D. C. D. Happold

Atelerix albiventris  White-bellied Hedgehog (Four-toed Hedgehog) Fr. Hérisson à ventre blanc; Ger. Weissbauchigel Atelerix albiventris (Wagner, 1841). In Schreber. Die Saugethiere, Suppl. 2: 22. Probably Senegal or Gambia.

Taxonomy  Originally described in the genus Erinaceus.The species, as now understood, includes many taxa originally considered to be species and subspecies; all are now treated as synonyms. Synonyms: adansoni, atratus, diadematus, faradjius, heterodactylus, hindei, kilimanus, langi, lowei, oweni, pruneri, sotikae, spiculus, spinifex. Subspecies: none recognized here; the large variation within populations suggests that subspecific differentiation is not justified (Corbet 1988, Reeve 1994). Chromosome number: 2n = 48, aFN = 96 (Hübner et al. 1991). Description  Small hedgehog with four digits on hindfoot. Dorsal pelage of dark spines; spines 15–20 mm, basal half off-white, terminal half dark brown or blackish-brown, often with white tip. Considerable variation in banding pattern on spines. Surface of spines smooth without papillae. Ventral pelage of non-spiny hairs; rather sparse; hairs white or buffy-white. Spiny dorsal pelage and hairy ventral pelage clearly delineated on lower flanks. Head with wide white forehead from cheek to cheek; narrow central parting of spines on crown of head; face-mask on muzzle and around eyes black, well-defined, extending posteriorly on lower cheek (see also below). Ears small, rounded, shorter than adjacent spines; ca. 73% of HF. Limbs short, white or pale; forefeet with

five digits; hindfeet with four digits, Digit 1 absent or rudimentary. Tail relatively very short (ca. 7% of HB), barely visible, with small pale hairs. Nipples: not known. Glans penis without spiny or papillate pads. Some ?? tend to be larger and heavier than //. Skull: auditory bullae comparatively small (see Measurements); P3 with two roots. See Table 6. Geographic Variation  Specimens from drier habitats appear paler because they have a greater number of white-tipped spines; some individuals do not have a black face-mask. Similar Species Paraechinus aethiopicus. On average larger (HB: 196.1 [169–217] mm); wide central parting on crown; spines with papillae and grooves; ears much longer (41–45 mm), longer than adjacent spines; hindfeet with five digits; Sahara and northern semi-arid regions. Atelerix sclateri. On average larger (HB: 225.0 [210–263] mm); Digit 1 of hindfoot present, small but not rudimentary; Somalia only. A. frontalis. On average slightly larger (HB (??): 185 [170–190] mm, HB (//): 196 [186–210] mm); face-mask extending ventrally to merge with black ventral pelage; south of Zambezi R. only. 31

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Family ERINACEIDAE

Distribution  Endemic to Africa. The most widespread of the African hedgehogs. Recorded from Sudan Savanna and Guinea Savanna BZs, Northern and Eastern Rainforest–Savanna Mosaics and most of the Somalia–Masai Bushland BZ. Penetrates the northern margins of the Rainforest BZ where habitat destruction has created grasslands and cultivations. Recorded from Senegal to Ethiopia, Djibouti and Somalia, and southwards through East Africa to Zambia (north of the Zambezi R.) and Malawi. Only known overlap in geographic range with other species of hedgehogs is with Paraechinus aethiopicus in C Sudan and N Ethiopia (and possibly at other places where the northern savannas meet the Sahara Desert), and with A. sclateri in N Somalia. (Note: the record of this species from Liberia [Lienhardt 1982] is presumably an error.) Habitat  Savanna and semi-arid habitats, including rocky insel­ bergs. Tends to avoid waterlogged habitats, marshes and swamps. Often found in suburban gardens and cultivated fields. Abundance  May be common in suitable habitats. Individuals are seen more often at the beginning of the wet season when more individuals are killed by vehicles on roads. Adaptations  Nocturnal and terrestrial. During day, rests under rocks and logs, and in crevices and termitaria. In captivity, two peaks of nocturnal activity are evident: 21:00–24:00h and around 03:00h (Herter 1965).When active, hedgehogs walk and trot on all four limbs. Body temperature (Tb) is maintained at 32.9–35.4 °C at normal ambient temperatures (Herter 1971). In captivity, when Ta is 19– 24 °C, individuals become torpid and less active than normal. A similar drop in Tb is likely to occur in those regions where cool nights are experienced at some times of the year. Foraging and Food  Detailed information unavailable; probably similar to other species of hedgehogs (see order and family profiles).

Atelerix albiventris

In East Africa, the diet is reported to consist of earthworms, snails, slugs, crabs, fruit, fungi, roots and groundnuts (Kingdon 1974). Hedgehogs usually forage alone. Social and Reproductive Behaviour  Primarily solitary. Several vocal sounds are emitted, which enable communication between individuals and express mood. Five categories of audible vocal sounds have been recorded (Gregory 1975). (1) Twitter – a very quiet sound emitted through the closed mouth, and often accompanied by sniffing; usually associated with unfamiliar situations. Each twitter is of very short duration (5–40 msec) and several are emitted in pulses lasting for several seconds. (2) Hiss – a short noise of lower pitch than the twitter, emitted during stressful situations. (3) Snort – similar to the hiss, but louder and emitted when severely stressed or attacked, often repeated rapidly. (4) Scream – a rare sound, emitted under extreme stress. (5) Serenade – a series of low-pitched sounds ranging from a pure whistle to a course squawk, emitted by ?? during courtship behaviour. The faecal pellets have a strong odour, and may be a means of advertising the presence of an individual. Likewise, ‘self-anointing’ (licking the spines of the flank with copious amounts of saliva) may be a means of advertising and ensuring recognition (Reeve 1994). Courtship behaviour is said to be similar to that of the European Hedgehog Erinaceus europaeus (M. W. Gregory, in Reeve 1994). When the ? approaches, the / reacts aggressively, bristling her spines and snorting. The ? attempts to circle the /, who may try to run away; the / responds by vigorously pushing the ? on his flank with the spines on her head. Such behaviour may last for minutes or hours. Copulation occurs when the ? mounts the / from behind; the ? has a particularly long penis, perhaps because the spines on the rump of the / prevent him from getting too close to the /. Reproduction and Population Structure Times of reproductive activity vary in different parts of the range: probably active all year in East Africa (Kingdon 1974), but seasonal in drier and cooler habitats. Collections in Nairobi for a complete year include the following: pregnancies in Jul and Aug; litters in Apr–May; juveniles (50%) due to loss of habitat.

296

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Scotonycteris zenkeri

Measurements Scotonycteris ophiodon FA (!!): 76.5 (74–79) mm, n = 19 FA (""): 77.5 (73–81) mm, n = 38 WS (c): 523 (469–548) mm, n = 10 HB: 117.1 (115–122) mm, n = 9 T: 0 mm E: 22.6 (20–25) mm, n = 14 Tib: 29.1 (26–32) mm, n = 10 HF: 18.1 (17–19) mm, n = 15 WT (!!): 67.2 (60–77) g, n = 12 WT (""): 74.4 (64–95) g, n = 30

GLS: 37.8 (35.3–40.0) mm, n = 8 GWS: 23.3 (22.3–24.3) mm, n = 14 C–M1: 12.5 (12.0–13.4) mm, n = 17 Liberia, Côte d’Ivoire, Ghana, Cameroon (BMNH [incl. holotype cansdalei], FC, SMF, SMNS, USNM, ZFMK) For specimens from Congo, see Geographic Variation Key References Bergmans 1973, 1990; Eisentraut 1959; Hayman 1945; Novick 1958a; Wolton et al. 1982. Jakob Fahr

Scotonycteris zenkeri ZENKER’S FRUIT BAT Fr. Scotonyctère de Zenker; Ger. Zenkers Harlekin-Flughund Scotonycteris zenkeri Matschie, 1894. Sitzb. Ges. Naturf. Fr. Berlin 1894: p. 202.Yaunde [= Yaoundé], Cameroon.

Taxonomy Synonyms: bedfordi, occidentalis. Subspecies: three, but limits uncertain (see Geographic Variation). Chromosome number (Gabon): 2n = 32 (Primus et al. 2006); as a result of conventional staining, no detailed comparison could be made with the banded karyotype of S. ophiodon, which is very distinct from other African fruit bats. Description Very small fruit bat with face-markings (white patch on forehead, white posterior eye-spots, partly white or pale lips); no conspicuous basal ear-patches; no epaulettes; snout and finger-joints not yellowish; forehead region of skull almost straight; bony palate extending well beyond last teeth; cusps of premolars and molars relatively weak; FA: 47–55 mm. Sexes similar in colour; "" on average with slightly longer forearms. Pelage dense, soft and woolly dorsally; shorter and sparser ventrally with stiff hairs on chest and belly; mid-dorsal hairs 9–10 mm. Dorsal pelage medium sepia brown to rusty-brown, mottled; hairs tricoloured, white or whitish with dark brown at base and sepia brown to rusty-brown at tip. Ventral pelage with lower breast and central belly whitish to pale grey, flanks medium to dark brown or greyish-brown, contrasting with the paler areas. No epaulettes or white markings on shoulders. Head with three conspicuous white patches, one on forehead (between anterior corners of eyes) and one at posterior corner of each eye; anterior basal ear-patches indistinct or absent; no posterior basal ear-patches; lips usually bordered by a narrow and indistinct band of white or pale hairs around each corner. Muzzle short, slender; lips only moderately expansible. Ears dark brown (paler near base), naked; oval with broadly rounded tip. Eyes large, dark brown. Palate with four thick and smooth ridges (the first three undivided, the fourth sometimes medially divided) and 6–9 very thin, serrated and irregular ridges (Figure 54a). Wing-membranes dark greenish-brown or brown, reticulated; attaching to first toe; finger-joints not yellowish (cf. S. ophiodon). No visible tail. Skull short and delicate for an African fruit bat. Braincase rounded; rostrum relatively short (30.4–34.6% of GLS), not upturned. Profile of forehead region (viewed laterally) almost straight (cf. Casinycteris). Zygomatic width relatively narrow (62–69% of GLS) and zygomatic arches comparatively lightly built and with lower

margin level with infraorbital foramen (cf. Casinycteris). Bony palate extending well beyond last teeth. Upper incisors comparatively short. Upper canines only moderately long, with faint inner groove, not conspicuously curved, and without secondary cusps or serrated inner edges (cf. S. ophiodon). Premolars and molars rounded to slightly oval in transverse section, with moderate outer cusps but only faint inner cusps (cf. S. ophiodon, Casinycteris argynnis). Dental formula usually 2121/2132 = 28 (variations known). Geographic Variation Three subspecies are tentatively recognized here with the following distributions: S. z. zenkeri: E Nigeria, Cameroon, Gabon and mainland Equatorial Guinea. S. z. bedfordi: Bioko I. S. z. occidentalis: Liberia, Guinea, Côte d’Ivoire, Ghana. Andersen (1912a), Eisentraut (1959) and Kuhn (1961) questioned subspecific differences. However, Bergmans (1990) demonstrated morphological differentiation between five disjunct populations in (1) Liberia, Côte d’Ivoire, Ghana, (2) W Nigeria, (3) Cameroon, mainland Equatorial Guinea and probably E Nigeria, (4) Bioko I. and (5) E DR Congo, which differ in size and/or cranial and dental morphology. This is not adequately reflected in the above subspecific classification. Furthermore, there are isolated records from Congo and Central African Republic and it is not yet clear if these should be assigned to one or more of the above populations or if they represent additional independent and disjunct enclaves. Apart from cranial and dental differences, the population from W Nigeria seems to average largest in measurements while that from Bioko I. is smallest (Bergmans 1990). Similar Species Only two other species of African fruit bats have white markings on the nose, upper lips and/or near the eyes: Casinycteris argynnis. Forearm often longer (50–62 mm). Fingerjoints pale yellowish. Forehead region of skull strongly concave; rostrum upturned; bony palate barely extending beyond last teeth. 297

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Family PTEROPODIDAE

Scotonycteris ophiodon. Much larger (FA: 73–87 mm). Finger-joints yellowish. Distribution Endemic to Africa. Mainly found in the Rainforest BZ (Western, West Central and East Central Regions) and marginally in the Afromontane–Afroalpine BZ. Recorded, disjunctly, from Guinea, Liberia, Côte d’Ivoire, Ghana, Nigeria, Cameroon, Equatorial Guinea (including Bioko I.), Gabon, Central African Republic, Congo and DR Congo. Not yet recorded from Dahomey Gap. A gap between records from W Nigeria and E Nigeria mentioned by Bergmans (1990) has been partly bridged by a record from Orashi, Niger Delta (Angelici et al. 2000). The records from Central African Republic, E DR Congo and Congo appear isolated. This species has been found at almost all localities where S. ophiodon occurs but is much more widely distributed than the latter. Habitat Recorded in lowland rainforest, coastal forest, montane forest, swamp forest, mangroves and marginally in forest–savanna mosaic vegetation zones. Recorded up to 1000 m on Mt Nimba (Verschuren 1976) and up to 1100 m on Mt Kupé and Rumpi Hills, Cameroon (Eisentraut 1973a). However, seems to prefer lowland rainforests and usually is not found higher than 500–800 m (Eisentraut 1973a, Coe 1975, Wolton et al. 1982, Fedden & Macleod 1986). On Bioko I., only found below 400 m (Juste & Perez del Val 1995). Occurs in both pristine and disturbed rainforest and has been mist-netted frequently near forest fringes, in gardens and forest gaps but never in extensively cleared areas or far from closed forest (Eisentraut 1959, Brosset 1966, Jeffrey 1975, Happold & Happold 1978, Wolton et al. 1982, J. Fahr unpubl.). On Bioko I., mist-netted significantly more often in cultivated clearings than in lowland rainforest (Juste & Perez del Val 1995). The habitats in Congo are described as degraded rainforest in an area with low mountains (Bergmans 1973) and heavily disturbed low-altitude forest dominated in the understorey by Haumania liebrechtsiana (Marantaceae) (Dowsett et al. 1991). Abundance Locally rare to very rare. In Taï N. P., comprised 3.6% of fruit bat catch (n = 1216 individuals); sixth most abundant species in community of eight species of fruit bats (J. Fahr & S. Pettersson unpubl.). At Mt Nimba, comprised 2.7% of the fruit bat catch (n = 979) and was the sixth most abundant species in a community of nine species of fruit bats (Wolton et al. 1982). On Bioko I., between 0 and 400 m, represented 13.0% of the catch of four sub-canopy species of fruit bats (n = 332) (Juste & Perez del Val 1995). Adaptations By day, usually roosts hanging from vegetation; has been found between plantain leaves and in trees or bushes, sometimes at the forest edge (Hayman 1946a, Kuhn 1961, Rosevear 1965). Recapture rate in Taï N. P. was 23.8% (n = 21); much higher than in most other fruit bats at this locality. Most were recaptured from six months to one year later; two were recaptured 1.7 and 2.1 years later. All individuals (including subadults and young-adults) were recaptured less than 400 m from the initial site; most less than 250 m away. This suggests that home-ranges are very small (several hectares) and site-fidelity is unusually high for a fruit bat (J. Fahr unpubl.).

Scotonycteris zenkeri

Frequencies of captures had two distinct peaks (21:00–23:00h and 02:00–04:30h), with very few captures between midnight and 02:00h. In contrast, almost the inverse pattern was shown by S. ophiodon at the same locality. Captive S. zenkeri were active for only ca. six hours per night (Wolton et al. 1982). The activity was either broken up into two 3-hour periods with a 2–3 hour rest commencing around midnight, or continuous for ca. six hours. Kuhn (1968) studied the innervation of the larynx of S. zenkeri and concluded that the pattern in this species is basal but derived in Epomophorus labiatus, Epomops buettikoferi and Hypsignathus monstrosus. Foraging and Food Frugivorous. Data regarding vertical preference are equivocal. Eisentraut (1959) and Fedden & Macleod (1986) caught S. zenkeri mainly in elevated mist-nets. Cosson (1995) and Happold & Happold (1978), using only mist-nets at ground-level, caught S. zenkeri in the understorey. The latter authors suggested that foraging is more likely to occur in the upper strata and canopy. However, captures in mist-nets set from 0 to 25 m above ground in Taï N. P. suggest that foraging occurs from ground to canopy level (median capture height 15.4 [1.5–23.6] m, n = 21) (J. Fahr unpubl.). Diet almost unknown. Eisentraut (1959) noted a honey-like smell when examining the stomach of a specimen but could not identify the ingested food. At Mt Nimba, this bat fed extensively on Solanum torvum and S. erianthum between Jul and Sep (Wolton et al. 1982). Captive individuals (in Gabon) refused various fruits, honey-water and flowers (Brosset 1966). Captive individuals (in Liberia) spent 2–2.5 h/night feeding; the mean ± S.D. nightly fruit consumption was 20.3 ± 9.7 g (n = 11); the dry weight assimilated was 90%; the mean consumption per unit body weight was 0.99 g/gbwt and the through-put time was 24 ± 8 min (Wolton et al. 1982). The ratio between body and intestine length is 1 : 3.6 and the intestine is rather short compared to that of other (mainly) frugivorous pteropodids (Eisentraut 1959).

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Social and Reproductive Behaviour Usually roosts singly. In Gabon, Brosset (1966) caught three "", each accompanied by volant young, which were almost the same size as their mothers; later, in captivity, these young huddled with their mothers. According to Wolton et al. (1982), captive "" appear to be more active and more vocal than !!. They emitted an abrupt high-pitched whistle throughout the night, with maximum frequency just before dawn. Adult !! in Taï N. P. had a remarkably strong and spicy scent. Reproduction and Population Structure Litter-size: one (n = 7), two (n = 1) (Jeffrey 1975, Wolton et al. 1982, J. Fahr unpubl.). Embryos are implanted in both uterine horns. In West Africa (E Liberia, Côte d’Ivoire, SW Ghana) the reproductive chronology is probably seasonal bimodal monoestry: out of 70 adult "", pregnancies have been recorded for Jan and Feb (n = 7) and from Jul to Nov (n = 12), with peaks in Feb (37.5% of 16 "") and between Sep (60% of 5 "") and Oct (83.3% of 5 ""). In the same region, lactating "" were found from Feb to Apr (n = 5) and from Oct to Dec (n = 5); notably, none of 10 "" was lactating in Jan (Kuhn 1961, Wolton et al. 1982, J. Fahr unpubl., IRSN, SMF, SMNS, USNM). None of the 29 reproductively active "" was simultaneously palpably pregnant and lactating, which suggests that the chronology is not polyoestry. In the rainforest region of West Africa, the first lactation period (Feb–Apr) would correspond to the onset of the wet season while the second period (Oct–Dec) would be at the end of the wet and start of the dry season, possibly coinciding with peaks in fruit production. Data from central Africa are inconclusive: in SW Cameroon, a " was pregnant with a peasized embryo in Feb, and another " was pregnant in Oct on Bioko I. (Eisentraut 1973a). In the Ituri region of DR Congo (near Epulu: 01° 23' N) a " had an embryo (CR: 15 mm) in Jun (FMNH). The ratio of !! to "" in 75 bats captured at Taï N. P. was 1 : 0.8 (J. Fahr unpubl.); the ratio in 16 bats captured on Bioko I. was 1 : 0.8 (Eisentraut 1973a), and the ratio in 22 museum specimens was 1 : 1.

Predators, Parasites and Diseases Two specimens were recovered from green mambas (Dendroaspis jamesoni) in Niger Delta, Nigeria (Luiselli et al. 2000). No other information. Conservation IUCN Category: Least Concern. Although this species probably depends on undisturbed forest to a lesser degree than S. ophiodon, it is most likely to be lost from areas with extensive clearings and land conversion. Population size unknown but decline inferred from loss of habitat, degradation and fragmentation. Recent records are mostly from undisturbed sites. Measurements Scotonycteris zenkeri FA (!!): 50.3 (47–55) mm, n = 25 FA (""): 52.2 (47–55)mm, n = 21 WS (c): 346 (330–372) mm, n = 10 HB: 77.5 (65–85) mm, n = 37 T: 0 mm E: 14.4 (12–17) mm, n = 38 Tib: 20.5 (18–24) mm, n = 11 HF: 12.7 (11–14) mm, n = 33 WT: 20.3 (16–24) g, n = 59 GLS: 25.9 (24.0–27.4) mm, n = 16 GWS: 16.9 (15.9–17.8) mm, n = 17 C–M1: 8.4 (7.9–9.0) mm, n = 18 Liberia, Côte d’Ivoire, Ghana, Cameroon, Equatorial Guinea, Central African Republic, Congo, DR Congo (BMNH [including holotypes bedfordi and occidentalis], FC, FMNH, IRSN, MNHN, RMCA, ROM, SMF, USNM, ZMB [including holotype zenkeri]) Key References Bergmans 1990; Eisentraut 1959; Hayman 1946a; Wolton et al. 1982. Jakob Fahr

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Family RHINOLOPHIDAE

Family RHINOLOPHIDAE HORSESHOE BATS

Rhinolophidae Gray, 1825. Zool. Journ., 2 (6): 242. Rhinolophus (27 species)

Horseshoe Bats

p. 303

This is a monotypic family, which is widely distributed in the tropical and sub-tropical regions of the Old World, with some species extending into temperate regions.All rhinolophids belong to the genus Rhinolophus. Currently, 77 extant species are recognized, of which 27 occur in Africa (Simmons 2005). Some of these contain subspecies that almost certainly should have specific rank. Rhinolophids have a large, complex noseleaf, the anterior component of which roughly resembles the underside of a horse’s hoof and is referred to as the horseshoe (Figure 49b).The posterior part of the noseleaf has a single, erect, pointed tip. Rhinolophids have large leaf-shaped ears without a tragus, toes with three phalanges, and a medium-length tail that is completely enclosed by an interfemoral membrane (Figures 33b and 55). Rhinolophids resemble hipposiderids (which are closely related) but differ externally in the form of the noseleaf and the number of phalanges in the toes. None are considered to be pests. Rhinolophids are small to medium-sized microbats with long, soft, dense fluffy pelage. Most African rhinolophids are greyish-fawn or greyish-brown in their grey-phase, and have an orange-phase.They have a small compact body and a rounded head with a short blunt muzzle. The ears are almost as broad as they are long and are well separated; the antitragus is conspicuous and there is no tragus. The eyes are very small. The noseleaf is very prominent and is comprised of an anterior horseshoe-shaped component (the horseshoe), a central component, which has a transverse projection (the sella)

and a longitudinal connecting process, and a subtriangular posterior component with an erect tip referred to as the lancet (Figure 56). Many species of rhinolophids are distinguished by the shape of the lancet, sella and connecting process. The wings are large and broad with rounded tips; the second finger has a long metacarpal but no bony phalanges. The hindlimbs are moderately long with small soles and relatively long toes, each (except the hallux) having three phalanges (cf. two in hipposiderids). The tail is relatively short to medium (30–37% of TL), and is completely enclosed by the interfemoral membrane; calcars are present. Females have one pair of pectoral nipples and one pair of pubic nipples; the !! of some species have a false nipple and tuft of stiff hairs (axillary tuft) in each armpit. The skull (Figure 57) has nasal swellings (sometimes called rostral swellings or rostral inflations), a low sagittal crest (usually more prominent anteriorly than posteriorly) and no postorbital processes. Supraorbital ridges vary from weak to prominent.The nasal branches of the premaxillae are lost. The palatal branches are reduced, partly cartilaginous, not fused with each other and not fused with the maxillary; they are usually lost when skulls are cleaned. Because of this, greatest length of skull (GLS) of rhinolophid bats is replaced by CrnC (the distance from the anterior of the upper canine to the most posterior part of the skull). The palate has a very deep, wide, roughly U-shaped anterior palatal emargination, and also a U-shaped posterior emargination, and is shorter than it is broad. The length of the palate between the two emarginations is referred to as the length of the palatal bridge. The tympanic bullae are relatively small

b d

a e c

f

Figure 55. Characters of African bats in the family Rhinolophidae. (a) Flight membranes and bones of wing, hindlimb and tail (e.g. Rhinolophus blasii). (b) Ventral view of armpit showing axillary tuft (e.g. R. alcyone; based on Rosevear 1965). (c) Ear with 11 internal folds, a large antitragus but no tragus (e.g. R. ziama; based on photo by T. Vierhaus). Grooves in the lower lip: (d) one well-defined median groove and no lateral grooves (e.g. R. eloquens, (e) one well-defined median groove and one poorly-defined lateral groove on each side (e.g. R. alcyone) and (f) three well-defined grooves (e.g. R. simulator).

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posterior component

a

central component

Figure 56. (a) Lateral and (b) frontal views of a noseleaf of Rhinolophus showing the components as defined and referred to in the profiles below (some components are defined differently by some other authors).

tip of lancet

b

lancet

connecting process top of sella sella

anterior component

narial cup narial lobe nostril lateral leaflet horseshoe median emargination

but the cochleae are well developed. The dental formula is variable, usually 1123/2133 = 32. However, the anterior upper premolar and/or the posterior middle premolar may be missing. In some species, the anterior upper premolar lies within the toothrow (albeit sometimes slightly displaced) and separates the canine from the posterior premolar: in other species, this tooth is fully displaced labially and the canine and posterior premolar are in contact. Similarly, the middle lower premolar may be within the toothrow or partly or fully displaced labially, or it may be absent, and the anterior and posterior nasal swellings

Figure 57. Rhinolophus hildebrandtii. Skull (RMCA RG 23816).

lower premolars may be separated or in contact accordingly. The upper incisors are very small; the lower incisors are tricuspid. The molariform teeth do not show any particular modification; M3 almost always has three ridges (Csorba et al. 2003). Most African rhinolophids have low to very low aspect ratios and low to very low wing-loadings; only one species is known to have medium wing-loading. They fly slowly with considerable manoeuvrability. They can hover briefly and take off from the ground. Rhinolophids are insectivorous: several feed mainly on moths and/ or small beetles, at least during some seasons. Some forage by slowhawking, and some are predominantly fly-catchers and/or gleaners that often forage within 6 m of the ground, often close to foliage and tree-trunks, in cluttered environments. At least one species catches flying insects in the tip of one wing, bending the phalanges to hold the prey before rapidly transferring it to the mouth (Webster & Griffin 1962). Rhinolophids do not ‘pouch’ their prey in the interfemoral membrane (cf. vespertilionids). They sometimes fly into lighted rooms in search of prey. Most forage alone. Their echolocation calls are typically FM/CF/FM calls with a sustained CF component and maximum energy in the second harmonic: some (perhaps all) species in Africa can be distinguished by the CF-frequency of their calls (Figure 58). The echolocation calls of rhinolophids are adapted to facilitate the detection of flutter (by exploiting Doppler-shifted echoes reflected from the fluttering wings of insects), and are particularly suitable for densely cluttered environments (Neuweiler 1989). Harmonics are of particular value to bats that emit long CF calls.The calls, which initially include the fundamental harmonic, are produced in the larynx, and the nasal swellings (swellings in the skull beneath the noseleaf) facilitate the production of harmonics and suppress the fundamental harmonic in the emitted sound (Suthers et al. 1988).Tracheal chambers suppress the fundamental harmonic in internally reflected sound, and this may enable rhinolophids to rely on the tissue-conducted fundamental as a reference or marker of their own laryngeal generated sound, which could be useful in processing sonar information (Suthers et al. 1988). Rhinolophids emit their echolocation calls through the nostrils and the noseleaf acts as a parabolic reflector to direct a beam of sound in front of the bat and shield the ears, to some extent, from the emitted sounds 301

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a 100 90

b c

d

Frequency (kHz)

80 70

e

60

f

50

g

40 30 20 10 Time (ms)

Figure 58. Sonograms of the FM/CF/FM echolocation calls of seven species of Rhinolophus from Malawi, presented in order of decreasing CF frequency: all bats hand-held (M. Happold unpubl.). (a) R. swinnyi, (b) R. blasii, (c) R. simulator, (d) R. clivosus, (e) R. fumigatus, (f) R. ? eloquens and (g) R. hildebrandtii. Time axis marked at intervals of 50 ms.

(Möhres 1953).While echolocating, the ears move independently with a very characteristic flickering movement, and the head moves up and down and from side to side to beam the sound in different directions. Rhinolophids are unable to scuttle over the ground, and they cannot climb. During the day, most roost in dark caves or cavelike day-roosts such as mine-adits, hollow trees and dark places in buildings. They hang freely from ceilings or hang in contact with vertical walls. When hanging, the tail and interfemoral membrane fold up over the back and the wings fold around the chest so the body is either mostly or fully enclosed by the flight-membranes. Rhinolophids roost singly or in small to very large groups. Very little is known about the social behaviour of African species. Maternity colonies are established by many species. When ! and " R. ferrumequinum begin to roost together at the end of summer, the !! become territorial and establish small harems (see profile); the mating systems of other species in Africa are apparently not known. All of the seven African rhinolophids for which data are available are monotocous and seasonally monoestrous. Delayed fertilization and sperm storage has been documented for R. capensis, R. clivosus, R. ferrumequinum and R. hipposideros, delayed implantation for R. landeri and retarded embryonic development for R. ferrumequinum (in Europe) (see species profiles). In temperate regions, rhinolophids mate in autumn, sperm storage and hibernation occur during winter, and ovulation followed by uninterrupted gestation occurs in spring. In the tropics, young are born at the end of the dry season or in the wet season (data available for five species). It is thought that the young cling to the pubic nipples with their toes and to the pectoral nipples with their teeth. Females carry their young while flying from perch to perch within a day-roost, and will carry the young away from a disturbed day-roost, but they probably do not carry their young while routinely foraging for food. The geological range of the Rhinolophidae is late Eocene to Recent in Europe, Miocene to Recent in Africa, early Miocene to Recent in Asia, middle Miocene to Recent in Australia and Recent in other regions now occupied (Bogdanowicz & Owen 1992, Corbet & Hill 1992, Csorba et al. 2003). The Rhinolophidae had probably diverged

from the Hipposideridae by the late Eocene (Csorba et al. 2003). According to Bogdanowicz & Owen (1992), the family probably originated in the Old World tropics, perhaps in Africa or perhaps in southern Asia. Maree & Grant (1997) also agree that morphological analyses indicate that the family probably originated in South-East Asia. In contrast, a European origin for the family, and monophyly for the African and Palaearctic species is proposed by Csorba et al. (2003). Based on genetic evidence, Csorba et al. (2003) suggest that there was an early emergence of a clade containing R. trifoliatus and R. hipposideros (of which only R. hipposideros occurs in Africa), and a clade containing, amongst others, an African clade, which probably contains all of the other African species. Within this African clade, the clade representing the most basal lineage comprises species linked to rainforest habitat (R. alcyone, R. landeri and probably R. guineensis). Subsequently, there seems to have been a series of radiations into dry environments. The first of these radiations seems to have given rise to a Mediterranean group (R. blasii, R. euryale and R. mehelyi) and R. blasii appears to have migrated southwards as far as southern Africa more recently. Subsequently, there appear to have been at least two radiations in arid areas of eastern and southern Africa, one giving rise to a lineage including R. capensis, R. denti, R. simulator and R. swinnyi (Maree & Grant 1997) and probably also R. adami and R. maendeleo (Csorba et al. 2003), and another giving rise to a lineage containing R. clivosus, R. darlingi, R. fumigatus and R. hildebrandtii (Maree & Grant 1997) and also R. ferrumequinum, R. maclaudi and R. eloquens (Csorba et al. 2003). Rhinolophids have radiated into forests, woodlands and semi-desert habitats at both high and low altitudes. The majority of species are tropical or sub-tropical and a few occur in temperate regions. Of the 27 species that occur in Africa, 14 are found only in the tropics, four are found only in temperate regions and seven are in tropical, sub-tropical and temperate regions. Seven species are only or mainly found in forest habitats, nine occur in both forests and savannas, four occur only in savannas, one occurs in both savanna and arid habitats, three occur in all of these habitats, and three which are found in the Mediterranean Coastal BZ extend marginally into the arid habitats of the Sahara Arid BZ. Thirteen species (48%) have been found in montane habitats but,

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of these, only R. ruwenzorii and the very closely related R. hilli are found only in montane habitats. It is not uncommon to find two species at any one locality, but the occurrence of three or more species seems to be very rare (except in Malawi where 3–5 species have been recorded at six localities Happold & Happold 1997). Some examples are:

(Simmons 1998, Simmons & Geisler 1998), but more recent molecular studies have contradicted many groupings based on morphological data and, pending resolution of the controversies, no chiropteran superfamilies are recognized by Simmons (2005). Some authors, including Koopman (1984, 1993, 1994) and Simmons (1998), follow Tate (1941) in considering the Rhinolophidae to Misserghin, Algeria (garrigue): R. blasii, R. ferrumequinum, R. include the Hipposideridae as a subfamily. However, the familial hipposideros and R. mehelyi (Kowalski & Rzebik-Kowalska 1991). status of the Hipposideridae has been retained by Maree & Grant Shimoni, Kenya (coastal forest mosaic): R. fumigatus, R. deckenii and R. (1997), Csorba et al. (2003), Simmons (2005) and in the majority landeri (Aggundey & Schlitter 1984). of books about African mammals and is retained here. For further Liwonde N. P., Malawi (miombo woodland): R. blasii, R. clivosus, R. comment, see Family Hipposideridae. fumigatus, R. hildebrandtii and R. simulator (Happold & Happold 1997). The Rhinolophidae have been comprehensively reviewed by Csorba et al. (2003). All rhinolophids belong to the genus Rhinolophus. Based on morphological data, the family Rhinolophidae was placed in the superfamily Rhinolophoidea with the families Nycteridae, Meredith Happold & F. P. D. Cotterill Megadermatidae and the very closely related Hipposideridae

GENUS Rhinolophus Horseshoe Bats Rhinolophus Lacépède, 1799. Tabl. Div. Subd. Orders Genres Mammifères, p. 15. Type species: Vespertilio ferrum-equinum Schreber, 1774.

This genus currently has 77 species of which 27 occur in Africa (Simmons 2005). There are no other genera in the family Rhinolophidae and the characters of this genus are given in the family profile. a

b

f

j

This genus is divided into 15 species-groups following Csorba et al. (2003) (see also Bogdanowicz 1992, Bogdanowicz & Owen 1992, Maree & Grant 1997, Fahr et al. 2002). Of these groups, eight are represented in Africa:

k

l

c

d

g

e

h

m

i

n

Figure 59. Variations in the shape of the connecting process of the noseleaf of African Rhinolophus. (a) Noseleaf showing (black) that part of the connecting process that is described by the following terms. Rounded as in (b) R. maendeleo (based on Kock et al. 2000), (c) R. adami (based on Kock et al. 2000), (d) R. simulator (M. Happold, Malawi) and (e) R. clivosus (M. Happold, Malawi). Subtriangular as in (f) R. landeri (based on Csorba et al. 2003), (g) R. guineensis (based on Csorba et al. 2003), (h) R. alcyone (based on Csorba et al. 2003) and (i) R. mehelyi (based on Csorba et al. 2003). Rising to a high horn as in (j) R. blasii (M. Happold, Malawi) and (k) R. euryale (based on Gaisler 2001a). Rounded but rising to a high peak as in (l) R. sakejiensis (based on Csorba et al. 2003). Very low, slightly rounded or flat as in (m) R. hipposideros (based on Csorba et al. 2003). Greatly reduced, very low and concave as in (n) R. ziama (from photo in Fahr et al. 2002).

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adami group: two species – R. adami, and R. maendeleo. capensis group: four species – R. capensis, R. denti, R. simulator and R. swinnyi. euryale group: two species – R. euryale and R. mehelyi. ferrumequinum group: eight species including R. clivosus, R. darlingi, R. deckenii, R. ferrumequinum, R. hillorum, R. sakejiensis and R. silvestris. fumigatus group: three species – R. eloquens, R. fumigatus and R. hildebrandtii. hipposideros group: one species – R. hipposideros. landeri group: four species – R. alcyone, R. blasii, R. guineensis and R. landeri. maclaudi group: four species – R. hilli, R. maclaudi, R. ruwenzorii and R. ziama. Rhinolophus hildebrandtii.

The African species are distinguished mainly on the basis of the following characters (Table 14):

Table 14.  Key to the African species in the family Rhinolophidae. Information from African material, mostly from species profiles. Sample sizes of R. adami, R. hilli, R. maendeleo, R. sakejiensis, R. silvestris and R. ziama are less than six. Distinguishing species that are morphologically very similar from the information below is not easy, and identifications should be confirmed using the additional information in the Description, Similar Species and Distribution sections of the species profiles.

Connecting process

Rounded Rounded

Position of anterior upper premolara (Proximity of canine and posterior premolar) Within toothrow (Well separated) Within toothrow (Well separated)

Rounded

Within toothrow (Well separated)

Rounded

Within toothrow (Well separated)

Rounded

Within toothrow (Well separated)

Rounded

Rounded Rounded Rounded Rounded Rounded

Usually partly displaced, sometimes fully displaced (Typically well separated, never in contact) Usually fully displaced or absent (Separated by very narrow gap) Fully displaced (In contact or almost so) Fully displaced or absent (In contact) Fully displaced or absent (In contact) Fully displaced or absent (In contact)

Sellab Hairiness and shape

Naked Sides slightly concave Naked Sides concave Naked Almost parallel-sided or slightly concave

Lancet

Subtriangular, sides slightly convex Tip broad and rounded Subtriangular, sides slightly concave Tip bluntly pointed

Horseshoe mean (range) (mm)

8.5, 9.0 8.2, 8.4

Subtriangular, sides slightly concave Tip bluntly pointed

7.1 (6.8–7.5)

Naked Sides slightly concave

Subtriangular, almost hastate, sides concave Tip bluntly pointed

6.8 (6.0–7.4)

Naked Sides slightly concave

Subtriangular or hastate Tip somewhat rounded

7.4 (6.7–9.0)

Naked Sides parallel or slightly concave

Hastate Tip bluntly pointed

7.7 (7.3–8.1)

Naked Sides parallel or slightly concave

Subtriangular, sides almost straight Tip bluntly pointed

10.3 (9.1–11.5)

Naked Sides concave Naked Sides concave Naked Sides slightly concave Naked Sides concave

Subtriangular, sides slightly concave Tip bluntly pointed Subtriangular, sides slightly concave Tip bluntly pointed Hastate Tip bluntly pointed Hastate Tip rounded

9.5, 10 8.0 (7.1–8.7) 8.1 (7.0–9.0) 7.8 (6.6–9.6)

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Connecting process. Can be rounded (with variable height relative to the height of the sella), rounded but rising to a high peak, rising to a high horn (not rounded), subtriangular, or greatly reduced (Figure 59). Position of the anterior upper premolar. Can be within the toothrow (so canine and posterior premolar are well separated), or fully displaced labially (so canine and posterior premolar are in contact) or partly displaced labially (so canine and posterior premolar are almost in contact). It is only the cingula of the canine and premolar that are ever in contact. The anterior upper premolar is, or can be, absent in some species (Figure 62). Sella. Can be hairy (well covered by longish, moderately conspicuous hairs) or naked (sparsely covered by inconspicuous short hairs). The shape of the sella is variable (Figure 60). Lancet. Can be subtriangular, subtriangular with slightly convex sides, subtriangular with slightly concave sides, hastate (having a broad base, markedly concave sides and narrow tip), or short, very narrow and almost parallel-sided (Figure 61). The tip of the lancet can be bluntly pointed or broader and more rounded.

First phalanx of fourth finger as % of 4th metacarpal: categoryc, mean (range)

Greatest breadth of horseshoe. Relative length of first phalanx of the fourth finger to the metacarpal of the fourth finger. Said to be relatively short if its mean relative length is less than 22% of the metacarpal, medium if its mean relative length is 22–25% of the metacarpal, and long if its mean relative length is >25%. Length of forearm, tibia and ear. Relative length of palatal bridge. This is the distance from anterior emargination to posterior emargination along mid-line of the bony palate, excluding the median spike projecting from the posterior margin (Figure 23g); expressed as a percentage of C–M3. Said to be short if less than 30%, medium if 31–37%, and long if >37% (all data from Csorba et al. 2003). Axillary tuft. Present (Figure 55b) or absent. Ear folds. This is the number of internal folds in the outer side of the pinna (Figure 55c). Particularly relevant to species in the maclaudi group.

FA mean (range) (mm)

Tib mean (range) (mm)

E mean (range) (mm)

46–50d

20, 20

25, 26

48, 49

19, 19

24, 25

Medium 24 (21–27)

42.0 (37–44)

16.7 (15–18)

18.0 (14–21)

Medium 22 (21–25)

41.7 (40–44)

18.4 (17–21)

17 (15–20)

Long 23 (20–25)

45.2 (42–49)

18.3 (18–20)

21 (18–23)

Long 26 (24–30)

48.8 (47–51)

18.7 (17–21)

23.8 (21–25)

Western and Eastern Cape Provinces, South Africa

R. capensis

Long 27 (24–30)

53.1 (48–56)

25.0 (24–28)

22.6 (18–27)

Uganda, Kenya, Tanzania, Pemba I., Mafia I. and Zanzibar I. Mainly in coastal forests

R. deckenii

53.6 (50–56)

23 (23–24)

22, 23

46.9 (42–51)

20.9 (20–22)

20.1 (15–23)

55.4 (51–59)

24.0 (23–25)

22.8 (19–25)

North-West Africa

R. ferrumequinum

51.5 (42–59)

21.9 (16–27)

20.1 (16–24)

Widespread but not north-west Africa and not West Africa

R. clivosus

Medium 23 Medium 21, 23

Long ca. 31 Long 27 (24–30) Long 28 (24–31) Long 28 (25–31)

Miscellaneous

Palatal bridge 42–44% of C–M3 Congo Palatal bridge 37, 39% of C–M3 Tanzania Sella breadth not known CrnC 16.5 (15.8–17.3) mm See Distribution Sella comparatively narrow (1.2–1.3 mm prior to preservation) CrnC: 17.4 (17.0–18.2) mm See Distribution Sella comparatively broad (1.5–1.7 mm after preservation) CrnC: 18.4 (17.3–19.3) mm See Distribution

Gabon and Congo CrnC: 23.1 (22.3–23.7) mm Angola to Tanzania and southwards to South Africa CrnC: 19.3 (18.4–20.5) mm

Species

R. adami R. maendeleo R. denti

R. swinnyi

R. simulator

R. silvestris R. darlingi

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Table 14. continued.

Connecting process

Position of anterior upper premolara (Proximity of canine and posterior premolar)

Sellab Hairiness and shape

Rounded

Absent (In contact)

Naked Sides almost parallel

Rounded

Fully displaced or absent (In contact or almost so)

Lancet

Horseshoe mean (range) (mm)

Fully displaced or absent (In contact or almost so) Fully displaced or absent (In contact) Absent (n = 3) (In contact)

Hairy Sides concave near base, parallel above Hairy Upper two-thirds parallel-sided Hairy Sides parallel or slightly concave Naked Sides slightly concave

Short, mostly very narrow and almost parallel-sided Tip hairy and rounded Subtriangular, sides straight or slightly concave Tip rounded Subtriangular, sides slightly concave Tip rounded Subtriangular, sides slightly concave Tip rounded Hastate Tip long, narrow, bluntly pointed

Rising to high horn

Within toothrow (Well separated)

Naked Wedge-shaped, top narrow

Subtriangular or hastate Tip rounded

8.1 (7.2–9.0)

Rising to high horn

Within toothrow (Well separated)

Naked Parallel-sided, top broad, rounded

Subtriangular Tip bluntly pointed

7.2 (6.5–7.5)

Subtriangular

Somewhat displaced (Separated)

Naked Parallel-sided

Hastate, upper half narrow and almost parallel-sided Tip bluntly pointed

6.4 (4.9–7.5)

Subtriangular

Within toothrow (Well separated)

Naked Sides slightly concave

Hastate Tip bluntly pointed

7.2 (6.0–8.0)

Subtriangular

Within toothrow (Well separated)

Naked Parallel-sided

Subtriangular

Within toothrow (Well separated)

Naked Parallel-sided

Very low, slightly rounded or flat

Within toothrow (Well separated)

Naked Long, narrow, wedge-shaped

Hastate Tip bluntly pointed Subtriangular, sides straight or slightly concave Tip bluntly pointed Subtriangular, sides slightly concave Tip bluntly pointed

Greatly reduced, very low, concave

Within toothrow or slightly displaced (Separated)

Naked Inclined forward, sides almost parallel, basal lobes very large

Greatly reduced, very low, concave

Within toothrow (Separated)

Greatly reduced, very low, concave

Within toothrow or slightly displaced (Separated)

Greatly reduced, very low, concave

Displaced (Almost in contact)

Rounded Rounded Rounded but rising to high narrow peak

Hairiness not known Inclined forward, sides almost parallel, basal lobes very large Naked Upright, sides concave, basal lobes very large Naked Upright, sides concave, basal lobes very large

8.8 (8.4–9.1)

13.0 (11.0–15.0) 11.6 (11.1–12.3) 10.3 (9.6–11.5) ca. 10–11

8.6 (8.3–9.3) 10.0 (8.3–11.2) 6.6 (6.1–7.1)

Subtriangular, moderately tall Tip pointed

15.0, 16.0 –

Almost parallel-sided, moderately tall Tip moderately broad and rounded

ca. 11.5

Subtriangular, moderately tall Tip truncated

11.9 (10.8–12.6)

Subtriangular, tall Tip bluntly pointed

12

a

The anterior upper premolar is said to be ‘within toothrow’ if it lies in the middle of the toothrow or is only slightly displaced labially, causing the canine and posterior upper premolar to be distinctly separated. b The sella is said to be naked unless it is well covered by longish and fairly conspicuous hairs (as opposed to inconspicuous short hairs). c The first phalanx of the fourth finger varies in its length relative to that of the fourth metacarpal: it is said to be ‘short’ if its mean relative length is less than 22% of the metacarpal, ‘medium’ if its mean relative length is 22–25% of the metacarpal, and ‘long’ if its mean relative length is >25%. d No mean given; sample size not known.

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First phalanx of fourth finger as % of 4th metacarpal: categoryc, mean (range)

FA mean (range) (mm)

Tib mean (range) (mm)

E mean (range) (mm)

Miscellaneous

Species

Long 28 (27–31)

54.5 (52–57)

23.6 (22–25)

22.4 (21–24)

West Africa, perhaps Sudan

R. hillorum

Long 27 (24–29)

63.9 (60–67)

27.9 (26–31)

33.2 (26–36)

CrnC: 27.4 (26.1–28.7) mm

R. hildebrandtii

58.4 (53–63)

23.9 (22–25)

27.5 (21–38)

CrnC: 25.4 (24.8–26.2) mm

R. eloquens

50.9 (47–60)

21.4 (19–24)

22.9 (19–28)

CrnC: 22.7 (21.6–24.3) mm

R. fumigatus

53, 55, 55

25

20, 22, 22

Long 26 (24–28)

46.0 (43–48)

19.3 (18–21)

18.0 (16–21)

Short 20 (17–22)

48.2 (46–50)

21.2 (21–22)

20.1 (19–22)

Short 21 (19–23)

50.1 (48–53)

20.9 (19–23)

20.9 (19–23)

Short 21 (19–23)

43.3 (35–49)

18.7 (17–21)

16.6 (13–20)

Short 20 (17–21)

46.3 (44–50)

21.1 (21–22)

19.1 (17–22)

Short 21 (19–22)

52.3 (48–56)

24.2 (21–27)

22.2 (19–25)

Rainforest. Senegal to Uganda Axillary tufts (!!) orange-red

R. alcyone

Long 26 (24–28)

36.8 (35–40)

17.2 (15–18)

16.1 (15–17)

North-West Africa, Sudan, Ethiopia, Eritrea, Djibouti, Sinai

R. hipposideros

Long 26 (23–28) Long 26 (24–28) Long 27

Long 26 (25–29)

65.8 (64–69)

29.0 (28–31)

41.4 (40–46)

Long 26, 29

60, 60

26, 27

35, 36

Long 27 (26–28)

57.6 (55–62)

23.6 (22–26)

34.7 (32–38)

Long 26, 27

54, 54

24

29

As yet, only NW Zambia No axillary tufts (!!) No marked contrast between crown areas of anterior and posterior lower premolars North-West Africa, and eastern Africa to South Africa Marked contrast between crown areas of anterior and posterior lower premolars North-West Africa Marked contrast between crown areas of anterior and posterior lower premolars Axillary tufts: n. d. Morocco to N Egypt, north of Sahara Widely distributed S of Sahara (but not Cape Provinces, South Africa) Axillary tufts (!!) reddish-brown Senegal, Guinea, Sierra Leone Axillary tufts (!!) usually white

Horseshoe without lateral leaflets, median emargination very small or absent Each ear with 10–12 folds Guinea Horseshoe without lateral leaflets, without median emargination Each ear with 11 or 12 folds Horseshoe with lateral leaflets and median emargination Each ear with eight folds Rwenzori Mts (DR Congo, Uganda), Rwanda Horseshoe with lateral leaflets and median emargination Each ear with nine folds SW Rwanda

R. sakejiensis R. blasii

R. euryale

R. mehelyi

R. landeri R. guineensis

R. maclaudi

R. ziama

R. ruwenzorii

R. hilli

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b

c

d

a

f

e

Figure 60. Variations in the shape of the sella in African Rhinolophus. (a) Frontal view of the noseleaf showing the relevant part of the sella (shaded). The sella can be described as (b) almost parallel-sided (e.g. R. alcyone; based on Rosevear 1965), (c) with upper two-thirds parallel-sided (e.g. R. hildebrandtii; M. Happold, Malawi), (d) concave sided (e.g. R. clivosus; M. Happold, Malawi), (e) wedge-shaped (e.g. R. blasii empusa; M. Happold, Malawi) or (f) parallel-sided with greatly expanded narial lobes (e.g. R. ziama; based on Fahr et al. 2002). Sellas which curve forwards have been pushed back to flatten the anterior surface and reveal its outline.

a

d

c

b

e f

Figure 61. Variations in the shape of the lancet in African Rhinolophus. (a) Frontal view of the noseleaf showing the lancet (shaded). The lancet can be (b) subtriangular (e.g. R. deckenii; based on Csorba et al. 2003), (c) subtriangular with slightly convex sides (e.g. R. adami; based on Kock et al. 2000), (d) subtriangular with slightly concave sides (e.g. R. maendeleo; based on Kock et al. 2000), (e) hastate (e.g. R. ferrumequinum; based on Csorba et al. 2003), or (f) short, very narrow and almost parallel-sided (e.g. R. hillorum).

Distribution. Some species are extremely similar morphometrically, and are best distinguished by their distributions. The validity of the specific status of some of these species is uncertain. Additional diagnostic characters include: Cranio-canine length (CrnC). For Rhinolophus, the distance from the anterior of the upper canine to the most posterior part of the skull (Figure 23f) is used instead of GLS. This is because the nasal branches

of the premaxillae are absent and the palatal branches are frequently lost during preparation of the skulls of these bats. Zygomatic width relative to mastoid width. Zygomatic width ranges from much greater than mastoid width to much narrower than mastoid width (see Figure 63b and c for same character in Hipposideridae). Lateral leaflets (on each side of horseshoe). Can be present, rudimentary (inconspicuous) or absent.

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Rhinolophus adami

Number of grooves in the chin region of the lower lip. There is invariably a well-defined median groove. Lateral grooves (one on each side of the median groove) can be well defined, indistinct or absent (Figure 55d–f).

a

Median emargination in horseshoe. Can be present (deep to shallow) or absent. Frontal and lateral views of the noseleaf, lateral views of the rostral part of the skull, and occlusal views of the upper canine and premolars, are given for almost all African species in Csorba et al. (2003).

b

The species are presented in alphabetical order irrespective of the species-group to which they belong. Meredith Happold c

Figure 62. Variations in the size and position of the upper premolars in African Rhinolophus. Left: occlusal views of teeth on left side of upper jaw. Right: lateral views of labial side of the same teeth. The anterior upper premolar can be (a) within the toothrow so canine and posterior premolar are well separated (e.g. R. denti), (b) partly displaced labially so canine and posterior premolar are almost in contact (e.g. R. hilli), (c) fully displaced labially so canine and posterior premolar are in contact (e.g. this specimen of R. fumigatus), or (d) absent (e.g. this specimen of R. hildebrandtii).

d

Rhinolophus adami ADAM’S HORSESHOE BAT Fr. Rhinolophe du Congo; Ger. Adams Hufeisennase Rhinolophus adami Aellen and Brosset, 1968. Rev. Suisse Zool. 75: 443. Grotte de Kimanika, Kouilou, Congo.

Taxonomy Species-group: adami (with R. maendeleo). Appears closely related to R. maendeleo (Kock et al. 2000). Synonyms: none. Chromosome number: not known. Description Small microbat with noseleaf (posterior component subtriangular with erect tip); medium-sized for an African rhinolophid; anterior upper premolar within toothrow; connecting process rounded and comparatively high; lancet subtriangular with slightly convex sides; sella with slightly concave sides and top which curves acutely forward and slightly downward; horseshoe breadth 8.5–9.0 mm. Sexual dimorphism: no information. Dorsal pelage brownish. Ventral pelage brownish-grey, sometimes becoming whitish on lower abdomen. Orange-phase: no information. Axillary tufts: no information. Ears comparatively and relatively of medium length (25–26 mm, 51–53% of FA), brown with darker rims. Noseleaf with lancet long, subtriangular with rounded tip, sides convex (Figure 61c) or almost so (cf. R. maendeleo). Connecting

process well developed, smoothly rounded, rising higher than sella. Sella naked, large and broad, sides slightly concave, top curving acutely forward and slightly downward. Basal lobes of sella poorly developed but surrounding a well-developed narial cup. Horseshoe narrow (8.5, 9.0 mm); almost covering muzzle; no lateral leaflets; median emargination well defined. Lower lip with three grooves. Wings dark brown; first phalanx of fourth finger of medium relative length (23% of fourth metacarpal, n = 2). Tibia 38–40% of FA (n = 2). Baculum with basal cone short with shallow dorsal and ventral invaginations (cf. R. maendeleo); shaft becoming dorsoventrally flattened distally; tip not expanded (Kock et al. 2000). Skull narrow; zygomatic width much less than mastoid width. Rostrum with anterior median swellings well developed, posterior swellings reduced (Csorba et al. 2003). Frontal depression moderately deep. Sagittal crest low. Infraorbital foramen covered by bony bar (cf. R. maendeleo). Palatal bridge 2.8–3.0 mm, 42–44% of C–M3. Anterior upper premolar small, within toothrow; canine and 309

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posterior premolar well separated. Middle lower premolar slightly displaced labially; anterior and posterior lower premolars separated. Dental formula 1123/2133 = 32. Geographic Variation None known. Similar Species Five other Rhinolophus in Africa have the anterior upper premolar in the toothrow and a rounded connecting process, but none are known to be sympatric with R. adami: Rhinolophus maendeleo. Lancet subtriangular with slightly concave sides; connecting process rising only just above sella; sella with markedly concave sides, top of sella curving obtusely forward; horseshoe narrower (8.2, 8.4 mm). Skull with infraorbital foramen open; palatal bridge 37–39% of C–M3. R. simulator simulator. Ears shorter (18–23 mm). Lancet hastate, horseshoe narrower (6.7–8.3 mm). Palatal bridge relatively short (29–34% of C–M3). R. swinnyi. Smaller (FA: 40–44 mm). Lancet hastate, horseshoe narrower (6.8–7.4 mm). R. denti. Smaller (FA: 37–44 mm). Horseshoe narrower (6.8– 7.5 mm). R. capensis. Horseshoe narrower (7.3–8.1 mm). Distribution Endemic to Africa. Known only from the type locality, Kouilou, Congo, in the Rainforest–Savanna Mosaic very near the Rainforest BZ. Habitat The vegetation of the Kouilou region is a mosaic of lowland rainforest and secondary grassland, and limestone caves are present. Abundance Only four specimens were described by Aellen & Brosset (1968): these, and an additional seven specimens (six in

MNHN, one in HNHM) recognized by Kock et al. (2000), appear to be the only known specimens. Probably very rare (see Conservation). Remarks The holotype, an adult ", and two subadult "" were collected on the same day in a limestone cave (Grotte de Kimanika) for which no details are available (Aellen & Brosset 1968). A fourth specimen, an adult !, was collected in another limestone cave (Grotte de Meya-Nzouari), parts of which were dry and ‘dead’ with respect to the growth of limestone formations, and parts of which were still ‘living’ and very humid, with water seeping over the formations and trickling into underground streams (Adam & Le Pont 1974). This cave was also inhabited by Rousettus aegyptiacus, Rhinolophus silvestris, Hipposideros ruber, H. gigas, Triaenops afer and Miniopterus minor. Predators, Parasites and Diseases Ectoparasites include a bat-fly Penicillidia penicillidia (Diptera: Nycteribiidae) (Anciaux de Faveaux 1984). Conservation IUCN Category: Data Deficient. Known from only four specimens from only two of 45 caves investigated for the presence of bats over a period of ca. seven years (Adam & Le Pont 1974), and an additional seven specimens from the type locality (Kock et al. 2000). Furthermore, not recorded from ten localities in nearby Mayombe and Lower Kouilou regions of Congo, where 80 specimens (14 species) were collected by Dowsett et al. (1991). Also, not recorded from Haut-Ivindo region of Gabon where 1732 individuals belonging to 27 species were captured from caves and by mist-netting by Brosset (1966), nor from Kikwit (05° 13' S, 18° 49' E, DR Congo) where 538 bats belonging to 18 species were collected in 1995 (Van Cakenberghe et al. 1999). As indicated by Dowsett et al. (1991), this species is potentially at risk because it roosts in caves occupied by the fruit bat Rousettus aegyptiacus, which is sometimes hunted for food. Measurements Rhinolophus adami FA: 46–50 mm, n = ?* WS: n. d. TL: n. d. T: 28, 27 mm E: 25, 26 mm NL (breadth): 9.0, 8.5 mm Tib: 20, 20 mm HF: 9, 9 mm WT: n. d. CrnC: 20.1 (19.8–20.6) mm, n = 7* GWS: 9.8 (9.5–9.9) mm, n = 7* C–M3: 7.3 (7.1–7.6) mm, n = 7* Congo (holotype ! and one adult " respectively; Aellen & Brosset 1968) *Csorba et al. 2003 Key Reference Aellen & Brosset 1968. Meredith Happold

Rhinolophus adami

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Rhinolophus alcyone

Rhinolophus alcyone HALCYON HORSESHOE BAT Fr. Rhinolophe alcyon; Ger. Temmincks Hufeisennase Rhinolophus alcyone Temminck, 1852. Esquisses Zool. sur la Côte de Guiné, p. 80. Boutry River, Ghana.

Taxonomy Species-group: landeri. Synonyms: none. Chromosome number: not known.

premolar within toothrow; connecting process triangular and pointed; first phalanx of fourth finger relatively short:

Description Small microbat with noseleaf (posterior component subtriangular with erect tip); medium-sized for an African rhinolophid; anterior upper premolar within toothrow; connecting process triangular; lancet subtriangular; sella parallel-sided; first phalanx of fourth finger relatively short; axillary tufts orange-red (if present); tibia 24 (21–27) mm. Sexes similar. Pelage dense, soft, fluffy; mid-dorsal hairs ca. 9 mm. Dorsal pelage dark to medium brown; hairs yellowishbeige with brown at tip. Ventral pelage slightly paler. Orange-phase: dorsal pelage pale brown to bright orange-red. Adult !! often with orange-red or brown axillary tufts. Ears comparatively and relatively short (19–25 mm, 36–49% of FA); each with 8–9 internal folds. Noseleaf with lancet subtriangular (margins straight or slightly concave), tip bluntly pointed. Sella with extremely short white hairs, narrow, with straight, almost parallel sides (Figure 60b); top rounded, curved forward. Connecting process well developed, subtriangular, with bluntly to sharply pointed tip. Horseshoe of medium breadth (8.3–11.2 mm) almost covering muzzle; lateral leaflets present, median emargination present. Lower lip with a well-defined median groove and two poorly defined lateral grooves. Wings medium to blackish-brown; first phalanx of fourth finger relatively short (20.8 [19–22]% of fourth metacarpal, n = 20). Interfemoral membrane paler. In one specimen (possibly abnormal) the flight-membranes are cream with dark brown reticulation (BMNH 66.6242). Tibia 46.1 (38–52)% of FA (n = 22). Skull robust; zygomatic arches moderately robust; zygomatic width greater than mastoid width. Nasal swellings relatively high, anterior median swellings globular and prominent, lateral and posterior swellings medium (Csorba et al. 2003). Frontal depression very shallow; supraorbital ridges weak. Sagittal crest variable – low to well developed. Palatal bridge 29–35% of C–M3. Anterior upper premolar within toothrow or only slightly displaced labially; canine and posterior premolar well separated. Middle lower premolar either within toothrow or displaced labially; anterior and posterior lower premolars separated. Anterior lower premolar at least two-thirds the height of the posterior premolar. Dental formula 1123/2133 = 32.

Rhinolophus landeri. Body measurements almost always smaller (FA: 35– 49 mm, Tib: 17–21 mm). Skull smaller (CrnC: 16.9–19.1 mm). R. guineensis. Body measurements usually smaller (FA: 44–50 mm, Tib: 21–22 mm). Skull smaller (CrnC: 19.2–20.6 mm). Sagittal crest less developed. Axillary tufts in !! usually white. Distribution Endemic to Africa. Mainly recorded from the Rainforest BZ and surrounding Rainforest–Savanna Mosaics, from Guinea to Ghana, from Nigeria to Central African Republic and southwards to S Congo and Bioko I., with some apparently isolated records in NE DR Congo, S Sudan and Uganda. Also recorded from the Sudan Savanna BZ in Senegal, and the Guinea Savanna BZ in Côte d’Ivoire and Ghana. Records are scattered but this probably reflects insufficient collecting. Predicted to occur throughout the Rainforest BZ. Mapped from country checklists (see order Chiroptera), other literature and museum records. Habitat Predominantly lowland rainforest, but also dense relict and riverine forests north of the Rainforest BZ. Abundance

Uncertain: rarely collected.

Remarks Day-roosts include caves, hollow trees and hollow logs, a mine-shaft and in the roof of a hut (Eisentraut 1956, Verschuren 1957, Rosevear 1965, Brosset 1966). Reported roosting singly and

Geographic Variation No subspecies. Eisentraut (1964) reported that six specimens from Bioko I. are larger than those from the mainland (especially ear length). However, the differences are small. For example: Bioko I.: FA: 53.5 (51–56) mm; T: 31.3 (26–38) mm; E: 24.0 (23– 25) (n = 6). Cameroon: FA: 52.4 (49–56) mm; T: 26.1 (22–32) mm; E: 22.0 (20–24) mm, n = 26. Similar Species Only two other Rhinolophus in sub-Saharan Africa have the following combination of characters: anterior upper

Rhinolophus alcyone

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in small groups of unknown composition (Rosevear 1965). Has been seen flying over water, swampy areas and along forest paths. Diet not known. Echolocation call-shape FM/CF/FM; CF-frequency (Uganda) 87 kHz (Pye & Roberts 1970, Roberts 1972). Littersize: one (n = 5). Reproductive chronology not known. At ca. 6° N in Côte d’Ivoire, 4 of 4 "" were pregnant in mid-Feb (CR: 19–26 mm) (Lim & Van Coeverden de Groot 1997). Ectoparasites include bat-flies Phthiridium inopinatum (Diptera: Nycteribiidae), Raymondia allisoni (Diptera: Streblidae) (Anciaux de Faveaux 1984). Conservation

IUCN Category: Least Concern.

Measurements Rhinolophus alcyone FA: 52.3 (48–56) mm, n = 100 WS (a): 342.4 (320–355) mm, n = 9

TL: 87.4 (75–100) mm, n = 49 T: 27.0 (18–32) mm, n = 57 E: 22.2 (19–25) mm, n = 74 NL (breadth): 10.0 (8.3–11.2) mm, n = 26 Tib: 24.2 (21–27) mm, n = 35 HF: 12.2 (11–13) mm, n = 25 WT: 15.4 (14–23) g, n = 23 CrnC: 22.3 (21.0–23.3) mm, n = 30* GWS: 11.8 (11.2–12.7) mm, n = 24 C–M3: 8.7 (7.9–9.2) mm, n = 32 Throughout geographic range (BMNH, HZM, ROM and literature) *Csorba et al. 2003 Key References

Rosevear 1965; Csorba et al. 2003. Meredith Happold

Rhinolophus blasii BLASIUS’S HORSESHOE BAT (PEAK-SADDLE HORSESHOE BAT) Fr. Rhinolophe de Blasius; Ger. Blasius Hufeisennase Rhinolophus blasii Peters, 1867. Monatsber. K. Preuss. Akad. Wiss. Berlin 1866: 17 [publ. 1867]. SE Europe; restricted to Italy by Ellerman et al. (1953: 59).

Taxonomy Species-group: landeri. Synonyms: andreinii, blasiusi, brockmani, clivosus Blasius, 1857, empusa, meyeroehmi. Subspecies: four; three in Africa. Chromosome number (South Africa): 2n = 58; aFN = 60. Two pairs biarmed chromosomes, 26 pairs acrocentric chromosomes; X = large submetacentric; Y = small metacentric (Rautenbach 1986). Description Small microbat with noseleaf (posterior component subtriangular with erect tip); small for an African rhinolophid; anterior upper premolar within toothrow; connecting process rising to high, narrow, forward-curving pointed horn; sella wedge-shaped; horseshoe breadth 7.2–9.0 mm; no axillary tufts; first phalanx of fourth finger relatively long; no marked contrast between crown areas of anterior and posterior lower premolars. Sexes similar. Pelage dense, soft, fluffy; mid-dorsal hairs 8–9 mm. Dorsal pelage greyishfawn to brownish-grey; hairs pale greyish-fawn or pale brownishgrey, with darker tip. Ventral pelage considerably paler. Only one individual in orange-phase has been recorded (Ansell 1974). No axillary tufts on adult !!. Ears comparatively and relatively short (16–21 mm, 33–44% of FA), dark greyish-brown. Noseleaf with lancet subtriangular with slightly concave sides (sometimes hastate), tip rounded. Connecting process well developed, rising to high, narrow, pointed horn (Figure 59j). Sella naked, wedge-shaped with sides converging towards top; top narrow and tilted forward (Figure 60e). Horseshoe narrow (7.2–9.0 mm), not covering whole muzzle but on average broader than in R. euryale; lateral leaflets absent, rudimentary or well developed (probably depending on subspecies); median emargination present but indistinct. Lower lip with three grooves: the two lateral grooves are poorly defined in R. b. empusa. Wings and interfemoral membrane dark greyish-brown. First phalanx of fourth finger relatively long (25.8 [24–28]% of fourth metacarpal, n = 30) and usually >50% of second phalanx (cf. R. euryale and R. mehelyi). Tibia 42.1 (39–45)% of FA (n = 31).

Skull delicate; zygomatic arches narrow; zygomatic width = mastoid width. Nasal swellings relatively low. Frontal depression shallow to very shallow; supraorbital ridges poorly developed (Csorba et al. 2003). Sagittal crest usually low. Palatal bridge 32–35% of C–M3. Upper incisors weakly bilobed. Upper canine with weak anterior and posterior cusps. Anterior upper premolar small (but moderate to relatively large for a rhinolophid), within toothrow; canine and posterior premolar well separated. Molar width more than half width of palate between molars (cf. R. landeri). Middle lower premolar minute and either within toothrow or displaced labially, or absent; anterior and posterior lower premolars usually well separated. Crown area of anterior lower premolar equal to or only slightly less than that of posterior lower premolar. Dental formula 1123/2133 = 32 or 1123/2123 = 30. Geographic Variation subspecies in Africa.

Koopman (1994) recognizes three

R. b. blasii: NW Africa (and extralimitally southern Europe and SW Asia). R. b. andreinii: Ethiopia and Somalia. R. b. empusa: S DR Congo to the former Transvaal. Similar Species Only one other Rhinolophus in Africa has a connecting process which rises to a high narrow pointed horn: Rhinolophus euryale. Sella parallel-sided, horseshoe usually narrower (6.5–7.5 mm). First phalanx of fourth finger relatively short (17– 22% of fourth metacarpal). Marked contrast between crown areas of anterior and posterior lower premolars. NW Africa. Two potentially sympatric species (with anterior upper premolar within toothrow) have triangular connecting processes, which, although lower, also rise to a point:

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R. mehelyi. Usually larger (FA: 48–53 mm). Sella parallel-sided; horseshoe usually narrower (4.9–7.5 mm). First phalanx of fourth finger relatively short (19–23% of metacarpal). Marked contrast between crown areas of anterior and posterior premolars. N Africa. R. landeri. Horseshoe narrower (6.0–8.0 mm). First phalanx of fourth finger relatively short (19–23% of metacarpal). Axillary tufts present in !!. South of Sahara. Distribution In Africa, occurs in three isolated regions, each occupied by a different subspecies. In NW Africa, found in the Mediterranean Coastal BZ (and marginally in the Sahara Arid BZ). In Ethiopia and Somalia, found in the Afromontane–Afroalpine BZ (and marginally in the Somalia–Masai Bushland BZ). In south-central and southern Africa, found in the Zambezian Woodland, Afromontane– Afroalpine and Coastal Forest Mosaic BZs. Two specimens in BMNH, recorded from ‘Cape of Good Hope’, probably came from Mozambique or KwaZulu–Natal (Roberts 1951). Extralimitally: Southern Europe and SW Asia. Mapped from country checklists (see order Chiroptera), other literature and museum records. Habitat In NW Africa, recorded from Mediterranean sclerophyllous forests, sub-Mediterranean semi-desert grassland and shrubland, and in stone or gravel desert habitats with wadis (but perhaps only where water is available). In Algeria, most localities are near caves and/or streams (Kowalski & Rzebik-Kowalska 1991). In Ethiopia and Somalia, most localities are in montane vegetation, evergreen and semi-evergreen bushland and thicket, and Acacia– Commiphora bushland and thicket. In Malawi and Zambia, found between 500–2300 m, in miombo woodland and montane forests (Ansell 1978, Happold et al. 1987). In KwaZulu–Natal, South Africa, most records are from major river valleys in Lowveld, Valley Bushveld and Mistbelt bioregions (Taylor, P. 1998). Abundance Uncertain. Appears common at least in some parts of geographic range (e.g. very commonly recorded in Malawi; Happold & Happold 1997). Adaptations Aspect ratio low; wing-loading low; wing-tip rounded (M. Happold unpubl.). Sometimes flies slowly with shallow wing-beats and some gliding and dipping, but can also put on bursts of fast flying with sudden turns. Can take off from ground, hover briefly; flies with great manoeuvrability; turns by banking (minimum radius mastoid width; infraorbital bridge much longer and slender. West Africa.

Rhinolophus hilli

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Measurements Rhinolophus hilli FA: 54, 54 mm WS: n. d. TL: 92, – mm T: 30, – mm E: 29, – mm NL (breadth): 12.0, – mm Tib: 24, – mm HF: 12, – mm WT: –, 16.5 g

CrnC: 23.0, – mm GWS (MW): 10.9, 11.2 mm C–M3: 8.1, 7.9 mm Rwanda (ZMUZ 126639 [holotype] and RMCA 82006-M-1 respectively; both "") Key References 1980.

Aellen 1973; Fahr et al. 2002; Smith & Hood Jakob Fahr

Rhinolophus hillorum UPLAND HORSESHOE BAT Fr. Rhinolophe des collines; Ger. Hochland-Hufeisennase Rhinolophus hillorum Koopman, 1989. Amer. Mus. Novit. 2946: 4. John Hegbe Farm, near Zozoma, 2 miles SW Voinjama, Lofa County, NW Liberia.

Taxonomy Originally Rhinolophus clivosus hillorum. Speciesgroup: ferrumequinum. Proposed as a subspecies of Rhinolophus clivosus by Hill (1968, 1982a) and described as such by Koopman (1989) but elevated to species rank by Cotterill (2002a). Synonyms: none. Chromosome number: not known.

Geographic Variation Quite uniform throughout geographic range but data are limited. A specimen from Lotti Forest, Sudan, which perhaps represents R. hillorum (see Distribution), is somewhat smaller than other specimens (FA: 53 mm, CrnC: 23.0 mm, GWS: 12.4 mm, C–M3: 8.9 mm).

Description Small microbat with noseleaf (posterior component subtriangular with erect tip); medium-sized for an African rhinolophid; anterior upper premolar absent; connecting process high but rounded; lancet short, mostly narrow and almost parallel-sided; sella parallel-sided; no axillary tufts. Sexes similar. Pelage soft, fluffy; mid-dorsal hairs ca. 10 mm. Dorsal pelage medium brown to greyish-brown. Ventral pelage paler. Orange-phase not yet reported. No axillary tufts on adult !!. Ears comparatively and relatively short (21–24 mm, 40.7 [37–44]% of FA); each with 11–12 internal folds. Noseleaf with lancet short and, except at base, very narrow and almost parallel-sided; tip hairy and slightly rounded (Figure 61f). Connecting process narrow and high but rounded, ellipsoid in profile and liberally furnished with hairs, much higher than sella. Sella naked, almost parallel-sided, diverging slightly towards the rounded top. Horseshoe narrow (8.4–9.1 mm); not completely covering muzzle; no lateral leaflets; median emargination distinct. Lower lip with one groove. Wings and interfemoral membrane blackish-brown. First phalanx of fourth finger relatively long (28.2 [27–31]% of fourth metacarpal, n = 10). Tibia 42.8 (41–44)% of FA, n = 10. Baculum trumpet-shaped, shaft dorsoventrally flattened, length 2.9–3.1 mm, n = 2 (Cotterill 2002a). Skull very robust; zygomatic arches thick and broad; zygomatic width much greater than mastoid width. Rostrum very broad. Nasal swellings very low, frontal depression very shallow. According to Csorba et al. (2003), supraorbital crests ill-defined. Sagittal crest anteriorly well developed, posteriorly moderately developed. Dentition robust. Anterior upper premolar absent; canine and posterior premolar in contact. Middle lower premolar absent; anterior and posterior lower premolars in contact. Anterior lower premolar half to two-thirds of the height, and half the crown area, of the posterior premolar. Dental formula 1113/2123 = 28.

Similar Species Only two other Rhinolophus occurring south of the Sahara and north of the Equator have the following combination of characters: anterior upper premolar fully displaced labially or absent; connecting process rounded; sella naked (or with sparse short hairs only) (Table 14, p. 304): Rhinolophus clivosus. Connecting process lower and broader in profile. Skull smaller and less robust (CrnC: 18.1–22.8 mm); dentition weaker, C–M3: 6.7–8.9 mm. Anterior upper premolar usually present. Not known in West Africa. R. deckenii. Horseshoe broader (9.1–11.5 mm). Skull with nasal swellings moderately high. Frontal depression moderately deep; supraorbital ridges prominent. Anterior upper premolar present or absent; canine and posterior premolar usually separated by narrow gap. East Africa. Distribution Endemic to Africa. Known only from three small areas within the Rainforest BZ (Western and West Central Regions) and Afromontane–Afroalpine BZ, and perhaps from one locality in the Eastern Rainforest–Savanna Mosaic. Recorded from 12 localities in Guinea, Liberia, Nigeria, Cameroon and possibly Sudan. Records from Mt Nimba, Guinea (MNHN, published as R. fumigatus by Brosset 1984) and from R. Peblei, Liberia (IRSN, tentatively identified as R. alcyone by Verschuren 1976) represent R. hillorum (Fahr et al. 2006). Specimens from Tokadeh, Mt Nimba (BMNH) were erroneously listed by Csorba et al. (2003) from both Guinea and Liberia; Tokadeh is on the Liberian side of Mt Nimba. A record from Sapoba F. R., Nigeria (M. E. Gartshore in Fedden & Macleod 1986, Cotterill 2002a), is unusual because the locality is situated in lowland rainforest and therefore this specimen should be re-examined. A specimen from Lotti Forest in the Imatong Mts, Sudan (FMNH 67500, published as R. clivosus keniensis by Koopman

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1975) has characters that indicate it might be referable to R. hillorum but the specimen needs re-examination. Habitat Found in montane forests in the highlands of SE Guinea– NW Liberia and the Cameroon Highlands, and in lowland rainforests and coastal forests in the immediate vicinity of hilly or mountainous landscapes. Found up to 1400 m on Mt Nimba, up to 1950 m on Mt Kupé, Cameroon, and at 1800 m at L. Manenguba, Cameroon. The vegetation has been described as montane grassland (Brosset 1984), montane forest (Fedden & Macleod 1986), dense rainforest (Verschuren 1976), secondary forest with surrounding primary forest (Wolton et al. 1982) and gallery forest amidst savanna at the foot of a mountain. The specimen from Lotti Forest, Sudan, was taken in East African Montane Forest at an altitude of ca. 1500 m. Abundance No detailed information but apparently very localized and rare. Adaptations Has been recorded roosting by day in caves and artificial equivalents such as mines and bridges (Verschuren 1976, Brosset 1984, Fahr et al. 2006). Has been found sharing roosts with Lissonycteris angolensis smithii, Rhinolophus simulator alticolus and R. guineensis (Brosset 1984, as R. fumigatus). During the day at ca. 1400 m, R. hillorum and R. guineensis were torpid in their roosts while R. simulator alticolus was active (Brosset 1984). Foraging and Food No information. Based on body size, probably forages, at least some of the time, by perch-hunting. The powerful skull and teeth suggest that this bat can feed on relatively large and hard-shelled insects. Echolocation

No information.

Social and Reproductive Behaviour There are three reports of these bats roosting in small groups. In Diécke Forest, Guinea, two colonies roosted under small concrete bridges; one colony comprised four !! and one ", and the other comprised one ! and three "" (Fahr et al. 2006). One specimen in Liberia was taken from a colony of 10 individuals (sexes not recorded) (Verschuren 1976). Reproduction and Population Structure Litter-size: one (n = 1). Reproductive chronology not known. A " from 07° 29' N (R. Peblei, Liberia) was pregnant in late Jan (Verschuren 1976). Predators, Parasites and Diseases Ectoparasites include batflies Brachytarsina africana (Diptera: Streblidae) and a species of the Raymondia intermedia-group (Diptera: Streblidae) (Wolton et al. 1982). Conservation IUCN Category: Near Threatened. Close to qualifying for Vulnerable. Distribution small and disjunct

Rhinolophus hillorum

(five locations). Population size low; trend inferred to be declining as result of deforestation of montane forest within area of occupancy. Small colonies roosting in caves are potentially threatened by exploitation for bushmeat. Populations in the highlands of SE Guinea–NE Liberia are particularly at risk from on-going and planned large-scale mining (Fahr & Ebigbo 2003). Measurements Rhinolophus hillorum FA: 54.5 (52–57) mm, n = 17 WS: n. d. TL: 101.5 (94–112) mm, n = 9 T: 36.4 (30–41) mm, n = 9 E: 22.4 (21–24) mm, n = 7 NL (breadth): 8.8 (8.4–9.1) mm, n = 7 Tib: 23.6 (22–25) mm, n = 10 HF: 13.2 (12–14) mm, n = 6 WT: 20.2 (16.5–25.0) g, n = 9 CrnC: 23.8 (23.3–24.4) mm, n = 7 GWS: 13.0 (12.3–13.5) mm, n = 9 C–M3: 9.2 (8.9–9.5) mm, n = 13 Guinea, Liberia, Cameroon (AMNH incl. holotype, BMNH, FC, IRSN, MNHN, SMNS) Key References Cotterill 2002a; Hill 1968, 1982a; Koopman 1989; Koopman et al. 1995. Jakob Fahr

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Rhinolophus hipposideros LESSER HORSESHOE BAT Fr. Petit rhinolophe; Ger. Kleine Hufeisennase Rhinolophus hipposideros (Bechstein, 1800). In: Pennant, Allgemeine Uebers. Vierfüss. Thiere 2: 629. France.

Taxonomy Originally Noctilio hipposideros. Species-group: hipposideros. Synonyms in Africa: escalerae, minimus (extralimitally 21 others listed by Simmons [2005]). Subspecies: controversial (see Geographic Variation). Chromosome number (Europe and Asia): 2n = 54, 56, 58, 62 (Zima et al. 1992, Horáček & Zima 1996, Benda & Horáček 1998). This is the only rhinolophid in which chromosomal polymorphism has been reported. However, the older 54 karyotype should be re-examined, and the 62 karyotype probably represents another species (Horáček et al. 2000). There are no data from Africa. In specimens from Europe and Asia, 4–10 metacentric and 44–52 acrocentric chromosomes are reported, X is metacentric or acrocentric, Y acrocentric or dot-like (Zima et al. 1992, Horáček & Zima 1996, Benda & Horáček 1998). Description Very small microbat with noseleaf (posterior component subtriangular with erect tip); smaller than any other rhinolophid in Africa (FA: 35–40 mm in Africa); anterior upper premolar within toothrow; connecting process very low and either slightly rounded or flat; lancet long and wedge-shaped; sella wedge-shaped and backward-sloping with the top curving forward; no axillary tufts. Sexes similar. Pelage soft, fluffy; mid-dorsal hairs ca. 10 mm. Dorsal pelage greyish-brown to medium brown (dark grey in juveniles); hairs of adults pale beige with greyishbrown or brown tip. Ventral pelage paler or grey to greyish-white. Apparently no orange-phase. No axillary tufts on adult !!. Ears comparatively short (15–17 mm) but of short–medium relative length (mean E ca. 47% of mean FA). Noseleaf with lancet subtriangular with slightly concave sides, tip bluntly pointed. Connecting process low (not rising above sella), slightly rounded (Figure 59m) or sometimes flat. Sella naked, long, narrow, wedgeshaped; top pointed and curves forward and downward. Horseshoe narrow (6.1–7.1 mm in Africa) but almost covering muzzle; no lateral leaflets; median emargination a distinct notch. Wings and interfemoral membrane medium to dark brown. First phalanx of fourth finger relatively long (26.0 [24–28]% of fourth metacarpal, n = 31). Tibia 45.4 (41–49)% of FA, n = 10. Skull very delicate; zygomatic arches extremely slender; zygomatic width slightly greater or almost equal to mastoid width. Nasal swellings of medium relative height. Frontal depression shallow, supraorbital ridges weak. Sagittal crest low and extending partway across parietals. Palatal bridge (African specimens) relatively short to medium (29–33% of C–M3; G. Csorba pers. comm.). Anterior upper premolar relatively large (reaching one-third height of canine), within toothrow; canine and posterior premolar well separated. Middle lower premolar very small, usually fully displaced labially; anterior and posterior lower premolars in contact or nearly so. Dental formula 1123/2133 = 32. Geographic Variation Controversial. Six subspecies are currently recognized by Csorba et al. (2003) and Simmons (2005), of which two occur in Africa:

R. h. escalerae: NW Africa. Considered subspecifically distinct on basis of size and also the narrowness of the bar between the infraorbital foramen and the orbit (Gaisler 1983, Kowalski & Rzebik-Kowalska 1991, Steiner & Gaisler 1994). FA: 35.8 (35–38) mm, n = 9. R. h. minimus. Southern Europe to eastern end of Mediterranean, including several islands, and southwards to Sudan, Ethiopia and SW Arabia. Possibly represents a distinct species (see Zagorodniuk 1999). FA: 37.7 (36–40) mm, n = 10. Extralimitally: majori (Corsica), minutus (Britain and Ireland), hipposideros (continental Europe to eastern end of Black Sea), midas (Transcaucasia and Iraq to Kazakhstan and Kashmir). In contrast, only hipposideros and midas have been recognized by majority of authors in the past (Horáček et al. 2000). Aulagnier & Thévenot (1986) included both minimus and escalerae in the nominate subspecies. Similar Species No other Rhinolophus in Africa is so small, and none has the combination of a connecting process that is very low and either slightly rounded or flat and a sella that is wedge-shaped and backward-sloping with the top curving forward (Table 14, p. 304). Distribution In Africa, recorded from almost all of the Mediterranean Coastal and Afromontane–Afroalpine BZs in NW Africa (Aellen & Strinati 1969, Aulagnier & Thévenot 1986, Kowalski & Rzebik-Kowalska 1991), and from the Afromontane–Afroalpine BZ of the Ethiopian Highlands and adjacent Somalia–Masai Bushland BZ in Sudan, Ethiopia, Eritrea and Djibouti (Largen et al. 1974, Koopman

Rhinolophus hipposideros

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1975, Pearch et al. 2001). Also one record from the Sahara Arid BZ in Sinai (Egypt) (Qumsiyeh 1985). Not known in Libya. In Morocco, recorded from the northern region south to Anti-Atlas and the preSaharan region (29° N); in Algeria from the coast to the southern limit of Saharan Atlas (33° N), and in N Tunisia to Kasserine (35° N) (Aellen & Strinati 1969, Aulagnier & Thévenot 1986, Kowalski & RzebikKowalska 1991). Extralimitally: Ireland to Tajikistan, Kashmir and Arabian Peninsula (see Geographic Variation).

are emitted during slow-hawking (Jones & Rayner 1989a). CFfrequency 105–113 kHz; call-duration 20–30 ms; maximum energy in second harmonic (but some in first harmonic); average terminal FM bandwidth 29.1 kHz suggesting R. hipposideros relies particularly heavily on FM information (Jones & Rayner 1989a). For hand-held bats, the CF-frequencies of calls emitted by bats more than one year old were higher than those emitted by younger bats, and "" emit higher frequencies than !! (Jones et al. 1992).

Habitat In NW Africa, this cave-roosting species seems to be rare and its foraging habitats are little known. In Algeria, recorded from sea level to ca. 1500 m; three!! mist-netted in a bushy forest with a permanent stream (Sebdour, Tlemcen Mts), one " mist-netted over water in a desert environment (Brezina, Saharan Atlas [Gaisler & Kowalski 1986, Kowalski & Rzebik-Kowalska 1991]). One specimen was taken beside a lake in an area of grassland, thorn scrub, volcanic rubble and lava blister caves, at ca. 1000 m, in Awash N. P. (Hill & Morris 1971).

Social and Reproductive Behaviour In North Africa in winter, single individuals, mostly !!, were sporadically seen roosting in caves (Kowalski et al. 1986); no other information. Extralimitally (Europe), during winter, hibernates singly or in colonies of up to 500 individuals, which hang apart, 25–50 cm from their neighbours (Macdonald & Barrett 1993). In summer, "" form maternity colonies of 10–800 individuals; immature !! sometimes present; "" hang apart except when heavily pregnant in cool weather, or when huddling with their young. Audible (to humans) vocalizations at roosts include chirping. Mating often takes place in autumn; preceded by chasing; ! hangs behind and over " during brief copulation; sometimes copulates in hibernacula.

Abundance Common in N Morocco but rare in Algeria and Tunisia, even in coastal areas. Probably rare in Sudan, Ethiopia and Eritrea. Adaptations Aspect ratio very low; wing-loading very low; wing-tip especially short and rounded (Norberg & Rayner 1987). Flight slow (possibly with some short bursts of speed), fluttering and butterfly-like with intermittent gliding; manoeuvrable. Can hover. Day-roosts in Africa include both natural and man-made underground spaces such as caves; individuals hang from ceilings or walls; when torpid and hibernating, they wrap themselves completely in the wing-membranes and draw close to the substrate by bending the legs. In winter in Algeria, they hibernate in the coolest places, usually close to openings (Kowalski et al. 1986); extralimitally, temperatures in hibernacula are 6–9 °C and humidity is high (Schober & Grimmberger 1989). Foraging and Food No information for African populations except that three foraging !! were mist-netted in bushy areas and one " above water in a desert environment in Algeria (Gaisler & Kowalski 1986, Kowalski & Rzebik-Kowalska 1991), and one individual was reported flying low over the muddy shore of a lake in Ethiopia (Hill & Morris 1971). In England and Eire, bats were observed foraging by gleaning (they usually picked non-volant prey [including larvae] from stones, rocks and vegetation without landing but sometimes pounced on prey on the ground) and by slow-hawking; not known to forage by fly-catching (Jones & Rayner 1989a). Foraged close to vegetation, either patrolling edges of river banks close to riverine vegetation, or close to walls. Some individuals regularly patrolled an ivy-covered bridge when ivy flowers were attracting moths. Apparently patrols well-defined beats. Foraging is strictly nocturnal; sometimes reported to continue for at least five hours after sunset. In Europe, diet is mainly small Lepidoptera and Diptera (Nematocera), and less often Neuroptera, Trichoptera, Coleoptera and Araneae. Echolocation No data for African populations. In England and Europe, call-shape usually FM/CF/FM with the terminal FM component greater in bandwidth; some CF and CF/FM calls

Reproduction and Population Structure Litter-size: no data for Africa. In Iran, ca. 65% of "" bear singletons, and 35% bear twins (DeBlase 1980 in Csorba et al. 2003); in Europe, litter-size is one. Reproductive chronology in Africa not known. In Europe, it is restricted seasonal monoestry with mating in autumn (Sep–Nov) or in hibernacula during winter; sperm-storage by "" until Mar–Apr when ovulation and fertilization occur; births in summer (mid-Jun to early Jul); lactation for 4–5 weeks (Gaisler 1966, Macdonald & Barrett 1993). A " with vaginal plug was recorded in Jan at Sig, NW Algeria (Kowalski et al. 1986). Extralimitally, "" reach sexual maturity in first year, but most give birth for first time when two years old. Maximum life-span: 21 years 3 months. Predators, Parasites and Diseases Preyed on, rarely, by Barn Owls Tyto alba (Cabrera 1932 in Aulagnier & Thévenot 1987). Ectoparasites include a bat-fly Phthiridium biarticulatum (Diptera: Nycteribiidae) for which R. hipposideros is the principal host (Corbet & Harris 1991). Conservation IUCN Category: Least Concern (assessed from extralimital as well as African data). Measurements Rhinolophus hipposideros FA: 36.8 (35–40) mm, n = 19 WS (d): 192–254 mm* TL: 63.0 (60–68) mm, n = 10 T: 24.2 (21–26) mm, n = 11 E: 16.1 (15–17) mm, n = 11 NL (breadth): 6.6 (6.1–7.1) mm, n = 10 Tib: 17.2 (15–18) mm, n = 10 HF: 7.0 (6–8) mm, n = 7 WT: 3.8 (3.5–4.0) g, n = 4 CrnC: 14.9 (14.3–15.5) mm, n = 12 GWS: 7.2 (6.8–7.6) mm, n = 11 339

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Family RHINOLOPHIDAE

C–M3: 5.0 (4.8–5.3) mm, n = 13 Morocco, Algeria, Sudan, Ethiopia (BMNH, Gaisler 1983, Kowalski & Rzebik-Kowalska 1991). For NW Africa, n = 9–11; for Sudan and Ethiopia, n = 1–3 (except FA, NL and Tib) *Europe (Schober & Grimmberger 1989)

Key References Aulagnier & Thévenot 1986; Csorba et al. 2003; Horáček et al. 2000; Jones & Rayner 1989a; Kowalski & RzebikKowalska 1991. Jiµí Gaisler

Rhinolophus landeri LANDER’S HORSESHOE BAT Fr. Rhinolophe de Lander; Ger. Landers Hufeisennase Rhinolophus landeri Martin, 1838. Proc. Zool. Soc. Lond. 1837: 101 [publ. 1838]. Bioko I., Equatorial Guinea.

Taxonomy Species-group: landeri. Synonyms: angolensis, axillaris, dobsoni, lobatus. Subspecies: three. Chromosome number (South Africa): 2n = 58; aFN = 60. Two pairs biarmed chromosomes and 26 pairs acrocentric chromosomes. X = large submetacentric; Y = no data (Rautenbach 1986). Description Small microbat with noseleaf (posterior component subtriangular with erect tip); small for an African rhinolophid; anterior upper premolar within toothrow; connecting process triangular; lancet hastate; sella parallel-sided; axillary tufts in adult !! reddish or reddish-brown; first phalanx of fourth finger relatively short; tibia 17–21 mm. Sexes similar. Pelage dense, soft, fluffy; mid-dorsal hairs 8–9 mm. Dorsal pelage greyish-fawn to brownish-grey; hairs pale greyish-fawn or pale brownish-grey with darker tip. Ventral pelage slightly paler. In orange-phase, dorsal pelage golden-brown, orangecinnamon to bright rusty-red. Adult !! with reddish or reddishbrown axillary tufts, which are sometimes sticky with yellow secretion. Ears comparatively and relatively short (13–20 mm, 34–42% of FA). Noseleaf with lancet hastate, tip bluntly pointed. Connecting process subtriangular with tip either sharply or bluntly pointed. Sella naked, narrow with slightly concave sides, top broad and rounded. Horseshoe narrow (6.0–8.0 mm) but covering whole

muzzle; no lateral leaflets; median emargination a deep notch. Lower lip with a well-defined median groove and two very poorly defined lateral grooves. Wings and interfemoral membrane dark greyishbrown to blackish-brown (grey-phase) or brown (orange-phase). First phalanx of fourth finger relatively short (21.0 [19–23]% of fourth metacarpal, n = 52). Tibia 42.4 (38–45)% of FA, n = 18. Skull of medium build; zygomatic arches of moderate breadth; zygomatic width slightly greater than mastoid width. Nasal swellings of medium relative height. Frontal depression usually shallow (Csorba et al. 2003). Sagittal crest low to moderately developed anteriorly, absent posteriorly. Palatal bridge 28–37% of C–M3 (Csorba et al. 2003). Anterior upper premolar small, within toothrow or only slightly displaced labially; canine and posterior premolar well separated. Molar width less than half width of palate between molars (cf. R. blasii). Lower canines not distinctly smaller than upper canines (cf. R. denti, R. simulator, R. swinnyi). Middle lower premolar small, slightly to fully displaced labially; anterior and posterior lower premolars separated by narrow gap or in contact. Anterior lower premolar (R. l. landeri) only a little smaller than posterior lower premolar; more than half and usually two-thirds of its height (Kock et al. 2002) (cf. R. guineensis). Dental formula 1123/2133 = 32. Geographic Variation Koopman (1994):

Three subspecies are recognized by

R. l. landeri: Gambia to Cameroon and south to mouth of Congo R. R. l. lobatus: Sudan and Ethiopia, and south to former Transvaal; also Zanzibar I. R. l. angolensis: W Angola. Similar Species Only two other sub-Saharan Rhinolophus have the following combination of characters: anterior upper premolar within toothrow; connecting process triangular and pointed; first phalanx of fourth finger relatively short (Table 14, p. 304): Rhinolophus guineensis. Body measurements usually larger (FA: 44– 50 mm [cf. 39–45 in R. landeri from West Africa],Tib: 21–22 mm). Skull larger (CrnC: 19.2–20.6 mm [cf. mastoid width. Rostrum broad. Sagittal crest low. Anteorbital foramen comparatively small (cf. H. cyclops), closed by narrow bar. Cochleae

Geographic Variation

None recorded.

Similar Species Only one other African Hipposideros has two median club-shaped processes on the noseleaf, and frosted blackishbrown woolly pelage (Table 15, p. 370): Hipposideros cyclops. Usually smaller (FA !!: 65.4 [61–75] mm, FA "": 68.0 [59–74]; CrnC: 28.2 [26.3–30.0] mm). Anteorbital foramen larger and closed by a moderately wide bar.

Hipposideros camerunensis

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Hipposideros curtus

Distribution Endemic to Africa. Known only, very disjunctly, from the Afromontane–Afroalpine BZ (Cameroon) (Eisentraut 1956), the Rainforest BZ (E DR Congo) (Hayman et al. 1966), and from two localities in the Eastern Rainforest–Savanna Mosaic in W Kenya (Schlitter et al. 1986). Subsequently also recorded from Budongo (00° 45' N, 31° 36' E) and Itama (00° 57' S, 29° 42' E), Uganda, by Thorn & Kerbis Peterhans (2009): not mapped. Habitat Afromontane forests at 1200–1400 m on Mt Cameroon; degraded afromontane forest with undergrowth of Acanthus arboreus and Brillantaisia in North Nandi Forest, Kenya; intermediate evergreen forest in Kakamega Forest, Kenya; and lowland rainforest at Shabunda, DR Congo. Abundance Uncertain. Rare in collections. Adaptations Based on measurements of a dried museum specimen, aspect ratio is low and wing-loading is high or very high, but this needs confirmation from unpreserved specimens. If correct, flight is predicted to be fast, agile and energetically expensive, and manoeuvrability is probably poor. Day-roosts include caves and hollow trees (Eisentraut 1973a). One individual was caught in the bottom shelf of a mist-net (Schlitter et al. 1986), suggesting that this species sometimes forages near the ground. Foraging and Food Predictably forages by fly-catching, as does H. cyclops and H. vittatus (see species profiles). Diet not known, but the massiveness of the zygomatic arches suggests that H. camerunensis can eat hard-shelled insects. Social and Reproductive Behaviour

Reproduction and Population Structure Twelve "" were pregnant in Oct in Cameroon (Eisentraut 1963). No other information. Conservation IUCN Category: Data Deficient. The possibility that the records comprise more than one species has been debated by the IUCN assessors and evaluators: because of habitat loss, the Mt Cameroon population is possibly a threatened endemic species. Measurements Hipposideros camerunensis FA: 75.9 (74–80) mm, n = 17 WS (c): ca. 340 mm, n = 1 TL: 127.7 (110–140) mm, n = 9 T: 31.8 (23–45) mm, n = 13 E: 33.6 (30–38) mm, n = 16 NL (breadth): 17.4 (14.8–18.7) mm, n = 5 Tib: 35.9 (35–39) mm, n = 21 HF: 19.0 (18–22) mm, n = 7 WT: 47.7 (39–53) g, n = 7 CrnC: 30.2 (29.2–31.5) mm, n = 14 GWS: 16.3 (15.7–16.9) mm, n = 16 C–M3: 10.7 (9.6–11.5) mm, n = 17 Cameroon, Kenya, DR Congo (BMNH, NMW, RMCA, SMNS, ZFMK and literature) Key References 1986.

Eisentraut 1973a; Hill 1963; Schlitter et al. Meredith Happold

No information.

Hipposideros curtus SHORT-TAILED LEAF-NOSED BAT Fr. Phyllorhine à queue courte; Ger. Kurzschwanz-Rundblattnase Hipposideros curtus G. M. Allen, 1921. Rev. Zool. Bot. Afr. 9: 194. Sakbayeme, Cameroon.

Taxonomy Species-group: bicolor. Synonyms: sandersoni. Subspecies: none. Chromosome number: not known. Description Small to very small microbat with noseleaf (posterior component roughly elliptical); sepia brown; ears large and separated; noseleaf with enlarged internarial septum, which partly conceals the nostrils, and two lateral leaflets on each side; frontal sac usually present in both sexes. Sexes similar. Pelage long, silky, fluffy. Dorsal pelage sepia brown; hairs buff with sepia brown at base and at tip. Ventral pelage same as dorsal pelage, or slightly paler. Orange-phase: no information. Ears separated, comparatively short but of medium relative length (35–49% of FA), rounded (length and breadth almost equal), tip bluntly pointed with sharp concavity in outer margin just below tip: each ear with 11 internal folds (n = 1). Antitragus well developed with small fold. Noseleaf as in Figure 68c. Posterior component not elongated; divided into four cells by three vertical septa; upper margin with low-arched outline. No club-like processes. Anterior component broad, almost covering muzzle. Internarial septum pad-like and moderately enlarged, forming a

longitudinally oval disc (longer than broad), which partly conceals the nostrils. Two weakly-developed lateral leaflets on each side. Frontal sac usually present in both sexes (sometimes absent in ""); opening horizontally. Wings and interfemoral membrane blackishbrown. Fifth metacarpal 89–109% of third metacarpal. Tibia 37– 49% of FA. Tail 35–51% of HB. Skull delicate, short and broad; zygomatic arches moderate; zygomatic width = mastoid width or slightly less. Sagittal crest low. Cochleae not enlarged, their breadth ca. equal to their distance apart or a little greater. Upper incisor slightly bicuspid. Upper canine relatively short (42 [38–48]% of C–M3, n = 4). Anterior upper premolar small, slightly displaced labially, canine and posterior premolar well separated. Further details in Hill (1963). Geographic Variation

None recorded.

Similar Species Only two other African Hipposideros have a noseleaf with the internarial septum enlarged (Table 15, p. 370): 379

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Family HIPPOSIDERIDAE

Hipposideros jonesi. Margin of posterior component elongated, subtriangular in outline; internarial septum larger, broader than long; one long, well-developed lateral leaflet on each side; no frontal sac. Ears usually larger (21–28 mm) and ca. 50% of FA. H. marisae. Usually smaller (FA: 40.6 [38–42] mm; CrnC: 15.6 [15.4–15.8] mm). Ears 15–17 mm; no antitragal fold. Internarial septum less enlarged, oval disc smaller; usually only one, rudimentary, lateral leaflet on each side. Frontal sac present in both sexes. Distribution Endemic to Africa. As yet, known only from the Rainforest BZ in Cameroon (where recorded from at least seven localities) and from Equatorial Guinea (both mainland and Bioko I.) (Allen 1921, Sanderson 1939, Aellen 1952, Perret & Aellen 1956, Eisentraut 1973a, J. Juste pers. comm.). Habitat

Apparently restricted to lowland rainforest.

Abundance Uncertain. Very rare in collections. Remarks Very little is known about this species (Rosevear 1965). Its flight is described as sluggish in comparison with that of H. caffer (Sanderson 1939). One individual foraged with numerous H. caffer in the verandah of a house. One was caught over a stream (Sanderson 1939). A small colony, which varied in size according to season, roosted in a shelter formed by big rocks; others have been found under a boulder in a forest (Perret & Aellen 1956). Also roosts in caves (Eisentraut 1964). Conservation IUCN Category: Vulnerable. Some of the few known roosts have disappeared and habitat is being lost as a result of selective logging, clear-cutting and other human activities. Population trend: declining. Measurements Hipposideros curtus FA: 43.7 (42–47) mm, n = 21 WS: n. d. TL: 71.5 (69–75) mm, n = 11

Hipposideros curtus

T: 20.7 (18–23) mm, n = 16 E: 18.1 (15–22) mm, n = 18 NL (breadth): 5.7 (5.1–7.2) mm, n = 12 Tib: 18.8 (16–21) mm, n = 16 HF: 7.3 (7–8) mm, n = 3 WT: 7.1 g, n = 1 CrnC: 17.0 (16.3–17.5) mm, n = 13 GWS: 9.3 (8.4–10.2) mm, n = 15 C–M3: 5.6 (4.9–7.1) mm, n = 17 Côte d’Ivoire, Cameroon, Bioko I. (BMNH, RMCA, ROM, SMNS, ZFMK and literature) Key References

Hill 1963; Rosevear 1965; Sanderson 1939. Meredith Happold

Hipposideros cyclops CYCLOPS LEAF-NOSED BAT Fr. Phyllorhine cyclope; Ger. Zyklopen-Rundblattnase Hipposideros cyclops (Temminck, 1853). Esquisses Zool. sur la Côte de Guiné, p. 75. Boutry River, Ghana.

Taxonomy Originally Phyllorrhina cyclops. Species-group: cyclops. Synonyms: langi, micaceus. Subspecies: none recognized. Chromosome number: not known. Description Medium-sized microbat with noseleaf (posterior component roughly elliptical); blackish-brown with woolly, frosted pelage; ears separated; noseleaf with two median club-shaped processes; FA: 66.6 (59–75) mm. Not easily distinguished from H. camerunensis. Females significantly larger and heavier than !! in most body measurements. Pelage dense, soft, woolly; extending along proximal half of forearm; mid-dorsal hairs 11–13 mm (18–

19 mm at high altitudes). Dorsal pelage blackish-brown with white or silvery frosting; hairs blackish-brown with curly, white or silvery tip. Head greyish-brown with darker eye-rings. Ventral pelage paler than dorsal pelage with less conspicuous frosting. No orange-phase. Ears separated, comparatively medium to long but of medium relative length (42–53% of FA), narrow, medium to dark brown, inner margin convex, outer margin convex becoming concave near tip, tip narrowly pointed. Eyes comparatively large. Noseleaf as in Figure 68a. Posterior component divided into four cells by three vertical septa, and with a club-shaped process arising from middle of posterior edge. A second median club-shaped process arises

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Hipposideros cyclops

from the central component above the nostrils. Internarial septum weakly developed, not concealing nostrils. Two well-developed lateral leaflets on each side, the lower pair continuous with the base of the posterior component. Frontal sac present and prominent in both sexes; opens vertically. It is lined with long, stiff, white hairs that form a conspicuous tuft when the sac is everted. At each side of the frontal sac there is a glandular patch or shallow sac. Wing and interfemoral membranes blackish-brown; skin on forearm, digits and tibia much paler reddish-brown. Fifth metacarpal 96 (92–100)% of third metacarpal. Tibia 42–52% of FA. Tail 45–49% of HB. Males have a (glandular?) sac lined with long, stiff, reddish-brown hairs, between penis and anus, opening posteriorly just in front of anus (Figure 74). Females lack this anal sac but have a patch of bare skin with scattered long and stiff hairs near the vagina. Skull robust, elongate with broad rostrum and elongated braincase; zygomatic arches massive; zygomatic width much greater than mastoid width. Rostrum broad. Sagittal crest low. Infraorbital (= anteorbital) foramen large, rounded and closed by a moderately wide bar (cf. H. camerunensis). Cochleae greatly enlarged, their breadth four times their distance apart (Hill 1963) (Figure 73d). Upper incisor slightly bicuspid. Upper canine powerful, relatively short (42 [37–46]% of C–M3, n = 8). Anterior upper premolar very small, fully displaced labially; canine and posterior premolar in contact. Anterior lower premolar ca. one-third to half the length and half the height of the posterior premolar.

a

b

c

Figure 74. Pubic region of adult ! Hipposideros cyclops showing (a) the anal sac in its invaginated state and (b) in its everted state. (c) Pubic region of adult " H. cyclops showing the patch of bare skin with scattered long, stiff hairs near the vagina. All based on Allen (1917a). The pubic region is the same in H. camerunensis.

Geographic Variation Variable throughout geographic range although no subspecies are currently recognized. Populations from drier habitats have significantly longer forearms than those from wetter habitats as exemplified (in Côte d’Ivoire) by bats from the Guinea Savanna BZ (Comoé N. P.) with FA: 67.1 (62–72) mm (n = 45) compared to those from the Rainforest BZ (Taï N. P.) with FA: 65.2 (62–70) mm (n = 28). Specimens from E DR Congo, W Uganda and N Burundi seem to attain larger dimensions, but samplesize is limited. Similar Species Only one other African Hipposideros has two median club-shaped processes on the noseleaf, and frosted blackishbrown woolly pelage (Table 15, p. 370): Hipposideros camerunensis. Usually larger (FA: 75.9 [74–80] mm; CrnC: 30.2 [29.2–31.5] mm). Infraorbital foramen smaller and closed by a narrow bar. Not known west of 9° E, but geographical ranges overlap from Cameroon to Kenya. Distribution Endemic to Africa. Recorded from the Rainforest BZ (Western, West Central and East Central Regions), the adjacent Northern and Eastern Rainforest–Savanna Mosaics and Afromontane–Afroalpine BZ, with an apparently isolated population in the Eastern Arc Mts and the Coastal Forest Mosaic BZ of Kenya and Tanzania. Recorded, somewhat disjunctly, from Senegal to coastal Kenya and Tanzania. Gaps in Guinea and Nigeria might reflect insufficient sampling. There are very few records from the central Congo Basin and it remains to be established if the species ranges throughout the entire Congolian rainforest zone. The record from Bamingui-Bangoran N. P., Central African Republic, appears to be very isolated.

Hipposideros cyclops

Habitat Recorded mainly from lowland rainforest but also from coastal forest, montane forest, swamp forest and mangroves vegetation zones. Also extends far into the Rainforest–Savanna Mosaic, and into the Guinea Savanna BZ where sufficiently large relict forests and gallery forests are present. Although mostly recorded in undisturbed forest, also found in secondary forest, highly degraded areas, and converted habitats such as cocoa and rubber plantations (Lang & Chapin 1917b, Jeffrey 1975, Fedden & 381

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Family HIPPOSIDERIDAE

Macleod 1986, Schlitter et al. 1986, Juste & Ibáñez 1994, USNM records). Highest reported altitudes are 1000–1200 m on Mt Nimba (Liberia), 1100 m on Mt Kupé (Cameroon) (Hill 1968, Wolton et al. 1982) and 1950 m in Kibira N. P., Burundi (FMNH). Abundance Widespread and comparatively abundant in suitable habitat. The most frequently captured hipposiderid in Comoé N. P., Côte d’Ivoire, and the second most frequently captured hipposiderid in Taï N. P. Local distribution and abundance is probably determined by the availability of suitable day-roosts. Adaptations Flight moderately fast, manoeuvrability poor; energetic cost of flight probably high (J. Fahr & K. Soer unpubl.). By day, most often found roosting in spacious hollow trunks of standing trees, including Borassus-palms, Ceiba pentandra, Cola cordifolia, Cordia sp., Hallea stipulosa, Klainedoxa gabonensis, Macaranga sp., Pseudospondias microcarpa, Ricinodendron heudelotii and Terminalia superba (Decher & Fahr 2005a). Very occasionally found inside fallen logs, old wells and disused mines, and once found in a belfry. Never found in hollow Cynometra megalophylla in Comoé N. P. although this tree is very abundant in gallery forests and its hollow trunks seem suitable. Often shares day-roosts with other bats (Lissonycteris angolensis, Nycteris arge, N. major, Rhinolophus alcyone, R. landeri), anomalures (Idiurus macrotis, Anomalurus spp.), African Dormice (Gliridae) and Tullberg’s Softfurred Mouse Praomys tullbergi (Decher & Fahr 2005a). The recapture rate was 17% (n = 35) in Taï N. P. and 20% (n = 59) in Comoé N. P. (J. Fahr unpubl.). Most individuals were recaptured up to one year after marking, and two individuals were recaptured after three years. All were recaptured less than 400 m from the initial site, most less than 250 m, suggesting that home-ranges are very small (several ha) and that site-fidelity is unusually high. The eyes of H. cyclops are large in comparison with those of most other African hipposiderids, suggesting that vision plays a more important role in orientation than in other species of bats. Together with its sister species H. camerunensis, it is possibly the only African Hipposideros that does not have an orange-phase. The frosted pelage perhaps helps to camouflage these bats while they are exposed to predators during perch-hunting (see Foraging). Foraging and Food Insectivorous. Forages by fly-catching and, very rarely, by slow-hawking. Usually forages not far from the ground or vegetation. Median foraging height in Côte d’Ivoire (as determined by captures in mist-nets set 0–25 m above ground) was 1.9 (0.4– 23.6) m, n = 91, J. Fahr unpubl.). Most individuals were caught 1–8 m above ground, but six were caught between 13.5 and 23.6 m. They are specialized fly-catchers and rarely fly except when commuting from day-roosts to feeding areas, flying from one perch to another, or attacking prey. They emerge from their day-roosts comparatively early (18:00–18:30h) and fly to perches, such as tree trunks, branches and twigs, which are usually 2–6 m above ground. There they rotate from side to side while echolocating and scanning the surroundings for flying insects. Detected insects are captured in flight, and either carried back to the perch, or to the day-roost, where they are divested of their wings and other hard parts, and then consumed. Verschuren (1957) analysed discarded wings and other remains collected in Garamba N. P., NE DR Congo, and found predominantly hawk-moths (Sphingidae) and cicadas but also owl-flies (Neuroptera:Ascalaphidae),

flat-bugs (Heteroptera:Aradidae), wasps (Hymenoptera: Eumenidae) and beetles (Coleoptera: Scarabaeidae, Elateridae). Additionally, Verschuren (1957) found bark-lice (Psocoptera), moth-flies (Diptera: Psychodidae) and ants (Hymenoptera: Formicidae) in stomach contents. In Comoé N. P. these bats predominantly took hawk-moths, cicadas and beetles, but also ant-lions (Neuroptera: Myrmeleontidae), grasshoppers and crickets (Orthoptera: Acrididae, Gryllidae) and winged male driver-ants (Hymenoptera: Formicidae: Dorylus spp.) (K. Soer & J. Fahr unpubl.). Echolocation Call-shape CF/FM. CF-frequency in resting bats (Côte d’Ivoire) 59.7 (58.4–60.8) kHz (J. Fahr & N. Ebigbo unpubl.). The CF-frequency of 101–109 kHz given by Novick (1958b) is erroneous and possibly the result of the recording equipment at that time (J. Fahr unpubl.). Social and Reproductive Behaviour Roosts singly or in small to medium-sized groups comprised of 1–3 !! and several "" (Aellen 1952, Verschuren 1957, Fedden & Macleod 1986, J. Fahr unpubl.). The largest group found in Comoé N. P. consisted of 18 individuals (not sexed). Lang & Chapin (1917b) and Eisentraut (1956) reported group-sizes of 12 individuals. The mean ratio of !! to "" in colonies was 1 : 1.8 in Garamba N. P. (Verschuren 1957). Individuals roosting singly are mostly !!. The frontal sac in both sexes, and the anal sac in !!, are likely to play an important role in olfactory communication although the specific function is not known. The anal sac, when everted, emits a very strong, almost pungent odour. Not yet known if the odour is produced by glands or by bacterial fermentation of excretions.Young are not left in dayroosts during the night but are carried by their mothers, even during perch-hunting (J. Fahr pers. obs.). Reproduction and Population Structure Litter-size: one. Reproductive chronology uncertain. In the Northern Rainforest– Savanna Mosaic (Fintonia, NW Sierra Leone; Comoé N. P., NE Côte d’Ivoire; Garamba N. P., NE DR Congo), 11 of 14 adult "" were pregnant, two were lactating and one was neither pregnant nor lactating in Feb–Apr; 10 of 15 "" were lactating and five were neither pregnant nor lactating in May–Jun; none of 18 "" was pregnant or lactating in Oct–Nov; no data for other months (Verschuren 1957, J. Fahr unpubl., USNM). According to Verschuren (1957), in Garamba N. P., "" are in fairly close reproductive synchrony; births take place in mid-Mar, lactation ends in mid-May, and there is no evidence of a second parturition season; young are born with a FA ca. 25 mm. In the Rainforest BZ of Côte d’Ivoire, none of 18 adult "" was pregnant or lactating in Jan–Feb; 1 of 11 was pregnant, one was lactating and nine were neither pregnant nor lactating in Mar; none of 3 "" was pregnant or lactating in Jun; 8 of 14 "" were pregnant and six were neither pregnant nor lactating in Jul–Oct; 1 of 1 was lactating in Dec; no data for other months (J. Fahr unpubl., CM, MHNG, ROM, SMF, USNM). None of the females was found simultaneously lactating and pregnant (n = 34). These data are not conclusive but they are compatible with restricted seasonal monoestry in the Rainforest–Savanna Mosaic (with lactation coinciding with the onset of the wet season), and extended seasonal monoestry in the Rainforest BZ (with the majority of "" giving birth towards the end of the wet season).

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Hipposideros fuliginosus

The ratio of !! to "" as determined by captures with mistnets (excluding recaptures) in Comoé N. P. was 1 : 0.6 (n = 55); in Taï N. P., it was 1 : 0.7 (n = 29; J. Fahr unpubl.). Predators, Parasites and Diseases Remains of one individual were found in scats of an unidentified small carnivore (probably genet Genetta sp.) in Central African Republic (Hutterer & Ray 1997). Ectoparasites include bat-flies Raymondia brachyphysa, R. intermedia (Diptera: Streblidae) and a mite Steatonyssus hipposideros (Acari: Macronyssidae) (Jobling 1956). In Taï N. P., 23% of the individuals (n = 35) were heavily infested with unidentified, bright orange mites (J. Fahr unpubl.). Conservation IUCN Category: Least Concern. Probably most threatened by destruction of suitable day-roosts and direct exploitation therein.

TL (!!): 109.6 (99.0–133.0) mm, n = 45 TL (""): 113.1 (95.0–128.0) mm, n = 79 T: 83.6 (18–36) mm, n = 124 E: 33.5 (28–38) mm, n = 125 NL (breadth): 14.9 (12.9–15.7) mm, n = 8 Tib: 32.2 (29–35) mm, n = 20 HF: 20.1 (18–22) mm, n = 114 WT (!!): 29.2 (21–40) g, n = 81 WT (""): 34.8 (24–45.5) g, n = 91* CrnC: 28.2 (26.3–30.0) mm, n = 14 GWS: 15.3 (14.0–16.3) mm, n = 17 C–M3: 10.4 (9.9–10.8) mm, n = 12 Gambia, Guinea-Bissau, Liberia, Côte d’Ivoire, Ghana, Togo, Benin, Cameroon, DR Congo, Uganda, Burundi, Tanzania (FC, FMNH, IICT/CZ, MZUF, RMCA, ROM, SMF, SMNS, USNM) *Non-pregnant "". Pregnant "" up to 58 g Key References Decher & Fahr 2005a; Eisentraut 1956; Hill 1963; Lang & Chapin 1917b; Verschuren 1957.

Measurements Hipposideros cyclops FA (!!): 65.4 (61–75) mm, n = 54 FA (""): 68.0 (59–74) mm, n = 47 WS (c): 400 (374–425) mm, n = 13

Jakob Fahr

Hipposideros fuliginosus SOOTY LEAF-NOSED BAT (TEMMINCK’S LEAF-NOSED BAT) Fr. Phyllorhine fuligineuse; Ger. Temmincks Rundblattnase Hipposideros fuliginosus (Temminck, 1853). Esquisses Zool. sur la Côte de Guiné, p. 77. Ashanti Land, Ghana (type locality ‘Côte de Guiné’ restricted by Jentink 1887, 1888).

Taxonomy Originally Phyllorrhina fuliginosa. Species-group: bicolor. Synonyms: currently none. Subspecies: currently none recognized (but see Geographic Variation). Often confused with Hipposideros ruber, H. caffer, H. lamottei and H. abae. Andersen (1906) recognized the confused taxonomy of this bat and pointed out some of the specific characters for a diagnosis of the species. The analyses of Koopman (1989) and Koopman et al. (1995) again confused the situation by focusing entirely on measurements of the skull while not considering external measurements, proportions and characters. The following account is therefore largely based on specimens examined by the author, disregarding most of the published records because of the difficulties in identifying this species. Chromosome number: not known. Description Small to medium-sized microbat with noseleaf (posterior component roughly elliptical); dark brown with orangephase; ears separated; noseleaf with two lateral leaflets on each side, no club-shaped processes and nostrils not concealed by internarial septum; no frontal sac; thumb well developed with comparatively large claw and basal pad. Sexes similar. Pelage slightly coarser than in H. caffer and H. ruber. Dorsal pelage (grey-phase) dark brown. Ventral pelage similar but paler. Ears separated; comparatively and relatively short (30–31% of FA), broad, triangular and pointed with slight concavity in outer margin just below tip: 11 internal folds. Antitragus with slight fold. Posterior component of noseleaf not divided into cells by vertical septa; upper margin with low-arched outline. Behind the upper margin, there is a low to well-developed transverse supplementary structure, which is sometimes smooth

(Hill 1963) and sometimes more or less serrated (Koopman et al. 1995); the variability renders this character of little value as a means of distinguishing H. fuliginosus from H. ruber and H. caffer in which the structure is serrated (Koopman et al. 1995). No club-shaped processes; internarial septum not concealing nostrils; two lateral leaflets on each side. Frontal sac absent in both sexes (but sometimes there is a patch of bare skin). No anal sac. Wings and interfemoral membrane blackish-brown. Thumb and claw comparatively long and powerful, claw length >3 mm, claw height >1.2 mm; pad at base of thumb well developed. Fifth metacarpal 84–92% of third metacarpal. Third finger with comparatively long phalanges (first phalanx 14.3– 18.7 mm, 28–31% of FA; second phalanx 17.8–21.9 mm, 33–37% of FA) (cf. H. lamottei). Tibia comparatively short, and 38–39% of FA (cf. H. lamottei). Tail comparatively short, and 45–49% of HB. Skull robust; zygomatic arches slender; zygomatic width > mastoid width. CrnC relatively long (36.3 [35–40]% of FA, n = 83) (cf. H. lamottei). Sagittal crest prominent in specimens from the eastern population, much less so in specimens from the western population (see below). Cochleae not enlarged, their breadth only a little greater than their distance apart. Upper incisor slightly bicuspid. Upper canine of medium relative length (48 [44–51]% of C–M3, n = 3). Anterior upper premolar small, somewhat displaced labially; canine and posterior premolar almost in contact. Anterior lower premolar ca. half the height and length of the posterior premolar. Geographic Variation Although no subspecies have been described, there are two morphometrically distinct populations – 383

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a western population in Guinea to Cameroon and Gabon, and an eastern population known from only a few specimens from DR Congo, Central African Republic and Uganda. Eastern specimens are much heavier, and larger in both body and craniodental measurements (see Measurements) and have proportionally broader zygomata. They probably represent an undescribed taxon closely related to H. fuliginosus. Similar Species Four other African Hipposideros have the following combination of characters: ears separated; noseleaf with two lateral leaflets on each side, no club-shaped processes and nostrils not concealed by internarial septum (Table 15, p. 370): Hipposideros ruber. Smaller on average in all measurements (FA: 51.1 [47–55] mm; CrnC: 19.1 [17.8–20.3] mm). Thumb, claw and basal pad smaller (claw length 50 cm; stall-and-twist turns possible but rare; able to fly across a 1×1×1 m enclosure, but a complete circuit was observed only once (4 bats, 10 flights each, M. Happold unpubl.). Although able to take off from ground, prefers to dive to gain initial speed for flight and roost sites are selected accordingly. Whirring sound made during flight – perhaps caused by vibration of radio-metacarpal pouch (Lang & Chapin 1917b). By day, clings to tree trunks, rockfaces and exterior walls of buildings, two or more metres above ground, where overhanging branches, rocks or eaves create shade. Often roosts on or near buildings occupied by humans; surprisingly tolerant of humans provided they ignore the bats and do not come too close. Holds on with hindfeet and thumbs; head facing downwards but raised, chest seldom in contact; grizzled pelage provides camouflage on some natural surfaces. Reported to roost with Coleura afra in coastal caves (Kingdon 1974) but this observation almost certainly refers to T. hildegardeae (see profile); apparently there are no other published records of cave-roosting in T. mauritianus. Individuals return to the same roosts each day, and these sites become stained (? urine and/or glandular secretions). In wet weather, they move temporarily leeward and, if possible, shelter under branches, leaves or eaves. In Malawi, does not become torpid (even at 21 °C); instead, remains vigilant using the eyes and seldom echolocating. If large birds fly overhead, or if humans come too close, moves out of sight by scuttling sideways around corners or under branches and eaves at great speed. Only flies to nearby sites if danger comes very close.The importance of constant daytime vigilance to avoid predators, and the concomitant need to avoid torpor, probably limits this species to warm environments. Three captive bats, eating winged termites and seldom flying, did not drink (Happold & Happold 1988). Kidneys well adapted for conservation of water; predicted mean maximum urine concentration is comparatively high (3921 mOsmol/kg) (Happold & Happold 1988). Perceives surroundings during day mainly visually (even in flight), but echolocates at night. Foraging and Food  Forages mainly by fast-hawking in open spaces (Fenton et al. 1980, M. Happold unpubl.); wing morphology

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Taphozous mauritianus.

and echolocation are primarily adapted for this. However, also forages by chasing insects over walls (Grubb et al. 1998), and by taking off and catching butterflies passing their roosts during the day (D. Rushworth in Smithers 1983). Fast-hawking occurs above treetops, between well-spaced trees, over clearings, along cliff-faces and over rivers (5–40 m above the water) (Smithers 1971, Fenton et al. 1980, M. Happold unpubl.). Sometimes forages up to 550 m above ground (Fenton & Griffin 1997). Kingdon (1974) reports three hours of intensive activity beginning after dark, followed by long rests (at favoured night roosts) interspersed with short flights. However, foraging sometimes begins at sunset (M. Happold unpubl.). Foraging is characterized by steep dives, up to 10 m, assumed to be pursuits of moths diving evasively in response to the bat’s echolocation calls (Fenton et al. 1980). Feeds mainly on moths (based on stomach contents examined by Lang & Chapin 1917b), but captive bats also eat beetles, winged termites, flies and many other insects, suggesting that opportunistic feeding would occur if moths were comparatively scarce. Echolocation  In Zimbabwe, one foraging bat emitted multiharmonic CF search-phase calls (second harmonic ca. 25 kHz) followed by shorter, multiharmonic shallow linear FM approachphase calls and then even shorter, angular shallow/steep FM terminal-phase calls. There were four harmonics – the second was dominant but the first and third contained appreciable energy, especially in the approach- and terminal-phases, and therefore these calls are audible to humans (Fenton et al. 1980). Search-phase callshape (Malawi): typically short, multiharmonic CF or very shallow linear FM; intensity high; bandwidth 0–2 kHz; start-frequency 26–29 kHz (second harmonic); end-frequency 24–27 kHz; callduration 5–13 ms (five bats foraging or flying in open, 50 calls; M. Happold unpubl.) (Figure 89). While orientating in clutter (near ground, trees, buildings), the calls are steep quasi-linear FM sweeps falling from ca. 30 to 24 kHz (second harmonic); mean duration 2.3–3.9 ms; call repetition-rate 10–27 calls/sec. As bats gain height, calls become narrower in bandwidth, longer in duration and sequences may contain some angular shallow/steep FM sweeps (M. Happold unpubl.). See Taylor (1999a) for data from South Africa and Swaziland.

Social and Reproductive Behaviour  Roosts singly (? adult ?? only) or in groups typically of 2–6 (but up to 12) of mixed composition, apparently including several adult ?? and // (with or without unweaned juveniles or subadults), but group composition and distance between neighbouring groups need further investigation. Group-members (except // with young) roost at least 10 cm apart, and up to several metres apart. They make brief contacts, including climbing-over, but then immediately move apart. Vocalizations include (a) single loud ‘ping’ emitted in contexts of threat and/or alarm to repel conspecifics that come too close, (b) a three-syllable call emitted at 2–3 second intervals when a groupmember returns to the roost area, (c) ‘twittering’ emitted by mothers and young (sustained if they are kept apart) and (d) several other vocalizations of unknown meaning (Happold et al. 1987). Individuals sometimes pursue each other in flight, and fight on the roosts, and they screech in these contexts (Lang & Chapin 1917b). Mating system not known, but fragmentary evidence suggests territoriality with defence of foraging areas, day-roosts and perhaps //, but the possibility that this indicates resource-defence polygyny and/ or female-defence polygyny needs confirmation. Resource-defence polygyny is exemplified by the South American emballonurid Saccopteryx bilineata (Bradbury & Vehrencamp 1977). Female T. mauritianus fly with their young attached to their underparts until the young are volant (D. Rushworth in Smithers 1971). Juveniles often roost on their mothers’ backs. Reproduction and Population Structure  Litter-size: one. Reproductive chronology probably bimodal polyoestry throughout geographic range. At ca. 4° N (DR Congo), limited data (Lang & Chapin 1917b, Verschuren 1957) suggest bimodal polyoestry with births ca. Nov–Dec and ca. Apr–May (perhaps less synchronized than in Kenya and Malawi). No conclusive data for more northern latitudes. At ca. 02° 18' S (Masalani, near Kibwezi, Kenya), births occur in Nov (peak of main wet season) and Mar–Apr (little wet season) (O’Shea & Vaughan 1980): polyoestry probable but not confirmed at this locality. At ca. 15° S (Liwonde N. P., Malawi), // bimodally polyoestrous with births in early wet season (Nov–Dec) and end of wet season (Mar–Apr) (Happold & Happold 1990a). At ca. 30° S (Durban area, South Africa), // are polyoestrous with births in Oct–Dec and Feb or Mar (F. Mackenzie in Taylor, P. 1998). Happold & Happold (1990a) did not state that T. mauritianus may be monoestrous in some regions of Africa (as erroneously reported by Dengis 1996); monoestry has not been confirmed in this species. Predators, Parasites and Diseases  Predators include owls (Demeter 1981, Taylor, P. 1998). Roosting behaviour implies danger of predation by hawks, other raptors, snakes and perhaps genets Genetta spp. Ectoparasites include bat-flies Basilia blainvillii, Phthiridium integrum (Diptera: Nycteribiidae) and a mite Olabidocarpus taphozous (Acari: Chirodiscidae) (Aellen 1952, Anciaux de Faveaux 1984). Conservation  IUCN Category: Least Concern. Measurements Taphozous mauritianus FA: 61.4 (58–65) mm, n = 103 WS (a): 420.8 (409–443) mm, n = 5* 433

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C–M3: 8.8 (7.7–9.4) mm, n = 25 Throughout geographic range except West Africa (SMNS, ZFMK and literature) *Malawi only

TL: 105 (91–116) mm, n = 55 T: 22.5 (15–28) mm, n = 55 E: 17.6 (13–22) mm, n = 63 Tr: 6.1 (5–7) mm, n = 7 Tib: 24.8 (19–27) mm, n = 19 HF: 13.4 (11–18) mm, n = 93 WT: 26.4 (20–36) g, n = 39 GLS: 21.0 (19.5–22.5) mm, n = 27 GWS: 12.9 (12.2–13.4) mm, n = 19

Key References  Dengis 1996; Fenton et al. 1980; Smithers, 1971, 1983. Meredith Happold

Taphozous nudiventris  Naked-rumped Tomb Bat Fr. Taphien à ventre nu; Ger. Nacktbauch-Grabfledermaus Taphozous nudiventris Cretzschmar, 1830. In: Rüppell, Atlas Reise Nordl. Afr., Zool. Säugeth., p. 70. Giza, Egypt.

Taxonomy  Subgenus Liponycteris. Synonyms (Africa only): assabensis, possibly serratus. Subspecies: five; one in Africa. Chromosome number (Egypt): 2n = 42; aFN = 64 (Hood & Baker 1986). Description  Medium-sized microbat without noseleaf and with terminal portion of tail projecting freely from middle of dorsal surface of interfemoral membrane; two lower incisors on each side; rump, lower belly and hindlimbs naked; dorsal pelage greyishbrown or dark brown; ventral pelage brown; wings dark brown; condylocanine length 23.0–25.2 mm (cf. Taphozous hamiltoni). Sexes similar in colour; ?? on average slightly larger than //. Pelage sleek; mid-dorsal hairs 6–7 mm. Dorsal pelage uniformly sepia brown, dark rusty-brown or ashy greyish-brown (not grizzled); hairs with basal half cream. Rump and flanks naked with clear demarcation between furred and naked areas; up to one-third of the dorsal surface of the body is naked. Ventral pelage paler than dorsal pelage; posterior third of ventral surface of body is naked. Gular pouch well developed in ??, less so in //. No black beard. Head moderately flat, subtriangular (viewed dorsally) with long conical muzzle and very shallow depression between the eyes. Lower lip with conspicuous grooved prominence. Eyes comparatively large. Ears subtriangular, backward-pointing, with papillae along the lower inner margin. Tragus axe-head-shaped with pronounced lobule at base of posterior margin (Figure 85e); antitragus large, almost reaching corner of mouth. Wings and interfemoral membrane very dark brown; radio-metacarpal pouch present. Hindlimbs naked. Skull (Figure 86b) medium-large for an African emballonurid, very broad and heavily built. Frontal depression shallow. Dorsal profile of skull (viewed laterally) almost smoothly convex; profile of forehead region very weakly concave (almost straight). Postorbital processes long, slender. Sagittal crest low; occipital helmet well developed. Anterior palatal emargination wide and U-shaped (Figure 87c). Tympanic bulla with inner face incomplete. Condylocanine length: 23.0–25.2 mm. Two lower incisors on each side.

Taphozous hamiltoni. Naked area usually less extensive. Gular pouch of // well developed. Skull with occipital helmet poorly developed; condylocanine length: 20–22 mm. Saccolaimus peli. Much larger (FA: 87–95 mm; GLS: 28.0–31.6 mm). Distribution  In Africa, recorded from the Sahara Arid, Sudan Savanna, Guinea Savanna and Somalia–Masai Bushland BZs. Distributed from Morocco, Mauritania and Senegal, to Egypt, Eritrea, Djibouti and Somalia, with a narrow southward extension through S Sudan, NE DR Congo and Kenya to Tanzania. Recorded from Guinea-Bissau by Seabra (1900) and Veiga-Ferreira (1949): however, Seabra’s identification has not been confirmed and VeigaFerreira’s record refers to T. perforatus (Lopes & Crawford-Cabral 1990). Its occurrence in Gambia is unproven although plausible (Grubb et al. 1998). The African distribution appears very disjunct: there are some clusters of localities and many isolated localities. Not known to what extent this reflects insufficient sampling and/or the isolated nature of suitable habitats. Extralimitally: eastwards across

Geographic Variation  None recorded in Africa, where only the nominate subspecies is recognized (Koopman 1994). Similar Species  Only two other African emballonurids have a band of naked skin on the posterior of the rump adjacent to the interfemoral membrane:

Taphozous nudiventris

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Arabian Peninsula, Iran, Afghanistan, Pakistan and India to Myanmar. Mapped from country checklists (see order Chiroptera), Kock 1969a, other literature and museum records. Habitat  Woodland savannas including undifferentiated woodland, Isoberlinia woodland, Acacia–Commiphora bushland and thicket, and more arid habitats including semi-desert grasslands and shrublands, and deserts. Very little known about habitat requirements, but the distribution is probably restricted by the availability of both flying insects and day-roosts (see below): possibly this species was more abundant and widespread when northern Africa was less arid. Abundance  No information. Adaptations  Aspect ratio high; wing-loading extremely high; wing-tip pointed; flies very fast with great agility but poor manoeuvrability (Rosevear 1965, Norberg & Rayner 1987). Roosts by day in crevices and narrow fissures between rocks or stone blocks, in caves, inselbergs, sandstone hills, wells, old ruins, mosques and the ancient Egyptian temples and tombs of the NileValley, including Karnak (Hoogstraal 1962, Gaisler et al. 1972, Qumsiyeh 1985, Happold 1987). Day-roosts may have an unpleasant, pungent smell (to humans). Tolerates quite bright light in roosts. April temperatures recorded near roosting bats in Egypt ranged from 23 °C in a mosque to 35 °C in an occupied crevice at Karnak (cf. 37 °C outside in shade) (Gaisler et al. 1972). Rhinopoma hardwickii, Asellia tridens, Nycteris thebaica and Rousettus aegyptiacus also roosted in the temple complex at Karnak, but each species in a different place. In Iraq, spends summer in cool buildings and caves: moves to old buildings with wooden or rush roofs for hibernation during winter (Al-Robaae 1968). Accumulates fat prior to hibernation. In Pakistan, makes seasonal migrations between summer and winter day-roosts, accumulates fat towards end of monsoon and spends winter in torpor (Roberts 1977). In India and East Africa, fat also accumulates seasonally despite lack of hibernation there (Kingdon 1974). No further information for Africa, where this species has received little attention. Foraging and Food  Forages by fast-hawking in open spaces. Foraging more than 100 m above ground has been inferred from erratic flight paths by T. nudiventris in India (Siefer & Kringer 1991 in Fenton & Griffin 1997). Prey includes beetles, moths, grasshoppers, cockroaches, crickets and winged-ants (Pearch et al. 1999). Becomes particularly numerous in Gharbiya Province, Palestine during Jul– Aug when Cotton Leaf-worm moths invade cotton fields, and large quantities of moth scales are found in the bats’ stomachs at this time (Madkour 1977 in Qumsiyeh 1985). Echolocation  No information for this species in Africa. Social and Reproductive Behaviour  Roosts gregariously; becomes very active ca. 30 minutes before sunset and emerges from the roost ca. 15 minutes after sunset. One colony in Egypt comprised ca. 50 individuals (Gaisler et al. 1972); 2000 recorded in Pakistan (Roberts 1977). In Iraq, colonies are of mixed composition throughout most of year.When // approach time of parturition, ?? leave or are driven away (sometimes to roosts 100–300 m away) where they are found in groups of 5–10.The ?? rejoin the // after the young become volant

(Al-Robaae 1968). Maternity colonies contain 200–1000 //. Large nuclear colonies with small colonies nearby have also been recorded in Pakistan (Roberts 1977). According to Al-Robaae (1968), the young bat clings to its mother’s back for two weeks, then roosts beside her and remains with other young while she forages. According to Roberts (1977), the young remains attached to a nipple for 3–4 weeks, then clings to its mother’s flank or back while she forages until eight weeks old.Young begin flying within and near the day-roost when five weeks old. Weaning occurs during sixth week. When foraging away from dayroost, young stay close to mother even after weaning. Reproduction and Population Structure  Litter-size: one. In Iraq, the reproductive chronology is restricted seasonal monoestry, with sperm storage and delayed fertilization (Al-Robaae 1968); no conclusive data for Africa. In Iraq, copulation occurs in Sep–Oct shortly before hibernation. Sperm are stored in the / until late Mar when hibernation ends and ovulation and fertilization take place.Young are born nine weeks later. All young are born during a ten-day period in late May: they are blind, naked, FA: 25–30 mm, HB: 45–50 mm. Eyes open after one week. Growth rapid. Insects in stomach by seven weeks. Reproductive chronology in Pakistan is also restricted seasonal monoestry with copulation in Sep, emergence from winter-roosts at beginning of Mar, birth of young in mid-Apr (Roberts 1977). Predators, Parasites and Diseases  Predators include Barn Owls Tyto alba in Palestine (Dor 1947 in Qumsiyeh 1985) and hawks that capture the bats as they leave their roosts in Pakistan (Roberts 1977). Ectoparasites in Africa include bed-bugs Leptocimex vespertilionis, L. duplicatus, Stricticimex puylaerti (Hemiptera: Cimicidae); a flea Xenopsylla conformis (Siphonaptera: Ischnopsyllidae); ticks Carios vespertilionis, C. boueti, C. confusus (Acari: Argasidae); and a mite Steatonyssus sudanensis (Acari: Macronyssidae) (Anciaux de Faveaux 1984). Conservation  IUCN Category: Least Concern. Measurements Taphozous nudiventris FA: 71.8 (67–79) mm, n = 103 WS: n. d. TL: 120.0 (110–132) mm, n = 37 T: 31.0 (20–37) mm, n = 64 E: 20.4 (16–25) mm, n = 62 Tr: 6.6 (6–7) mm, n = 26 Tib: 29.1 (27–31) mm, n = 24 HF: 16.3 (11–18) mm, n = 50 WT: n. d. GLS: 25.7 (23.2–28.5) mm, n = 69 GWS: 15.5 (14.5–16.6) mm, n = 72 C–M3: 10.9 (10.1–11.6) mm, n = 72 Burkina, Mali, Egypt, Sudan, Ethiopia (BMNH, SMNS, ZFMK and literature) Key References  Al-Robaae 1968; Gaisler et al. 1972; Kock 1969a; Qumsiyeh 1985. Meredith Happold 435

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Taphozous perforatus  Egyptian Tomb Bat Fr. Taphien perforé; Ger. Ägyptische Grabfledermaus Taphozous perforatus E. Geoffroy, 1818. Descrip. de L’Egypte 2: 126. Kom Ombo, between Edfu and Aswan, Upper Egypt.

Taxonomy  Subgenus Taphozous. Synonyms in Africa: haedinus, maritimus, rhodesiae, senegalensis, sudani, swirae. Extralimitally: maritimus. Subspecies: four (three in Africa); boundaries unclear. Harrison (1958, 1962) considered sudani to be a distinct species, a view not shared by more recent authors (Rosevear 1965, Kock 1969a, Hayman & Hill 1971, Koopman 1975, Simmons 2005). Chromosome number (Egypt): 2n = 42; aFN = 64 (Yaseen et al. 1994). Description  Medium-sized microbat without noseleaf and with terminal portion of tail projecting freely from middle of dorsal surface of interfemoral membrane; two lower incisors on each side; pelage covering all of body; dorsal pelage not grizzled; ventral pelage pale grey to dark greyish-brown; wings almost white to pale brown. Sexes similar. Pelage soft, fine, silky; covering all parts of body; mid-dorsal hairs 6–7 mm. Dorsal pelage uniformly dark chocolate brown, sepia brown, greyish-brown or ashy-brown; hairs white at base. Ventral pelage pale grey, pale greyish-brown to dark greyishbrown; chin and throat usually sepia brown and darker than dorsal or ventral pelage. Adult ?? and // with a poorly defined patch of longer darker hairs on the throat (perhaps not always present). No gular pouch in either sex according to Rosevear (1965), Hayman & Hill (1971) and Harrison & Bates (1991), but Rosevear mentions that a shallow fold of skin or merely a crescentic mark may be present. In contrast, Koopman (1975) implies that a gular pouch is present in some ?? (see Geographic Variation). Head moderately flat, subtriangular (viewed dorsally) with long pointed muzzle and deep depression between the eyes. Eyes comparatively large for a microbat. Lower lip with conspicuous grooved prominence. Ears subtriangular, backward-pointing, with small papillae on lower inner margin. Tragus axe-head-shaped with poorly developed lobule at base of posterior margin (Figure 85f). Wings variable: almost white in individuals with darker brown dorsal pelage, to pale brown in individuals with paler ashy-brown dorsal pelage. Radio-carpal pouch present in both sexes. Interfemoral membrane pale brownish. Skull (Figure 86) small for an African emballonurid. Frontal depression deep. Profile of forehead region of skull (viewed laterally) strongly concave. Postorbital processes long and slender. Sagittal crest absent, no occipital helmet. Anterior palatal emargination comparatively narrow, and more angular and more V-shaped than in other African Taphozous (Figure 87d). Inner face of tympanic bulla incomplete. Two lower incisors on each side.

Similar Species  Three other African emballonurids have pelage covering all of the body: Taphozous mauritianus. Dorsal pelage grizzled (salt and pepper effect); ventral pelage pure white (sometimes stained yellowish). T. hildegardeae. Dorsal pelage uniformly pale greyish-brown. Adult ?? with blackish ‘beard’ on throat, ventral pelage white stained yellowish-brown. Adult // with throat and belly pure white. Coleura afra. Three lower incisors on each side. Prominence on lower lip not divided by median groove. FA: 44–53 mm. Wings brown to blackish-brown. Ventral pelage brown. Distribution  In Africa, found along Nile Valley in the Sahara Arid BZ and in some parts of the Sahel Savanna, Sudan Savanna, Guinea Savanna, Somalia–Masai Bushland, Coastal Forest Mosaic and Zambezian Woodland BZs, and in the Northern and Eastern Rainforest–Savanna Mosaics. Not recorded from the Rainforest BZ except in NE DR Congo. Except along the Nile Valley, the distribution appears very disjunct (possibly because of insufficient sampling and/or the absence of suitable day-roosts). In West Africa, recorded from Mauritania, Senegal, Mali, Ghana and E Burkina to NW Nigeria. On eastern side of Africa, recorded contiguously from Egypt, Sudan, NE DR Congo, Uganda, NW Kenya and the Ethiopian Rift Valley to the Red Sea in Djibouti and NW Somalia, and there are apparently isolated populations in E Sudan, the Ethiopian Highlands, C to SE Kenya, Tanzania, S DR Congo to N Zambia, the Okavango Swamp of N Botswana, and the low-lying area of S Zimbabwe and E Botswana. Over much of its wide distribution in Africa, records are

Geographical Variation  Four subspecies are recognized in Africa by Simmons (2005), but their boundaries (based on Koopman 1994) and diagnostic characters are not clear. T. p. senegalensis: West Africa. T. p. perforatus: Egypt and N Sudan. T. p. sudani: C and S Sudan, E DR Congo, Botswana and Zimbabwe. T. p. haedinus: Tanzania to Ethiopia and, extralimitally, across S Asia to India.

Taphozous perforatus

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extremely sparse. Extralimitally: S Arabia, Jordan, S Iran, Pakistan and NW India. Habitat  Mainly open woodland savannas, including Acacia woodland, Isoberlinia woodland, Acacia–Commiphora bushland and thicket, and miombo and mopane woodlands, where suitable dayroosts are present. Also recorded from flooded savanna in the Nile Delta, in seasonally moist habitats along the Nile Valley, in rainforest–savanna mosaic, in East African coastal forest mosaic, and in the Okavango Swamp, Botswana. There are some records within the rainforest zone in NE DR Congo, but no details are available. Near the confluence of the Shashi and Limpopo Rivers in Zimbabwe, T. perforatus was found near a range of sandstone hills with Acacia woodland on the flat country, and well-developed riverine woodland along the seasonally dry river bed (Smithers 1983). Evidently avoids forests, montane habitats, semi-deserts and deserts. Abundance  Uncertain. Over much of its wide distribution, records are extremely sparse, e.g. in East Africa, only one specimen published from Uganda (Kock 1974b), two from Tanzania (Allen & Loveridge 1933, Harrison 1961) and seven from Kenya (Aggundey & Schlitter 1984). Also, comparatively poorly represented in museum collections. In contrast, large numbers (e.g. ca. 200) have been reported from day-roosts in the Nile Valley in Egypt (Gaisler et al. 1972, Qumsiyeh 1985) and Hoogstraal (1962) reports it as one of the most common bats in Egypt. Also, a colony of 150–200 was found in NE DR Congo (Lang & Chapin 1917b) and one of >100 in S DR Congo (Anciaux de Faveaux 1978). Adaptations  Aspect ratio medium; wing-loading mediumhigh. Based on wing morphology, flight-speed estimated to be ca. 8.0 m/sec (Rydell & Yalden 1997), which is medium for bats. By day, roosts in caves, crevices in rocky outcrops and old buildings, including Egyptian pyramids, tombs and other ancient monuments, gaining access by scuttling and climbing as well as by flying. Unlike T. mauritianus which roosts in the open, T. perforatus roosts tucked away in darkened narrow crevices in rocks or brickwork. Sometimes roosts quite close to the ground (Hoogstraal 1962). Foraging and Food  Forages by fast-hawking. Based on analysis of faeces (Rydell & Yalden 1997), the diet comprises predominantly, in order of priority, moths (56% by volume), termites (14%), beetles (10%) and, to a lesser extent, crickets and katydids (8%), bugs (3%), lacewings (2%), ants (1%) and flies (1%). Echolocation  No information. Social and Reproductive Behaviour  Unlike the less gre­garious T. mauritianus, T. perforatus roosts in groups of several individuals to at least 200 (see Abundance), with individuals huddled together in dense associations (Lang & Chapin 1917b, Hoogstraal 1962). Groupmembers scuttle and crawl about, but Lang & Chapin (1917b) noted that they never crawled over each other.

Reproduction and Population Structure  Litter-size (Egypt, Nigeria, Zimbabwe): one (n = 19). Reproductive chronology uncertain. At 25° 41' N (Luxor, Egypt), 16 of 16 // were in advanced pregnancy in Apr (no data for other months) (Gaisler et al. 1972). At 13° 04' N (Sokoto, N Nigeria), 6 of 6 adult // were lactating in mid-Jun (wet season) and, of these, one was examined histologically and found to be in early pregnancy (no data for other months) (Harrison 1958). This suggests that the chronology in N Nigeria is seasonal polyoestry with postpartum oestrus, but the number of litters/year, the timing of other births and the proportion of // that have more than one litter/year, are not known. In Zimbabwe, two pregnant // were reported in Nov (Smithers & Wilson 1979). The data from Nigeria and Zimbabwe indicate that at least some births occur at the beginning of the unimodal wet seasons, and that there are boreal and austral cycles. Data from elsewhere, summarized by Anciaux de Faveaux (1978), are not conclusive. Predators, Parasites and Diseases  Predators include Spotted Eagle-owls Bubo africanus (Demeter 1982), and probably Lanner Falcons Falco biarmicus, which have been observed feeding on bats emerging from a quarry where T. perforatus was known to roost (Butler 1905, D. Kock pers. comm.). Ectoparasites include fleas Xenopsylla cheopis (Siphonaptera: Pulicidae), Araeopsylla wassifi, Chiropteropsylla aegyptia, C. brockmani (Siphonaptera: Ischnopsyllidae); bat-flies Phthiridium integrum (Diptera: Nycteribiidae), Brachytarsina divsersa, B. alluaudi (Diptera: Streblidae); ticks Carios vespertilionis, C. boueti, C. confusus, Alectorobius salahi (Acari: Argasidae); and mites Steatonyssus sp. (Acari: Macronyssidae), Ugandobia barnleyi (Acari: Myobiidae), Alabidocarpus taphozous (Acari: Chirodiscidae) (Anciaux de Faveaux 1984). Dakar bat 249 virus has been isolated from T. perforatus (Anciaux de Faveaux 1984). Conservation  IUCN Category: Least Concern. Measurements Taphozous perforatus FA: 62.6 (56–67) mm, n = 129 WS: n. d. TL: 101.6 (90–112) mm, n = 46 T: 26.2 (19–32) mm, n = 92 E: 17.7 (15–21) mm, n = 94 Tr: 6.2 (4.9–7.5) mm, n = 47 Tib: 23.0 (21–24) mm, n = 18 HF: 12.4 (10–14) mm, n = 21 WT: 29.6 (20–39) g, n = 13* GLS: 19.7 (18.1–21.7) mm, n = 70 GWS: 11.7 (11.3–13.0) mm, n = 70 C–M3: 8.5 (7.8–9.0) mm, n = 13 Throughout geographic range (HZM, MNHN and literature) *Zimbabwe (Smithers & Wilson 1979) Key References  Harrison 1958, 1962; Rosevear 1965; Rydell & Yalden 1997; Taylor 2000. P. J. Taylor

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Family NYCTERIDAE

Family NYCTERIDAE Slit-faced Bats

Nycteridae Van der Hoeven, 1855. Handb. Dierkunde, 2nd edn., 2: 1028. Nycteris (13 species)

Slit-faced Bats

p. 440

All nycterids belong to the genus Nycteris. There are 16 extant species: 13 occur in Africa, one in Madagascar and two in South-East Asia (Simmons 2005). They are found mainly in rainforests and woodland savannas, but some species inhabit semi-arid habitats. Nycterids are unique in having a deep longitudinal cleft on the head (from the forehead to the nostrils), which is bordered by fleshy outgrowths whose outlines are usually obscured by the pelage (Figure 32g). Like the noseleaves on the muzzles of bats in the families Rhinolophidae, Hipposideridae, Megadermatidae and Rhino­ pomatidae, these outgrowths play a role in echolocation. Nycterids are also distinguished by a long tail that is completely enclosed by a very large interfemoral membrane and which terminates in a uniquely Y-shaped or T-shaped cartilaginous process (Figure 33g). None are considered to be pests. African nycterids are very small to medium-sized microbats with moderately long, soft, loose, fluffy pelage, usually brown or grey, occasionally orange. Males and // are similar in size and colour. The body is small and compact; the shape of the head is obscured by

pelage. The muzzle has a deep longitudinal cleft; the nostrils are located in the anterior end of this cleft, and the cleft expands into a deep pit on the forehead. The noseleaf is comprised of fleshy outgrowths along the margins of the cleft; its shape is not used diagnostically. The ears are enormous! They are longer than the length of the head, rounded and are united at the base by an inconspicuous low membrane. The tragus is well developed and its shape is an important diagnostic character. The eyes are minute. The wings are large and broad; the second finger has a long metacarpal but no phalanges. The hindlimbs are relatively long with small soles, short toes and small claws; the toes (except hallux) have three phalanges. The tail is relatively long (44–52% of TL), and is completely enclosed by a very large interfemoral membrane, which is supported and manipulated by long calcars and the Y-shaped or T-shaped terminal process of the tail. There are no pubic nipples. The interfemoral membrane is reinforced by numerous strands of collagen fibres, which form patterns of broken and/or unbroken but dotted transverse lines across the membrane. The skull is long and narrow, with a deeply concave frontal shield in the interorbital region; the shield is formed by the fusion of the postorbital processes

Figure 92. Characters of bats in the family Nycteridae. Flight membranes and bones of wing, hindlimb and tail (e.g. Nycteris thebaica). Skull (e.g. Nycteris grandis (RMCA 80-36-M-32).

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with broad supraorbital ridges. Selected external and craniodental characters of representatives of the genus are illustrated in Figure 92 but there are important inter-specific variations in dentition (see below). Posteriorly, the frontal shield rises to the most elevated part of the skull where it merges with the sagittal crest. The cleft or ‘slit’ in the face leads into a large chamber supported by the frontal shield. The sagittal crest is usually low or absent (but well developed in two species) and there are no lambdoid crests. The nasal branches of the premaxillae are lost; the palatal branches are partly developed and not fused to each other but are lightly fused to the maxillae and other bones of the palate; they may fill, or only partly fill, the space between the maxillae. Skull characters are not as useful as external characters and dental characters for distinguishing between species of Nycteris, and therefore skulls are not described in the species profiles. The dental formula is 2113/3123 = 32. The karyotypes of nycterids are characterized by high fundamental numbers (aFN) compared to other families of bats (Lee et al. 1989). The five species of African nycterids for which data are available have very low aspect ratios and low to very low wing-loadings (very low in all but one species). They can fly slowly with great manoeuvrability, hover, take off from the ground (even when carrying a young bat or heavy prey) and they can turn around in very confined spaces. These adaptations enable nycterids to forage close to the ground and in dense vegetation. Because of their wing morphology, flight is energetically expensive, so they spend minimal time in flight. Although they sometimes forage by slow-hawking, they are usually fly-catchers and perch-hunting gleaners and often carry larger prey back to the perch before eating it. Most species are insectivorous, feeding on a wide variety of taxa, but one African species (Nycteris grandis) also eats small vertebrates. Prey is detected from sounds made by the prey as well as by echolocation. Nycterids are referred to as ‘whispering’ bats because they emit multiharmonic CF echolocation calls of short duration and low intensity (Griffin 1958, Neuweiler 1989). These calls are particularly suited for densely cluttered environments. Nycterids can hover and glean prey from vegetation and other surfaces and, because they can take off from the ground, they can land to attack non-volant prey. However, they are unable to scuttle over the ground and they cannot climb. During the day, nycterids roost in hollow trees, caves, mines, buildings, road culverts, hollow logs, abandoned burrows and similar semi-dark places. They hang freely from footholds on horizontal ceilings, and maintain a space between adult individuals. Their exceptional manoeuvrability enables them to fly into confined spaces to find roosts inaccessible to other bats. They perform an aerobatic somersault in mid-air in order to gain a holdfast from which to hang. Nycterids do not hibernate. The social behaviour and mating systems of nycterids are not well known. Bats of one species appear to be mainly solitary; some apparently form monogamous family groups (Hill & Smith 1984), and it seems likely, from the very limited data, that some might establish harems (but this needs confirmation). All nycterids for which data are available are monotocous. Surprisingly, the reproductive chronology of only one African species (N. thebaica) has been ascertained with

certainty, and only in South Africa where it exemplifies restricted seasonal monoestry. Polyoestry has been reported for two species (N. hispida and N. macrotis) in Tanzania, but the chronologies of these species are not known in detail. Bernard (1982a) and Bernard & Cumming (1997) suggest that polyoestry is the ancestral strategy for nycterids and that monoestry evolved as an adaptation to higher latitudes where conditions are optimal only once per year. The geological range of the family Nycteridae is not known (Koopman 1984). It has been suggested that there have been at least two major dispersals of the genus in the Old World tropics (Thomas et al. 1994). First, the arge group (see below) dispersed from Africa, during a Miocene or Pliocene pluvial period, across forests that spread uninterrupted from Africa to eastern Asia and later, when drier conditions prevailed, two South-East Asian species became isolated from the remainder in Africa. Secondly, N. thebaica, which is adapted to xeric conditions, dispersed from Africa into Israel and Arabia. In Africa, adaptive radiation has been extensive: three species are found mainly in rainforest, six occur in both forests (including rainforest) and woodland savannas, three occur mainly in woodland savannas, and one (N. thebaica) occurs in diverse habitats including clearings in rainforest, woodland and grassland savannas, and oases and riverine habitats in arid regions. Four species (31%) have been found in montane habitats, but no species are mountain specialists. It is not uncommon to find more than one species at any one place, and 4–5 species have been recorded at several localities. For example: Comoé N. P., Côte d’Ivoire (Sudan Savanna BZ): N. arge, N. intermedia, N. hispida, N. macrotis, N. nana (Fahr & Kalko 2010). Ituri Forest, E DR Congo (Rainforest BZ): N. arge, N. grandis, N. hispida, N. intermedia, N. nana (Van Cakenberghe & De Vree 1985, 1993). Taï N. P., Côte d’Ivoire (Rainforest BZ): N. arge, N. grandis, N. intermedia, N. major, (Fahr & Kalko 2010). Adiopodoumé, Côte d’Ivoire (Rainforest BZ): N. arge, N. hispida, N. intermedia, N. macrotis (Van Cakenberghe & De Vree 1985, 1993). Avakubi, NE DR Congo (Rainforest BZ): N. arge, N. hispida, N. intermedia, N. major (Van Cakenberghe & De Vree 1985, 1993). Liwonde N. P., Malawi (Zambezian Woodland BZ): N. grandis, N. hispida, N. macrotis, N. thebaica (Happold & Happold 1997). Based on morphological data, the family Nycteridae was placed in the superfamily Rhinolophoidea together with the families Rhinolophidae, Hipposideridae and Megadermatidae (Simmons 1998, Simmons & Geisler 1998), but more recent molecular studies have contradicted many groupings based on morphological data and, pending resolution of the controversies, no chiropteran superfamilies are recognized by Simmons (2005). The systematics of African nycterids has been revised by Van Cakenberghe & De Vree (1985, 1993, 1998). All nycterids belong to the genus Nycteris. Meredith Happold

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GENUS Nycteris Slit-faced Bats Nycteris G. Cuvier & E. Geoffroy, 1795. Mag. Encyclop. 2: 186. Type species: Vespertilio hispidus Schreber, 1774 (nomen nudum, validated by Opinion 111 of the International Commission on Zoological Nomenclature).

a

b c

d

f

e

g

h

i

Nycteris thebaica.

This genus has 16 extant species of which 13 occur in Africa. There are no other genera in the family Nycteridae and the characters of the genus Nycteris are given in the family profile. African species of Nycteris can be distinguished by the combinations of characters given in Table 18, but see also additional information under Similar Species. Characters of particular diagnostic value are: Shape of upper incisors. Bicuspid (Figure 93a) or tricuspid (Figure 93b). Relative size of the posterior lower premolar. Can be large (reaching at least half the height of the anterior lower premolar Figure 93c) or small (reaching less than half height of anterior lower premolar (Figure 93d and e). Shape of posterior margin of the tragus in conjunction with the shape of the tragus. The posterior margin can be smoothly convex and the shape semi-lunate (Figure 93f) or smoothly convex and the shape narrow (not semi-lunate) (Figure 93g); or the posterior margin can be deeply notched and the tragus inverted pear-shaped (Figure 93h), or the posterior margin can be concave or slightly concave (Figure 93i). FA (length of forearm). Mean ± 1 S.D. is given for some species. E (length of ear). In African species, mean length ranges from 21.1– 32.0 mm, so 21.1–24.7 mm is comparatively short, 24.8–28.3 mm is medium, and 28.4–32.0 mm is comparatively long.

Figure 93. Diagnostic characters of species of Nycteris. (a) Upper incisors bifid (e.g. N. capensis). (b) Upper incisors trifid (e.g. N. hispida). (c) Posterior lower premolar relatively large (e.g. N. intermedia). (d) and (e) Posterior lower premolar relatively small (e.g. N. macrotis, N. thebaica). (f) Tragus broad with posterior margin smoothly convex; shape semi-lunate (e.g. N. macrotis). (g) Tragus narrow with posterior margin smoothly convex; shape not semi-lunate (e.g. N. hispida). (h) Tragus inverted pear-shaped with posterior margin deeply notched (e.g. N. thebaica). (i) Tragus with posterior margin concave or slightly concave (with ‘kink’ at centre) (e.g. N. arge; after Rosevear 1965). Left tragus illustrated; all drawn to same length.

Relative length of ear. In African species, ears range in relative length from 52–80% of FA, so 52–61% of FA is relatively short, 62–71% of FA is medium, and 72–80% of FA is relatively long. Tib (length of tibia). In African species, mean length ranges from 15–30 mm, so 15–20 mm is comparatively short, 21–25 mm is medium, and 26–30 mm is comparatively long. Relative length of tibia. In African species, mean relative length ranges from 44–56% of FA, so 44–48% is relatively short, 49–52% is medium and 53–56% is relatively long. Shape of the baculum. The tip can be simple (Figure 94a–d), ventrally hooked (Figure 94e–h) or trifid (Figure 94i–k); for minor intraspecific variations, see Thomas et al. 1994.

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Family NYCTERIDAE

1 mm

a

b

c

d

e

f

g

h

i

j

k

Figure 94. Bacular morphology in Nycteris. Bacula tips simple: (a) N. arge, (b) N. nana, (c) N. gambiensis (specimen 1) and (d) N. gambiensis (specimen 2). Bacula tips ventrally hooked: (e) N. hispida (specimen 1), (f) N. hispida (specimen 2), (g) N. hispida (specimen 3) and (h) N. grandis. Bacula trifid-tipped: (i) N. macrotis (specimen 1), (j) N. macrotis (specimen 2) and (k) N. macrotis (specimen 3). For each species, left illustration is the dorsal view with tip uppermost; right illustration is the lateral view of the right side with tip uppermost. Bacula for up to three different specimens are included to indicate intraspecific variation. All bacula traced from Thomas et al. (1994).

Table 18.  Key to the African species in the genus Nycteris. Forearm lengths for N. arge, N. intermedia, N. major and N. nana from J. Fahr (see profiles), all others from Van Cakenberghe & De Vree (1985, 1993, 1998). Upper incisors

Posterior lower premolar

Tricuspid

Small

Tricuspid

Small

Tricuspid

Small

Posterior margin of tragus (Tragus shape)

FA mean ± 1 S.D. (mm)

E (mm)

Tib (mm)

Smoothly convex Narrow Smoothly convex Narrow Smoothly convex Narrow

58.25 ± 2.6 (52–65) 38.72 ± 1.65 (34–44) 41.63 ± 1.72 (37–45)

31.6 (24–35) 22.3 (16–24) 27.8 (25–31)

30.1 (25–35) 18.1 (14–24) 20.3 (18–22)

Bicuspid

Small

Smoothly convex (Semi-lunate)

40.0 (38–41)

21.8 (20–23)

20.7 (18–23)

Bicuspid

Small

Smoothly convex (Very narrowly semi-lunate)

39.26 ± 1.91 (35–42)

31.7 (28–35)

19.5 (16–22)

Bicuspid

Small

Bicuspid

Small

Bicuspid

Small

48.19 ± 2.03 (40–55) 40.15 ± 1.44 (35–44) 44.52 ± 2.32 (34–52)

31.4 (27–35) 27.3 (24–30) 31.7 (26–37)

22.7 (18–26) 19.9 (17–22) 22.0 (17–27)

Bicuspid

Small

51

22

25

Bicuspid

Large

Concave with ‘kink’

34.4 ± 1.7 (32–37)

21.1 (19–23)

15.3 (14–17)

Bicuspid

Large

Concave with ‘kink’

36.8 ± 0.5 (36–38)

22.7 (20–24)

19.8 (18–21)

Bicuspid

Large

Slightly concave with ill-defined ‘kink’

42.1 ± 1.0 (39–46)

27.9 (25–34)

22.0 (19–26)

Bicuspid

Large

Concave with ‘kink’

47.5 ± 1.0 (45–49)

29.2 (27–31)

26.3 (24–28)

Smoothly convex (Semi-lunate) Deeply notched (Pear-shaped) Deeply notched (Pear-shaped) ? Deeply notched (? Pear-shaped)

Miscellaneous

Species

Baculum tip ventrally hooked

N. grandis

Baculum tip ventrally hooked

N. hispida

Baculum: no data

N. aurita

Baculum tip trifid Ventral pelage dark grey (few specimens) Ears 54.4 (51–56)% of FA (n = 3) Cameroon, Ethiopia, Somalia Baculum tip simple Ventral pelage not dark grey Ears 79.5 (74–89)% of FA Tanzania, Zambia, Malawi, Mozambique Baculum tip trifid Ears 65.0 (54–80)% of FA Baculum tip simple W Africa and Cameroon Baculum tip simple Widespread S of Sahara Baculum: no data As yet, only Mozambique Baculum tip simple Tibia 44 (40–47)% of FA Concavity of tragus well defined Baculum: no data Tibia 54 (50–58)% of FA Concavity of tragus well defined Baculum tip simple Tibia 52 (48–58)% of FA Concavity of tragus poorly defined Baculum: no data Tibia 56 (53–59)% of FA Concavity of tragus well defined

N. parisii

N. woodi

N. macrotis N gambiensis N. thebaica N. vinsonia N. nana

N. intermedia

N. arge

N. major

a

The taxonomic status of N. vinsoni is uncertain: the tragi of the only two specimens are damaged and their shape uncertain. If deeply notched and pear-shaped, the specific status is probably valid, but if smoothy convex and semi-lunate, vinsoni is probably as synonym of N. macrotis (see profile N. vinsoni).

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Family NYCTERIDAE

Andersen (1912b) divided the African species of Nycteris (as Petalia) into four species-groups on the basis of morphological characters, including the size of the lower premolar, the shape of the upper incisors and the shape of the tragus. Using multivariate statistical analyses of morphological characters,Van Cakenberghe & De Vree (1985, 1993, 1998) confirmed that four African species-groups can be recognized. So too did Griffiths (1994) based on hyoid and other morphology. We follow the species and species-group compositions suggested by Van Cakenberghe & De Vree, with the exception that N. parisii is considered distinct from N. woodi on the basis of bacular morphology (Thomas et al. 1994) and the relative size of the ears (see profiles). The African species-groups, their characters and species are:

macrotis group. Posterior lower premolar small; upper incisors bicuspid; tragus semi-lunate, posterior margin smoothly convex. N. macrotis, N. parisii and N. woodi. hispida group. Posterior lower premolar small; upper incisors tricuspid; tragus not semi-lunate, posterior margin smoothly convex. N. aurita, N. grandis and N. hispida. thebaica group. Posterior lower premolar small; upper incisors bicuspid; tragus pear-shaped, posterior margin deeply notched. N. gambiensis, N. thebaica and ? N. vinsoni.

arge group. Posterior lower premolar large; upper incisors bicuspid; tragus with posterior margin concave. N. arge, N. intermedia, N. major and N. nana.

Meredith Happold

The species are presented in alphabetical order, irrespective of the species-group to which they belong.

Nycteris arge  Bates’s Slit-faced Bat Fr. Nyctère de Bates; Ger. Bates’ Schlitznasen-Fledermaus Nycteris arge Thomas, 1903. Ann. Mag. Nat. Hist., ser. 7, 12: 633. Efulen [= Efulan], SW Cameroon.

Taxonomy  Species-group: arge. Synonyms: none. Some authors considered N. intermedia as a synonym. Chromosome number: not known. Description  Small to very small microbat with noseleaf comprised of a longitudinal cleft bordered by fleshy outgrowths; medium-small for a Nycteris (FA: 39–46 mm, GLS: 19.2–20.4 mm); upper incisors bicuspid; posterior lower premolar large; tragus with posterior margin slightly concave; E: 25–34 mm; Tib: 19–26 mm, 48–58% of FA. Not easily distinguished from N. major. Sexes similar. Pelage loose, fluffy; mid-dorsal hairs 9–10 mm. Dorsal pelage medium brown, rusty-brown or dark brown, sometimes strawcoloured on forehead between ears and along their outer margin. Ventral pelage similar but paler; some individuals with irregular whitish or pale brown patches. (One albino individual was found in Taï N. P., Côte d’Ivoire; J. Fahr unpubl.) Ears broad, rounded, of medium length for a Nycteris and also of medium relative length (67 [60–79]% of FA); uniformly blackish-brown. Tragus with slight and ill-defined concavity (‘kink’) at centre of posterior margin (Figure 93i). Muzzle of naked appearance and pink. Wings and interfemoral membrane blackish-brown. Tibia of medium length for a Nycteris (22.0 [19–26] mm) and of medium relative length (52 [48–58]% of FA). Baculum tip simple (Figure 94a; Thomas et al. 1994). Skull with sagittal crest low. Upper incisors bicuspid. Posterior lower premolar comparatively large (rising well above cingula of adjacent teeth and equal or almost equal to height of anterior cusp of first molar) and within toothrow. Geographic Variation  None. Similar Species  Three other African Nycteris have the following combination of characters: upper incisors bicuspid; posterior lower premolar large; tragus with posterior margin concave (Table 18, p. 441):

Nycteris nana. Much smaller (FA: 32–37 mm; GLS: 15.6–17.0 mm). Tibia 4 mm), with tip expanded and trifid. Ventral pelage dark grey in the very few known specimens. Cameroon, Ethiopia, Somalia. Distribution  Endemic to Africa. Zambezian Woodland BZ. Recorded from C and E Zambia, S and N Zimbabwe at lower altitudes ( distance [septum] between pits), medium (width of pit ca. = distance between pits), or

Figure 102. Variations in the length of the third ridge relative to the second ridge of M3, the third upper molar, in Tadarida. (a) Left upper toothrow showing M3. (b) Left M3 with ridges one, two and three labelled. (c) Third ridge > second ridge. (d) Third ridge = second ridge. (e) Third ridge > half of second ridge. (f) Third ridge < half of second ridge. (g) Third ridge absent.

small (width of pit noticeably < distance between pits) (Figure 101c, d and e, respectively). Length of third ridge relative to length of second ridge of M3, the posterior upper molar. The third ridge (the most posterior ridge) can be (a) equal in length or slightly longer than the second, (b) equal to the second, (c) more than half of the second but never equal to it, (d) less than half of the second but not completely absent or (e) absent (Figures 102c–g, respectively). Anterior upper premolar. Can be distinctly shorter than the cingulum of the posterior premolar, and either displaced labially or in toothrow,

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Family MOLOSSIDAE

anterior upper premolar

a

a

b

b

c

d

Figure 104. The upper incisors of Tadarida can be procumbent or not procumbent. (a) Lateral and occlusal views of the procumbent upper incisor in Tadarida nanula that is not obscured by the canine cingula. (b) Lateral and occlusal views of the non-procumbent upper incisor in T. spurrelli that is, in this specimen, almost entirely obscured by the canine cingula. Incisors shown in solid black; both specimens are !!.

e

a Figure 103. Sizes of the anterior upper premolar in African Molossidae relative to the height of the cingulum of the posterior premolar, and its position. Left: occlusal views of teeth on left side of upper jaw. Right: lateral views of labial side of the same teeth. (a) Anterior upper premolar distinctly shorter than the cingulum of the posterior premolar and displaced labially (e.g. Tadarida condylura). (b) Anterior upper premolar distinctly shorter than cingulum of posterior premolar and in toothrow (e.g. T. major). (c) Anterior upper premolar approximately same height as cingulum of posterior premolar and in toothrow (e.g. Sauromys petrophilus). (d) Anterior upper premolar distinctly taller than cingulum of posterior premolar and in toothrow (e.g. T. aloysiisabaudiae). (f) Anterior upper premolar absent (e.g. Myopterus whitleyi).

ca. same height as cingulum of posterior premolar, distinctly taller than cingulum of posterior premolar, or absent (Figures 103a–e, respectively). Upper incisors. Can be procumbent (angled forward), or not procumbent (pointing downwards and/or somewhat inwards) (Figures 104a and b, respectively).

b

Figure 105. Lower incisors and canines of African Tadarida. (a) Canine cingula not greatly enlarged and not in contact; incisors crowded (e.g. T. aegyptiaca). (b) Canine cingula greatly enlarged and in contact; incisors reduced in number and crowded (T. spurrelli !).

Lower incisors. There can be one, two or three on each side, and they can be crowded by the cingula of the lower canines, or not crowded (Figure 105). Lower canines. The cingula are always large and conspicuous compared with bats in other families but, in some species of molossids, they are greatly enlarged (Figure 105). The cingula are often larger in !! than in "" of the same species. The six genera represented in Africa are presented in alphabetical order. Meredith Happold & F. P. D. Cotterill

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Table 19.  Key to the African genera in the family Molossidae (based on African species only). For details of the differences in these characters, see Family Molossidae.

Inner margins of ears (Folding of ears)

Length of third ridge (3) in relation to second ridge (2) of posterior upper molar (M3)

Widely separated (Without complex folding)

3 = 2

Extremely flattened

Meeting to form V-shaped valley (Without complex folding)

3 = 2

Low but not extremely flattened

Distinctly separated or almost touching (Without complex folding)

3≥2

Not extremely flattened but sometimes low

Meeting (or almost meeting) to form V-shaped valley or Joined by band (With complex folding)

Varied: 3 = 2 3 > half 2 3 < half 2 3 absent

Low but not extremely flattened

Widely separated or Meeting to form V-shaped valley (Without complex folding)

3 absent

Low but not extremely flattened

Forward-projecting and joined to forehead and along extended muzzle (With moderate folding)

Head and skull

Extremely flattened

3≤2

Miscellaneous

Forearm with warty granulations FA: 27–36 mm Upper lip with many fine wrinkles and many stiff bristles Forearm without warty granulations FA: 37–50 mm Upper lip smooth with only a few stiff bristles Forearm without warty granulations FA: 38–43 mm Ears never white or translucent Upper lip with many well-defined wrinkles but few spoonhairs FA: 27–67 mm Ears never white or translucent Ears 10–33 mm Sometimes with yoke of almost naked, pale skin across nape of neck Upper lip usually with well-defined wrinkles and many spoon-hairs FA: 35–54 mm Ears whitish or translucent Ears 14–22 mm Upper lip smooth with many spoon-hairs FA: 61–74 mm Band of dense white or pale pelage across top of shoulders and along flanks Dorsal hairs conspicuously bicoloured Ears very long (31–42 mm) Upper lip with fine, microscopic wrinkles and no spoon-hairs

Genus

Platymops (1 species) Sauromys (1 species) Mormopterus (1 or 2 species)

Tadarida (27 species)

Myopterus (2 species)

Otomops (1 species)

Genus Mormopterus Little Mastiff Bats and others Mormopterus Peters, 1865. Monatsber. K. Preuss. Akad. Wiss. Berlin 1865: 258. Type species: Nyctinomus (Mormopterus) jugularis Peters, 1865.

Mormopterus was considered to contain ten species (Simmons 2005) but another species, M. francoismoutoui, has been described by Goodman et al. (2008). Species of Mormopterus include M. acetabulosus from Mauritius and M. francoismoutoui from Réunion, at least one of which has been recorded in Africa (but probably only as vagrants), and M. jugularis from Madagascar, which presumably has the potential to reach Africa as a vagrant although it has not yet been recorded there. The other species occur in the Malay archipelago, South America, Cuba or Australia.There are about seven more undescribed species in Australia (Churchill 1998). Mormopterus is included in Tadarida by some authors including Freeman (1981), but considered distinct by Legendre (1984), Meester et al. (1986), Corbet & Hill (1991, 1992) and others. Mormopterus is sometimes considered to include Platymops and Sauromys (e.g. Koopman 1993, 1994) but these genera are currently considered

distinct (Corbet & Hill 1992, Peterson et al. 1995, Simmons 2005). The only three species of Mormopterus likely to be recorded in Africa are distinguished from species in other molossid genera in Africa by the following combination of characters: head, body and skull low but not extremely flattened (cf. Platymops and Sauromys); ears black and not translucent (cf. whitish or translucent in Myopterus); inner margins of ears distinctly separated although close; third ridge of posterior upper molar (M3) equal in length or longer than second ridge; anterior palatal emargination wide; four upper cheekteeth on each side (cf. five in Tadarida except rarely); three lower incisors on each side (cf. one in T. spurrelli, two in all other African Tadarida except T. teniotis, which also has three; see also Table 19, p. 472). Dental formula (M. acetabulosus, M. francoismoutoui, M. jugularis): 1113/3123 = 30. Craniodental characters of Mormopterus are illustrated in Figure 106.

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Mormopterus acetabulosus and M. francoismoutoui

When the Mormopterus profiles were submitted, only two specimens of one species – M. acetabulosus – were said to have been recorded from Africa; one from Ethiopia and one from South Africa. Now, the Ethiopian specimen has been re-identified as M. francoismoutoui but the identity of the South African specimen could not be confirmed because its skull is missing: it could represent either M. acetabulosus or M. francoismoutoui (Goodman et al. 2008). Information about both species and the two African specimens is included in the following profile. Meredith Happold

Figure 106. Skull of Mormopterus francoismoutoui (BMNH 6.11.1.9 from Ethiopia). This specimen was previously considered to represent M. acetabulosus (Goodman et al. 2008).

Mormopterus acetabulosus and M. francoismoutoui MAURITIAN LITTLE MASTIFF BAT AND RÉUNION LITTLE MASTIFF BAT Fr. Mormoptère à lèvres ridées; Ger. Mauritius Mastino-Fledermaus, Réunion Mastino-Fledermaus Mormopterus acetabulosus (Hermann, 1804). Observ. Zool. p. 19. Port Louis, Mauritius. For several reasons, Goodman et al. (2008) designated FMNH 1984.368, from Palma Cave, Palma, Black River District, Mauritius, 20° 16.405' S, 57° 27.147' E, as the neotype of this species. Mormopterus francoismoutoui Goodman, van Vuuren, Ratrimomonarivo, Probst & Bowie, 2008. J. Mammal. 89: 1318. Commune de La Possession, Pont de Balthazar, 2.2 km SSW La Possession, La Réunion, 20° 56.732' S, 55° 19.484' E, 40 m.

Taxonomy Mormopterus acetabulosus was originally described as Vespertilio acetabulosus. Until recently, it was thought to occur mainly on Mauritius and Réunion, but with two specimens (probably vagrants) recorded from Africa. However, based on both molecular and morphological evidence, specimens from Mauritius and Réunion are now considered specifically distinct and the Réunion bats have been described as a new species, Mormopterus francoismoutoui (Goodman et al. 2008) (details in Appendix). Of the two specimens from Africa, one (BM 6.11.1.9 from Ethiopia) is considered by these authors to represent M. francoismoutoui. The other (MNHN 1984-201 from ‘Port Natal’ [now Durban]) is in poor condition and its skull could not be located: consequently Goodman et al. (2008) could not ascertain if it represents M. acetabulosus or M. francoismoutoui. Synonyms: natalensis. Subspecies: none. Chromosome number: not known. Description Very small microbats without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; small for an African molossid; head and body not extremely flattened; ears separated and without complex folds, upper

lip with many well-developed wrinkles; wings dark; forearms without warty granulations; anterior upper premolar absent. Sexes similar. Pelage short (mid-dorsal hairs 3–4 mm). Dorsal pelage uniformly dark brown; hairs unicoloured.Ventral pelage dark brown becoming slightly paler on abdomen. Head somewhat (but not extremely) flattened. Upper lip with many (>7) well-developed wrinkles, a few spoon-hairs and many fine hairs. Ears relatively short, erect, widely separated (by 2–4 mm) in M. acetabulosus, almost touching and joined by a slender band of skin in M. francoismoutoui, subtriangular with slight emargination below tip, without complex folds (cf. Tadarida). Tragus small but not concealed by antitragus. Antitragus low, indistinct. No interaural crest. Gular gland present in !!. Wings and interfemoral membrane dark brown. No wart-like granulations on forearm. Hindfeet relatively large; toes with long hairs and many short, stout bristles. Skull (Figure 106) low but not extremely dorsoventrally flattened (cf. Platymops and Sauromys) (maximum height ca. 41% of GLS as in Tadarida pumila). Rostrum relatively narrow (cf. Platymops). Zygomatic arches slender. Anterior of braincase rising above plane of rostrum; forehead slightly concave. Sagittal crest low or absent; lambdoid crest 473

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low or absent. Lachrymal ridges indistinct and forming a comparatively small tubercle on each side of rostrum (cf. Platymops). Anterior palatal emargination wide. Basisphenoid pits shallow, much wider than their distance apart. Upper incisors well separated. Anterior upper premolar absent. M3 with third ridge ≥ second ridge (Figure 102c and d). Three lower incisors on each side. Lower canines with cingula not greatly enlarged and not in contact. Dental formula: 1113/3123 = 30. Geographic Variation None recorded for either M. acetabulosus sensu Goodman et al. (2008) or M. francoismoutoui. Similar Species complex folds:

on Madagascar (e.g. Hayman & Hill 1971, Kingdon 1974, 1997, Koopman 1993) but no valid specimen appears to be available and the inclusion in Madagascar apparently arises from the confusion of names listed in synonymy dating back to Peters (1865), who proposed that the subgenus Mormopterus includes taxa from South Africa, Madagascar and the Mascarene Is. (Peterson et al. 1995). The lack of further records from Africa suggests that the African records are those of vagrants and that Mormopterus does not normally occur on the mainland. A specimen of ‘Mormopterus sp.’ (FA: 32 mm) from Meru N. P. (Webala et al. 2004) does not represent M. acetabulosus or M. francoismoutoui (M. Happold pers. obs.).

Four other African molossids have ears without

Myopterus daubentonii. Much larger (FA: 48–56 mm; GLS: 21.3– 25.7 mm). Dorsal pelage with contrasting lateral bands of paler colour. Wings with arm-wing white, hand-wing transparent. M3 with third ridge absent. M. whitleyi. Forearm shorter (35–37 mm); skull larger (GLS: 16.6– 18.5 mm). Arm-wing yellowish-white, hand-wing transparent. M3 with third ridge absent. Platymops setiger. Head extremely flattened. Forearms with warty granulations. Skull with very prominent lachrymal ridges. Sauromys petrophilus. Inner margins of ears separated by narrow gap. Head extremely flattened. Anterior upper premolar present. Distribution One specimen (of uncertain identity according to Goodman et al. 2008) was recorded in 1933 from Durban, KwaZulu–Natal, South Africa in the Coastal Forest Mosaic BZ, and one specimen (identified with M. francoismoutoui by Goodman et al. 2008) was recorded in 1905 from Ethiopia (between L. Turkana and Shewa Province), presumably in the Afromontane–Afroalpine BZ of the Ethiopian Highlands. Extralimitally: Mauritius and Réunion (but see Taxonomy). Mormopterus acetabulosus was said to occur

Mormopterus francoismoutoui and ? M. acetabulosus

Habitat Recorded from the edge of a forest near Durban (Smithers 1983); no data for the Ethiopian locality. On Réunion, M. francoismoutoui is widespread and found from sea level to ca. 2000 m. Abundance Only two specimens recorded in Africa. Mormopterus francoismoutou appears common on Réunion. Remarks On Réunion, M. francoismoutoui roosts in caves, crevices in rockfaces, and similar sites in bridges and buildings (Goodman et al. 2008). Anciaux de Faveaux (1984) catalogued a mite (Ewingana lamorali [Acari: Myobiidae]) from a Kenyan specimen said to represent M. acetabulosus, but the identity of the specimen has not been confirmed. Apparently nothing else is known about Mormopterus in Africa. Conservation IUCN Category (M. acetabulosus): Vulnerable (based on data from Mauritius and Réunion). Measurements Mormopterus acetabulosus FA: 40.2 (38–43) mm, n = 35 WS: (d) 242.5 (240–245) mm, n = 4* TL: 98.2 (92–102) mm, n = 34 T: 44.9 (43–50) mm, n = 34 E: 17.0 (15–18) mm, n = 34 Tr: 3.6 (3.3–3.8) mm, n = 4* Tib: n. d. HF: 6.8 (6–7) mm, n = 34 WT: 7.3 (6.0–8.2) mm, n = 34 GLS: 15.1 (14.6–15.7) mm, n = 34 GWS: 8.8 (8.5–9.1) mm, n = 30 C–M3: 5.6 (5.3–5.9) mm, n = 36 Mauritius (NMW, Goodman et al. 2008) *NMW Mormopterus francoismoutoui FA: 39.6 (38–42) mm, n = 33 WS: n. d. TL: 93.4 (89–97) mm, n = 33 T: 40.2 (38–45) mm, n = 33 E: 16.7 (15–18) mm, n = 33 Tr: n. d. Tib: n. d. HF: 6.1 (5–6.5) mm, n = 33 WT: 6.0 (5.0–7.2) mm, n = 33 GLS: 14.8 (14.1–15.3) mm, n = 35

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GWS: 8.5 (8.1–8.9) mm, n = 33 C–M3: 5.5 (5.2–5.8) mm, n = 33 Réunion (Goodman et al. 2008) Key References

Goodman et al. 2008; Peterson et al. 1995. Meredith Happold

GENUS Myopterus Winged-mouse Bats Myopterus Oken, 1816. Lehrbuch der Naturgeschichte, pt 3 sect. 2: 932. Type species: Myopterus senegalensis Oken, 1816 (not available). Myopterus E. Geoffroy, 1818. Description de l’Egypte 2: 113. Type species: Myopterus senegalensis Oken, 1816 (not available = Myopterus daubentonii Desmarest, 1820).

Myopterus (including Eomops as a synonym) is a polytypic genus with two species, endemic to tropical Africa from Senegal in the West to DR Congo and Uganda in the East, occurring in both rainforests and moist savannas. Distinguished from other genera of molossids in Africa by the following combination of characters: head, body and skull low but not extremely flattened dorsoventrally (cf. Platymops and Sauromys), wings translucent or whitish, ears whitish or translucent, without complex folding, either widely separated or meeting medially to form V-shaped valley; upper lip smooth with many spoon-hairs; third ridge of posterior M3 absent; dental formula: 1113/1123 = 26 (see also Table 19, p. 472). Bats in this genus also have white ventral pelage;

relatively short, narrow and erect ears; a large tragus (not concealed by the antitragus); upper incisors in contact; only one upper premolar on each side; one pair of lower incisors; anterior palatal emargination closed; and deep basisphenoid pits (distinct hole >1 mm in depth). Selected characters of a representative of the genus are illustrated in Figure 107, but there are some inter-specific differences.The savannadwelling species M. daubentonii is much larger than M. whitleyi, which is restricted to the rainforest zone. The latter species is one of the smallest molossids in Africa. In collections, both species are among the rarest African molossids. Almost nothing is known about their biology. They have been rarely captured except from their day-roosts,

Figure 107. Myopterus. Flight membranes of M. daubentonii. Unique dorsal pelage markings of M. daubentoni. Skull of M. whitleyi (RMCA 97-021-M-0825).

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which are sometimes rather unusual for molossids as M. whitleyi has been found roosting, unconcealed, in vegetation. In her study of the family Molossidae, Freeman (1981) found phenetic similarity between Myopterus and the Neotropical genera Molossops, Neoplatymops and Cynomops. According to her study, Myopterus is one of the most derived molossid genera and the most derived of all African molossids. Freeman (1981) speculated that Myopterus has no extant relative in the Old World but rather a Neotropical origin, with Molossops and Cynomops being the most closely related genera. Likewise, based on dental characters, Legendre (1984) would place Myopterus in the subfamily Molossinae, which in his arrangement

comprises otherwise New World genera, whereas all other molossid genera occurring in Africa would be included in the subfamily Tadaridinae. In a recent cladistic analysis by Jones et al. (2002), Myopterus grouped with the Asian Cheiromeles and Neotropical Molossops, again without any close relationship to African molossids. The two species are: M. daubentonii. FA: 48–56 mm; GLS: 21.3–25.7 mm; WT: 20–22 g. M. whitleyi. FA: 35–37 mm; GLS: 16.6–18.5 mm; WT: 10–12.0 g. Jakob Fahr

Myopterus daubentonii DAUBENTON’S WINGED-MOUSE BAT Fr. Myoptère de Daubenton; Ger. Daubentons Pergamentflügel-Fledermaus Myopterus daubentonii Desmarest, 1820. Encyclopédie Méthodique Mammalogie 1: 132. Senegal (no specified locality).

Taxonomy Synonyms: albatus. Subspecies: two. The taxonomic identity of this bat has been debated for a long time (Rosevear 1965). A specimen from Senegal was described by Daubenton (1765) and named ‘rat-volant’, which is not a proper binomial name. Based on the description of Daubenton, it was renamed Myopterus senegalensis by Oken (1816) and the specimen was examined by Peters (1869). The holotype was subsequently lost and therefore Brosset & Vuattoux (1968) designated a new ‘type’ and ‘cotype’ based on specimens from Côte d’Ivoire. Hill (1969) pointed out that the name senegalensis was not available and referred the specimens from Côte d’Ivoire to M. albatus Thomas, 1915, at that time known from two localities in DR Congo. Finally, Adam et al. (1993) designated one neotype and four neoparatypes for M. daubentonii Desmarest, 1820, a name that was also based on the ‘rat-volant’ of Daubenton, from a series collected in Senegal. They synonymized M. albatus as a subspecies of M. daubentonii, referring specimens from Côte d’Ivoire and Central African Republic to M. d. albatus, while restricting the nominate form, M. d. daubentonii, to the Senegal population (but see Geographic Variation). Chromosome number: not known.

patch of spoon-hairs on upper lip below nostrils; upper lip smooth. Ears erect (without complex folds), narrow, with outer side partly pigmented and inner side whitish; bases of inner margins meeting on forehead to form V-shaped valley. Tragus relatively very large for a molossid, simple, anterior and posterior margins nearly parallel, tip rounded; not concealed by antitragus. Antitragus semi-circular, ca. same size as tragus. No interaural crest. No gular gland. Wings with arm-wing membranes white and hand-wing membranes strikingly transparent and showing red veining and some enclosed white spots of fat. Wing-membranes of living bats feel moist and somewhat sticky. Interfemoral membrane dorsally pale brown, ventrally white. Skin of forearm, tibia, fingers and tail pink. Skull large and robust; not extremely dorsoventrally flattened. Sagittal and lambdoid crests absent or weakly developed. Anterior palatal emargination closed. Basisphenoid pits very deep, large, their width much greater than their distance apart. Upper incisors in contact. Anterior upper premolar absent. M3 with third ridge absent. Only one lower incisor on each side, bicuspid (inner cusp distinctly higher than outer cusp), not crowded. Lower canines with cingula not greatly enlarged, well separated. Dental formula 1113/1123 = 26.

Description Small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; medium-sized for an African molossid (FA: 48–54 mm); head not flattened; ears meeting on forehead to form V-shaped valley and without complex folds, inner side whitish; upper lip smooth; dorsal pelage usually with longitudinal banding, wings partly white and partly colourless and transparent. Sexes similar in colour; "" on average slightly larger than !!. Pelage short (mid-dorsal hairs 4–5 mm). Dorsal pelage dark cream, reddishbrown or greyish-brown usually with two lateral bands of whitish or pale yellowish-fawn on each side of the mid-dorsal line (Figure 107); hairs (except those of bands) whitish or cream-coloured with darker tip. Crown and nape with large patch of same colour as the lateral bands. Ventral pelage uniformly dirty- or creamy-white to pure white, much paler than, and conspicuously contrasting with, the dorsal pelage; no mid-ventral markings; ventral flank-stripe same colour as flanks. Head not dorsoventrally flattened; muzzle subcylindrical, smooth and rounded, almost naked except for dense

Geographic Variation Two subspecies are recognized (Adam et al. 1993), but variability seems to be high in this species: M. d. daubentonii. Originally considered to be restricted to Senegal (Adam et al. 1993), but a re-examination and comparison of published data with data from new specimens from Côte d’Ivoire show that the dimensions of specimens from Senegal, Côte d’Ivoire and Central African Republic overlap sufficiently to justify assigning all of them to the nominate subspecies. FA: 50.4 (48– 54) mm; TL: 110.9 (104–120) mm; GLS: 22.5 (21.3–24.9) mm; GWS: 13.4 (13.0–14.5) mm (see Measurements). M. d. albatus. Considered to include all populations east of Senegal by Adam et al. (1993), but considered here to include only specimens from DR Congo, which are clearly larger than those from Senegal to Central African Republic. For five "" from Niangara, DR Congo, FA: 53.7 (52–56) mm; TL: 126.0 (123–132) mm; GLS: 25.2 (24.2–25.7) mm; GWS: 14.5 (14.1–14.7) mm (Allen 1917a, Hill 1969).

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Adaptations Most of the known specimens were taken in their day-roosts in hollow trees. These include the hollow trunk of a dead Palmyra Palm Borassus aethiopum in which the bats roosted at a height of ca. 12 m (Brosset & Vuattuox 1968). According to Freeman (1981), species of Myopterus are characterized by broad wing-tips. She concluded that these species are probably more manoeuvrable than most other molossids and possibly fly more slowly. In contrast, preliminary flight-cage experiments indicate that M. daubentonii is less manoeuvrable than 16 other species of molossids tested (J. Fahr unpubl.). Individuals from Comoé N. P., Côte d’Ivoire, had a strong, spicy and almost pungent smell. Foraging and Food Foraging strategy: fast-hawking. In Comoé N. P., in mist-nets set 0–25 m above ground, one ! and two "" were caught at the same time, at heights of 9, 14 and 20 m (J. Fahr unpubl.). Possibly these individuals were foraging together as groupforaging seems to be common in molossid bats. The mist-nets were set between two large trees in woodland savanna not far from gallery forest. The teeth and mandibles are very robust and powerful; from this, Freeman (1981) deduced that this bat is probably able to consume large and hard-shelled prey. Myopterus daubentonii

Similar Species complex folds:

Four other African molossids have ears without

Myopterus whitleyi. Much smaller (FA: 35–37 mm; GLS: 16.6– 18.5 mm). Dorsal pelage without lateral bands. Mormopterus spp. Much smaller (FA: 38–43 mm; GLS: 14.1–15.7 mm). Dorsal pelage without lateral bands. Wings dark brown. M3 with third ridge ≥ second ridge. Sauromys petrophilus. Dorsal pelage uniformly dark brown. Wings blackish-brown, brown or semi-translucent greyish-brown. M3 with third ridge present. South of 15° S. Platymops setiger. Much smaller (FA: 27–36 mm; GLS: 14.7–17.0 mm). Dorsal pelage uniformly coloured. Distribution Endemic to Africa. Mainly found in the Guinea Savanna BZ and the Northern Rainforest–Savanna Mosaic, but extending into the Sudan Savanna BZ in Senegal. Known from only eight localities in Senegal, Côte d’Ivoire, Central African Republic and DR Congo (Lang & Chapin 1917b, Hayman et al. 1966, Adam et al. 1993, J. Fahr unpubl.). Habitat Predominantly a savanna species recorded in the following vegetation zones: Sudanian undifferentiated woodland savanna in Senegal, Isoberlinia woodland (e.g. N’Délé in Central African Republic) and Rainforest–Savanna Mosaic (e.g. Lamto and Comoé N. P. in Côte d’Ivoire). Also recorded at 1250 m in montane grassland (Bunia, DR Congo). Specimens from Lamto were found roosting in a palm in grassland ca. 100 m from gallery forest (Brosset & Vuattuox 1968). Abundance Uncertain. Only 23 specimens known. Apparently quite localized and rare.

Social and Reproductive Behaviour Very little is known. One group in a hollow tree included a subadult ! and six "", but not all group-members were captured (Lang & Chapin 1917b). Another group included an adult ! and four adult "" (not stated if all group-members were captured) (Brosset & Vuattuox 1968), and a third group included three adult !!, one subadult ! and one subadult " (Adam et al. 1993). Reproduction and Population Structure Predators, Parasites and Diseases

No information.

No information.

Conservation IUCN Category: Data Deficient. Distribution rather broad, but only eight localities known. Population trend inferred to be declining, in both past and future. Major threats: loss and degradation of habitat, particularly as result of logging and agriculture. Measurements Myopterus daubentonii daubentonii FA: 50.4 (48–54) mm, n = 14 WS (c): 368 (350–380) mm, n = 4 TL: 110.9 (104–120) mm, n = 14 T: 40.8 (37–44) mm, n = 14 E: 20.1 (18–22) mm, n = 14 Tr: 4.7 (4.6–5.0) mm, n = 3 Tib: 14.9 (14–17) mm, n = 13 HF: 11.2 (9–13) mm, n = 9 WT: 21.3 (20.0–22.0) g, n = 4 GLS: 22.5 (21.3–24.9) mm, n = 11 GWS: 13.4 (13.0–14.5) mm, n = 10 C–M3: 8.5 (8.0–9.2) mm, n = 11 Senegal, Côte d’Ivoire, Central African Republic (FC, MHNG, MNHN, SMF, USNM) 477

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Specimens from DR Congo (M. d. albatus) are on average larger (see Geographic Variation) Key References Adam et al. 1993; Allen 1917a; Brosset & Vuattoux 1968; Hill 1969; Lang & Chapin 1917b; Rosevear 1965. Jakob Fahr

Myopterus whitleyi BINI WINGED-MOUSE BAT (WHITLEY’S WINGED-MOUSE BAT) Fr. Myoptère de Whitley; Ger. Bini Pergamentflügel-Fledermaus Myopterus whitleyi (Scharff, 1900). Ann. Mag. Nat. Hist., ser. 7, 6: 569. Benin City, Nigeria.

Taxonomy Originally Mormopterus whitleyi. Synonyms: none. Chromosome number: not known. Description Very small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; small for an African molossid (FA: 35–37 mm); head not flattened; ears well separated and without complex folds, whitish; upper lip smooth; wings partly white and partly transparent, contrasting with uniformly dark brown dorsal pelage. Sexes similar in colour, "" on average slightly larger than !!. Pelage short (mid-dorsal hairs 4–5 mm). Dorsal pelage dark brown without stripes; hairs whitish with medium to dark brown at tip. Ventral pelage uniformly dirty-white, cream or pure white, in marked contrast to dorsal pelage; no midventral markings; ventral flank-stripe white. Head not dorsoventrally flattened; muzzle smooth and rounded; almost naked except for dense patch of very stiff spoon-hairs on upper lip below nostrils, and fringe of stiff hairs along edge of upper lip; upper lip smooth. Ears erect (without complex folds), narrow and pointed, with apex unpigmented; bases of inner margins widely separated. Tragus large, squarish, not concealed by antitragus. No interaural crest. Gular gland in !!.Wings with armwing membranes yellowish-white, hand-wing membranes transparent. Interfemoral membrane dorsally grey or pale brown, ventrally whitish. Skin on forearm, tibia and third finger dark pink. Skull (Figure 107) low but not extremely dorsoventrally flattened. Sagittal crest absent or very indistinct except at the very back of the braincase where, in fully mature adults, it rises gradually or abruptly to form a small wing ca. 4 mm long. Lambdoid crest absent or weakly developed. Anterior palatal emargination closed. Basisphenoid pits deep. Upper incisors in contact. Anterior upper premolar absent. M3 with third ridge absent. Only one lower incisor on each side; bicuspid, not crowded. Lower canines with cingula not greatly enlarged, well separated. Dental formula: 1113/1123 = 26.

Sauromys petrophilus. Inner margins of ears meet to form V-shaped valley. Head extremely flattened. Wings blackish-brown, brown or semi-translucent greyish-brown. South of 15° S. Platymops setiger. Head extremely flattened. Forearms with warty granulations. M3 with second and third ridges equal in length. Distribution Endemic to Africa. Known only from the Rainforest BZ (Western, West Central, South Central and East Central Regions) and margins of the Eastern and Southern Rainforest–Savanna Mosaics. Known only from 20 localities in Ghana, Nigeria, Cameroon, Central African Republic, Gabon, DR Congo and Uganda (Hayman et al. 1966, Happold 1987, Adam et al. 1993, Van Cakenberghe et al. 1999, Barrière et al. 2002, J. Fahr unpubl.). Although based on rather few records, it seems that the distribution pattern is patchy or even disjunct. Conspicuously absent from central Congo Basin but its distribution in DR Congo is not confined to the Kasai River System as proposed by Hayman & Hill (1971) because it has also been found along the lower stretches of the Congo R. A record from Eala (Schouteden 1944, 1948, mentioned by Aellen [1952] as from

Geographic Variation None recorded, but data are limited. Similar Species complex folds:

Four other African molossids have ears without

Myopterus daubentonii. Much larger (FA: 48–56 mm; GLS: 21.3–25.7 mm). Dorsal pelage with contrasting lateral bands of paler colour. Mormopterus spp. Forearm longer (38–43 mm); skull smaller (GLS: 14.1–15.7 mm). Wings and interfemoral membrane dark brown. M3 with third ridge ≥ second ridge.

Myopterus whitleyi

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Coquilhatville) was based on a specimen (RMCA RG 5392) that Hayman et al. (1966) re-identified as Tadarida pumila. Habitat Mainly recorded in rainforest–savanna mosaic, lowland rainforest and coastal forest vegetation zones, but also found in swamp forest and mangroves. Most of the localities are situated along the periphery of the Rainforest BZ. A single record from Kasenye near L. Albert, DR Congo, is from Isoberlinia woodland. Specimens from Calabar, Nigeria, were caught in a garden (ROM). Abundance No information, but localized. Remarks Day-roosts of M. whitleyi are variable and in some cases unusual for a molossid: specimens have been taken among the leaves of plantains, clinging to the outside of a Cola tree and in houses (Rosevear 1965). Surprisingly, a single, uninjured individual was captured on the ground in a bucket in a pitfall-trap line (Barrière et al. 2002). Predators, Parasites and Diseases Ectoparasites include fleas Lagaropsylla obliqua, Allopsylla hetera and A. lobayensis (Siphonaptera: Ischnopsyllidae) and a mite Dentocarpus tenuis (Acari: Chirodiscidae) (Beaucournu & Fain 1982, Anciaux de Faveaux 1984, Barrière et al. 2002).

Conservation

IUCN Category: Least Concern.

Measurements Myopterus whitleyi FA: 35.8 (35–37) mm, n = 7 WS: n. d. TL: 81.7 (78–87) mm, n = 7 T: 27.7 (25–31) mm, n = 7 E: 14.8 (14–16) mm, n = 7 Tr: 4.6, n = 1 Tib: 10.2 (9–12) mm, n = 6 HF: 7.0 (5.0–8.3) mm, n = 7 WT: 10.7 (10.0–12.0) g, n = 3 GLS: 17.6 (16.6–18.5) mm, n = 8 GWS: 10.5 (10.2–10.8) mm, n = 8 C–M3: 6.5 (6.1–6.9) mm, n = 8 Ghana, Nigeria, Cameroon, Gabon, Uganda, DR Congo (BMNH, HZM, MHNS, MNHN, USNM, Van Cakenberghe et al. 1999) Key References 1905.

Adam et al. 1993; Rosevear 1965; Thomas Jakob Fahr

GENUS Otomops Giant Mastiff Bats Otomops Thomas, 1913. J. Bombay Nat. Hist. Soc., 22: 91. Type species: Nyctinomus wroughtoni Thomas, 1913.

Otomops martiensseni.

Otomops currently contains seven species of which only one occurs in Africa; the others occur in Madagascar, S India, Java, Lesser Sundas Is. or New Guinea (Simmons 2005). Distinguished from other genera of molossids in Africa by the following combination of characters: head, body and skull low but not extremely flattened dorsoventrally; ears uniquely shaped with only moderate folding, forward-projecting and joined to the long muzzle (Figure 97b); no tragus; no antitragus; dorsal hairs bicoloured; a band of pale or white pelage across shoulders and along each flank above the wings; upper lip with fine, microscopic wrinkles and no spoon-hairs; basisphenoid pits very deep; third ridge of posterior upper molar (M3) as long or almost as long as the second ridge; dental formula: 1123/2123 = 30 (see also Table 19, p. 472). Selected characters of Otomops martiensseni are illustrated in Figure 108.

Otomops martiensseni showing ear flap folded down (upper) and open (lower).

The genus was reviewed by Peterson et al. (1995). Based on nonmetric multidimensional scaling,these authors recognized all previously named forms of Otomops as distinct species (with the exception of 479

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Figure 108. African Otomops. Flight membranes, and bones of wing, hindlimb and tail of O. martiensseni. Skull of O. martiensseni (BMNH 1975-914).

O. johnstonei from Indonesia, which was not described until 1992 and not included in their study), but they considered O. icarus very similar to O. martiensseni, and were uncertain of its specific status. This form is now considered to be a subspecies of O. martiensseni (see species profile). Peterson et al. (1995) found O. madagascariensis from Madagascar to be morphologically quite different from the African

O. martiensseni, and more closely related to O. formosus from Java, Indonesia than to any other species. The species found in Africa is O. martiensseni. Meredith Happold

Otomops martiensseni LARGE-EARED GIANT MASTIFF BAT Fr. Grand molosse oreillard; Ger. Großohrige Riesen-Bulldoggfledermaus Otomops martiensseni (Matschie, 1897). Arch. Naturgesch. 63 (1): 84. Magrotto Plantation, near Tanga, SE Usambara Mts, Tanzania.

Taxonomy Originally Nyctinomus martiensseni. Synonyms: icarus. Subspecies: two. A Madagascan form, madagascariensis, is considered to be a distinct species by Peterson et al. (1995) and Goodman (2011) although often considered a subspecies of O. martiensseni (e.g. Harrison 1957a, Koopman 1993, 1994). Chromosome number (Kenya): 2n = 48; aFN = 56 (Duli´c & Mutere 1973b) or 58 (Warner et al. 1973). Description Medium-sized microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; the largest African molossid (FA: 61– 74 mm); ears very large, forward-projecting with inner margins joined together and to muzzle; dorsal pelage with band of pale pelage across shoulders and along flanks; dorsal hairs bicoloured. Sexes similar in colour; !! on average larger than "". Pelage soft, dense; mid-dorsal hairs 6–7 mm. Dorsal pelage dark brown with dark rusty-brown tinge, or blackish-brown, darker on head and body, with pale brown or white band across shoulders, and a thin band of pale brown separating the dark dorsal pelage from the dark wing-membrane on each side; mid-dorsal hairs dark brown with whitish base. Ventral pelage dark brown, throat paler. Head not flattened; face pink; snout ‘pig-like’. Upper lip expansible with many fine wrinkles, no spoon-hairs. Ears rounded, fairly stiff, projecting forward well beyond snout; inner margins joined together and to muzzle (Figure 97b). Tragus minute; antitragus absent. A semi-circular flap extends forward from base of ear;

can be folded down to seal auditory meatus. No interaural crest. Gular sac (gular gland) present in both sexes; well developed in adult !! (Harrison 1957b). Wings and interfemoral membrane blackish-brown. Skull (Figure 108) not extremely dorsoventrally flattened. Anterior of braincase moderately elevated above plane of rostrum. Each zygomatic arch with a very prominent vertical projection. Sagittal crest moderate; lambdoid crest low or absent; no helmet. Anterior palatal emargination very narrow and deep. Basisphenoid pits very deep and much wider than the very narrow septum between them (as in Figure 101b). Dentition weak. Anterior upper premolar small but rising well above cingulum of posterior premolar. M3 with third ridge equal or just a little shorter than second. Two lower incisors on each side, bicuspid, not crowded. Lower canines with cingula weakly developed (cf. Tadarida) and well separated. Dental formula: 1123/2123 = 30. Similar Species

None.

Geographic Variation subspecies:

Koopman (1994) lists two African

O. m. icarus: Angola and Malawi to KwaZulu–Natal. Smaller (FA [KwaZulu–Natal]: 62–68 mm). O. m. martiensseni: remainder of African geographic range. Larger (FA [Ethiopia]: 69–74 mm).

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Otomops martiensseni

Otomops martiensseni

The validity of icarus as a subspecies, and its distribution, have been questioned (Hill & Carter 1941, Meester et al. 1986).The possibilities of a clinal decrease in size with increasing latitude within one species, or the existence of two distinct species, need investigation. See also Lamb et al. (2006, 2008) and Monadjem et al. (2010). Distribution In Africa, widely but apparently disjunctly distributed in the Somalia–Masai Bushland, Coastal Forest Mosaic, Zambezian Woodland and Afromontane–Afroalpine BZs in the eastern side of Africa from Djibouti to KwaZulu–Natal, South Africa. Also three isolated records in the Guinea Savanna BZ (Côte d’Ivoire, Ghana, Central African Republic), one record from the Rainforest BZ (DR Congo), and one isolated record from the Zambezian Woodland BZ (Angola). This bat is difficult to catch (except at day-roosts), therefore the disjunctions probably reflect insufficient sampling. Extralimitally:Yemen. Habitat Found in a wide range of habitats, from semi-arid scrub to montane forests, from urban and agricultural areas to undisturbed areas, and from sea level to 2000 m. Abundance Uncertain. Seldom recorded throughout most of geographic range, but two colonies numbering >300 and several thousand were reported in 1960s by Mutere (1973b). Kock et al. (2005) provides further evidence that the two breeding colonies in Kenya may fluctuate in numbers (possibly due to migration) but can number tens of thousands. Common in certain residential suburbs in and around Durban, KwaZulu–Natal (Fenton et al. 2002). Adaptations Aspect ratio very high; wing-loading high to very high; wing-tips broad; flight fast (9.5 m/sec for bat flying 20 m above ground); agile with poor manoeuvrability; uses side-slips to decrease altitude rapidly (Norberg 1976, Norberg & Rayner 1987, Rydell & Yalden 1997, Fenton et al. 2002). During flight, the somewhat rigid

ears project forward; the flap behind the antitragus can either open or close the auditory meatus – Kingdon (1974) suggests that the flap closes to increase streamlining; Valdivieso et al. (1979) suggest that it closes to protect the ear from intense noises. In Kenya, roosts by day on ceilings of caves including lava tunnels, preferring darker, poorly ventilated positions (Mutere 1973b). Becomes torpid during day (Mutere 1973b, D. W. Yalden pers. obs.). Emerges well after dark. Sometimes shares caves with Rhinolophus sp., Miniopterus sp. and Triaenops afer. In KwaZulu–Natal, roosts by day under roofs of houses and apartments in built-up areas, preferring old 2–3 storey houses, some of which have been occupied by bats for at least nine years (Taylor, P. 1998, Fenton et al. 2002). Entrances are high to allow diving to gain sufficient speed for flight, and so small that the bats have to scuttle in and out of them. Individuals hang or cling, face downwards, with their bellies in contact with vertical surfaces, or cling to horizontal surfaces. They huddle together and do not appear to become torpid during the day. Radio-tracking revealed that !! and also "" in different reproductive conditions used at least two day-roosts and up to four different day-roosts over a 12day period (Fenton et al. 2002). Bats begin to emerge 15–30 min after sunset. Otomops martiensseni is docile to handle and, despite its pugnacious appearance, makes no attempt to bite handlers. For further adaptations, see Long (1995). Foraging and Food Forages by fast-hawking in open spaces, probably at fairly high altitudes (see below). Because food resources near caves sheltering thousands of individuals are likely to be limited, these bats presumably travel long distances from their day-roosts while foraging. Local fluctuations in populations suggest that some individuals migrate seasonally at times of food-shortage (Mutere 1973b), but this needs confirmation. Radio-tracked bats in the Durban area, South Africa, covered considerable distances while foraging; most bats foraged more than 3 km from their day-roosts for most of the night; one ! night-roosted 6.8 km from his dayroost, and two "" night-roosted at least 10 km away (Fenton et al. 2002). Analysis of faecal pellets from Kenya and Rwanda indicated that moths comprise 97–100% by volume of the diet (Rydell & Yalden 1997) and, as in other molossid moth-specialists, the jaws are slender and the dentition weak (Freeman 1981). Echolocation Search-phase call-shapes (South Africa): (a) very shallow linear or quasi-linear FM; start-frequency ca. 18 kHz; endfrequency ca. 9 kHz; peak-frequency 10–12 kHz; call-duration 5–50 ms; maximum energy in fundamental harmonic, other harmonics uncommon; and (b) lilt calls – as above but end with a shallow upward sweep (lilt), and include the second harmonic (Fenton et al. 2002). Lilts are most common in calls lasting over 25 ms. In cluttered surroundings (e.g. when emerging from day-roosts) individuals emit pairs of multiharmonic calls (7–16 kHz, peak-frequency 11 kHz), which are audible to humans (Taylor 2000). No feeding-buzzes were heard during more than 20 h of monitoring. Calls with frequencies 10–13 kHz, recorded 500–550 m above ground, were assumed to have been emitted by O. martiensseni by Fenton & Griffin (1997) because they resembled the low frequency calls of a light-tagged individual (Fenton & Bell 1981). These audible calls are lower in frequency than those detected by species of moths that can hear ca. 20–40 kHz and dive to avoid capture by bats echolocating at these frequencies. 481

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Social and Reproductive Behaviour In Kenya, O. m. martiensseni roosts in very large groups (colonies) of several hundred to several thousand individuals of both sexes; social organization not known. In KwaZulu–Natal, O. m. icarus roosts singly, or in groups of up to 29 individuals that are typically comprised of one adult !, several adult "" and young: this suggests a harem mating system (Taylor, P. 1998, Fenton et al. 2002). Discounting four records of single bats, and two records of only two bats, 24 colonies averaged 11.2 ± 6.7 individuals (mean ± S.D.). Males with well-developed gular sacs (gular glands) were found with "" and young outside the mating season, suggesting that the harem associations are not just related to mating, and yearround maintenance of harems is suggested by Fenton et al. (2004). Radio-tagged bats from different colonies were never observed roosting in the same houses. Roosting group-members usually hang in physical contact with each other, and make social vocalizations, which are audible to humans. Group-members emerge from a roost singly, at 3–10 min intervals, but remain in the vicinity and emit FM sweeps that have individual signatures permitting individuals to be recognized; these seem to have a communication function and perhaps are not used in echolocation (Fenton et al. 2004). This behaviour continues for ca. 5–30 min, apparently until other bats from the roost join those calling in flight. Because, on any one day, group-members are sometimes in different buildings, their signature-calls perhaps play an important role in synchronizing group activities away from the roosts (Fenton et al. 2004). Reproduction and Population Structure Litter-size: one. Left ovary non-functional; implantation always in right horn. At 1–3° S (Ithundu and Mt Suswa, inland Kenya), the reproductive chronology is mainly restricted seasonal monoestry with 86% of 153 "" pregnant in Nov and most births at beginning of wet season (Nov–Dec) (Mutere 1973b). Gestation suggested to be ca. three months. A few pregnant "" were recorded in Jan (4%), Feb (1%) and May (2%) but no evidence of polyoestry was recorded. Testes size maximum in Aug, low Dec–May; gular sac of !! shows similar cycle. Neonates naked and pink; seldom cling to their mothers; form clusters often apparently surrounded by their mothers. Sex ratio of juveniles usually 1 : 1, but the ratio of !! : "" in adults apparently varies from 1 : 2 to 3 : 1 in different sites and years. At 29–30° S (KwaZulu–Natal, South Africa) the chronology appears to be extended seasonal monoestry, with juveniles recorded Oct–Jan and in May and Jun; but data are limited and it is not yet known if some "" are polyoestrous (Taylor, P. 1998, Fenton et al. 2002). The ratio of adult !! : "" in colonies was skewed towards "" and, in five colonies, ranged from 1 : 2 to 1 : 11 (Fenton et al. 2002). The sex ratios at birth and in the adult population as a whole are not known.

Predators, Parasites and Diseases Ectoparasites include a flea Araeopsylla scitula (Siphonaptera: Ischnopsyllidae); a tick Carios boueti (Acari: Argasidae); and mites Chiroptella suswaensis (Acari: Trombiculidae), Olabidocarpus otomops, Labidocarpellus abyssinicus (Acari: Chirodiscidae) (Anciaux de Faveaux 1984, Beaucournu & Kock 1996). Bat fleas parasitizing O. martiensseni increase to tremendous numbers at certain periods of the year; e.g. 662 A. scitula on one juvenile (Beaucornu & Kock 1996). Conservation IUCN Category: Near Threatened. Guano collection from one breeding-cave was thought to be the cause of variation in apparent reproductive success there from year to year (Mutere 1973b).This thinly distributed and highly colonial species would be very vulnerable to disturbance of this sort. In and around Durban, South Africa, where O. martiensseni roosts in houses and is one of the most commonly encountered bats, the bats themselves, and the amateur and professional people studying them, have received wide positive local publicity and this has made many of the human residents of the area more interested in the bats and more tolerant of them as co-inhabitants (Fenton et al. 2002). In this area, O. martiensseni is clearly able to find and exploit roosts in urban areas and thrive in areas of intensive agricultural operations, and could have a role as a flagship species representing the resilience of nature (Fenton et al. 2002). Measurements Otomops martiensseni FA: 66.8 (61–74) mm, n = 93 WS (d): 489.7 (432–549) mm, n = 38 TL: 148.7 (127–163) mm, n = 37 T: 48.9 (39–54) mm, n = 48 E: 39.0 (31–42) mm, n = 38 Tr: n. d. Tib: 21.2 (20–23) mm, n = 11 HF: 13.6 (12–15) mm, n = 12 WT: 34.4 (26–47) g, n = 80 GLS: 26.3 (24.5–28.3) mm, n = 38 GWS: 13.8 (12.9–14.7) mm, n = 49 C–M3: 10.2 (9.4–11.3) mm, n = 40 Ethiopia, Kenya, Uganda, South Africa (BMNH, RMCA, ROM, TM, Fenton et al. 2002, D. W.Yalden unpubl.) Key References Fenton et al. 2002, 2004; Freeman 1981; Long 1995; Mutere 1973b; Rydell & Yalden 1997 D. W. Yalden & Meredith Happold

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Platymops setiger

GENUS Platymops Peters’s Flat-headed Bat Platymops Thomas, 1906. Ann. Mag. Nat. Hist., ser. 7, 17: 499. Type species: Platymops macmillani Thomas, 1906 (= Platymops setiger Peters, 1878).

Platymops is a monotypic genus known only from S Sudan, Ethiopia and Kenya. Distinguished from other genera of molossids in Africa by the following combination of characters: head and body extremely flattened (as in Sauromys); ears widely separated and without complex folding; wart-like granulations on FA (unlike all other African genera); gular sac (gular gland) well developed; upper lip with many fine wrinkles and many stiff bristles; dental formula probably 1123/2123 = 30 but with some variations (see species profile; see also Table 19, p. 472). Selected characters of Platymops setiger are illustrated in Figure 109. Platymops was elevated to generic status (distinct from Mormopterus) by Thomas (1906c), but subsequently considered a subgenus of Mormopterus by some authors including Freeman (1981), Legendre (1984) and Koopman (1993, 1994). Considered here as distinct following Harrison & Fleetwood 1960, Peterson (1965, 1985), Meester et al. (1986), Peterson et al. (1995) and Simmons (2005). The only species currently recognized is P. setiger. Meredith Happold

Platymops setiger.

Figure 109. Platymops setiger. Flight membranes (traced from photo of dried skin in Harrison & Fleetwood 1960), and bones of wing and hindlimbs. Skull (HZM 6.3776): the anterior premolar is missing in this specimen.

Platymops setiger PETERS’S FLAT-HEADED BAT Fr. Molosse à tête plate de Peters; Ger. Peters’ Flachkopf-Bulldoggfledermaus Platymops setiger (Peters, 1878). Monatsber. K. Preuss. Akad. Wiss. Berlin 1878: 196. Taita, Kenya.

Taxonomy Originally Mormopterus setiger but see Genus Platymops. Synonyms: barbatogularis, macmillani, parkeri. Subspecies: none currently recognized. Reviewed by Peterson (1965). Chromosome number (Kenya): 2n = 48; aFN = 54 (Warner et al. 1974).

Description Very small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; small for an African molossid; head and body extremely flattened; ears well separated and without complex folds; upper lip without well-developed wrinkles; dorsal pelage without stripes; wart-like granulations on forearms and elsewhere; anterior 483

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upper premolar minute or absent. Sexes similar. Pelage short (middorsal hairs 3–4 mm). Dorsal pelage pale sepia brown, rusty-brown, brownish-grey or blackish; hairs with basal half dark cream. Ventral pelage rusty-yellow or creamy-white; sometimes with a darker to much darker lateral stripe beginning on neck and extending a variable distance along each flank, with a narrow band of brownish, rustyyellow or creamy-white pelage between the lateral stripe and the wingmembrane. Head flattened dorsoventrally. Upper lip thick, relatively rigid, with many very fine microscopic wrinkles (giving lip a skirtlike appearance), and many dense, stiff bristles. Ears dark brown, well separated, without complex folds (cf. Tadarida). No interaural crest. Both sexes with small gular sac (gular gland) containing tuft of coarse, brownish hairs in the form barbatogularis, untufted in other forms (details in Harrison & Fleetwood 1960). Wings long, narrow, semitranslucent, lightly pigmented with dark brown, especially over bones; wart-like granulations on forearm, thumb and third metacarpal, and also on tail. Toes with long bristles (up to 8 mm). Skull (Figure 109) extremely dorsoventrally flattened (maximum height 28–34% of GLS). Anterior of braincase not elevated above plane of rostrum. Sagittal crest low or absent anteriorly, moderate or absent posteriorly; lambdoid crests moderate to well developed (especially in adult !!). Lachrymal ridges well developed and forming a prominent tubercle on each side of rostrum (cf. Mormopterus and Sauromys). Anterior palatal emargination narrow. No basisphenoid pits. Upper incisors bicuspid, well separated; tall, curved, deciduous milk incisors may be retained in immature specimens. Anterior upper premolar absent in adults but sometimes there is a minute spicule (probably a deciduous milk tooth) in younger animals, displaced labially; canine and posterior premolar in contact or almost so. M3 with second and third ridges equal in length. Usually two lower incisors on each side, bicuspid, crowded. Lower canines with cingula not greatly enlarged and not in contact. Dental formula: probably 1123/2123 = 30 (but see above). Geographic Variation Uncertain. Peterson (1965) recognized two subspecies on the basis of size: P. s. setiger in S Kenya, and a smaller form, P. s. macmillani, in the remaining geographic range. However, Peterson’s samples were very small (eight adult setiger, 11 adult macmillani). Furthermore, as the distribution of this species is apparently contiguous, the variation is possibly clinal. In barbatogularis from Sudan (described as a species but now considered a synonym of macmillani by Simmons 2005), the gular sac contains a prominent tuft of brownish hairs (cf. macmillani) (Harrison 1956). Similar Species complex folds:

Four other African molossids have ears without

Sauromys petrophilus. Ears meeting to form V-shaped valley. Forearms without warty granulations. Upper lip smooth. No gular gland. Skull with lachrymal ridges indistinct. Anterior upper premolar present and about same height as cingulum of posterior premolar; canine and posterior premolar well separated. Mormopterus spp. Head not extremely flattened. Forearms without warty granulations. Skull without very prominent lachrymal ridges. Myopterus daubentonii. Much larger (FA: 48–56 mm; GLS: 21.3– 25.7 mm). Dorsal pelage usually with contrasting lateral bands of paler colour.

Platymops setiger

Myopterus whitleyi. Head not extremely flattened. Forearms without warty granulations. M3 with third ridge absent. Distribution Endemic to Africa. Apparently confined to a narrow belt through the Somalia–Masai Bushland BZ (and marginally into the Sudan Savanna BZ) from SE Sudan and SW Ethiopia to SE Kenya (Largen et al. 1974, Koopman 1975, Aggundey & Schlitter 1984). Not known to extend into the Horn of Africa (contra Kingdon 1974). Habitat At Masalani, near Kibwezi in inland Kenya, recorded from dense thorn scrub (Combretum, Grewia, Premna) with scattered patches of grassland (O’Shea & Vaughan 1980). Elsewhere, recorded from dry stony areas, and areas with rocky hills, in Ethiopian undifferentiated woodland, Somalia–Masai Acacia–Commiphora bushland and thicket, and mosaics of East African evergreen bushland and secondary Acacia wooded grassland (terminology: White 1983). Abundance

Uncertain.

Adaptations Wings short for an African molossid; metacarpal of third finger only ca. one-third longer than metacarpal of fifth finger (Thomas 1906c). Flight described as fast, direct or erratic. Roosts by day in very narrow crevices under stones or rocks lying on the ground, and in fissures in granite hills: sometimes found cohabiting with scorpions (Harrison & Fleetwood 1960). Able to squeeze into these confined spaces because head and body are flattened. Tactile hairs on feet and muzzle (as in other molossids) also facilitate movement into the day-roosts. According to Kingdon (1974), the tip of the tail is held aloft to gauge heights of crevasses, and the ears and raised phalanges of third and fourth fingers are also used for this purpose. Roosts have strong smell. Emergence is at dusk (Williams 1967). Foraging and Food Forages by fast-hawking (O’Shea & Vaughan 1980). Has been observed flying over waterholes and marshes, at

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Family MOLOSSIDAE

heights of 9 m and less (Nowak 1999). Small beetles recorded in diet. Social and Reproductive Behaviour Recorded roosting singly and in groups of up to five (Nowak 1999). No other information. Reproduction and Population Structure Litter-size: one (n = 4; Anciaux de Faveaux 1972, O’Shea & Vaughan 1980). Reproductive chronology not known. Predators, Parasites and Diseases No information. Conservation

TL: 91.3 (72–114) mm, n = 31 T: 29.9 (22–36) mm, n = 31 E: 15.0 (11–17) mm, n = 33 Tr: 3.8 (3.2–4.2) mm, n = 6 Tib: n. d. HF: 7.3 (7–8) mm, n = 10 WT: 12.6 (7–18) g, n = 20 GLS: 16.0 (14.7–17.0) mm, n = 13 GWS: 11.9 (11.2–12.3) mm, n = 8 C–M3: 5.9 (5.0–6.4) mm, n = 27 Throughout geographic range (BMNH, NMK, ROM, SMF, SMNS, Peterson 1965, Koopman 1975)

IUCN Category: Least Concern. Key References Harrison 1956; Harrison & Fleetwood 1960; Nowak 1999; Peterson 1965.

Measurements Platymops setiger FA: 32.6 (27–36) mm, n = 36 WS (a): 257.4 (227–280) mm, n = 20

Meredith Happold

GENUS Sauromys Roberts’s Flat-headed Bat Sauromys Roberts, 1917. Ann. Transvaal Mus., 6: 5. Type species: Platymops petrophilus Roberts, 1917.

Sauromys is a monotypic genus almost certainly found only in southern Africa. Simmons (2005) lists Ghana as a possible locality, and Meester et al. (1986) refer to an immature specimen from that country in the USNM collection. However, D. E. Wilson (pers. comm. in Jacobs & Fenton 2001) found that the USNM collection contained no specimens other than 38, all of which came from southern Africa. No records from Ghana are reported by Grubb et al. (1998). Sauromys is distinguished from other genera of molossids in Africa by the following combination

of characters: head, body and skull extremely flattened (as in Platymops); ears without complex folding and with inner margins separated by narrow gap (cf. ears widely separated in Platymops); no wart-like granulations on forearm; no gular sac (gular gland); upper lip smooth with few coarse bristles; third ridge of posterior upper molar (M3) equal in length to second ridge; dental formula: 1123/2123 = 30. The flight membranes, and bones of wing, hindlimb and tail, and the lateral, dorsal and ventral views of the skull are illustrated in Figure 110.

Figure 110. Sauromys petrophilus. Flight membranes, and bones of wing and hindlimbs (traced from a dried skin). Skull (ZFMK 77.489).

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Sauromys was described as a subgenus of Platymops, then raised to generic status by Peterson (1965). Subsequently, it has been considered a subgenus of Mormopterus by Freeman (1981) and others including Legendre (1984) and Koopman (1993, 1994). It is considered a distinct genus here, following Peterson (1965), Meester

et al. (1986), Corbet & Hill (1992), Peterson et al. (1995) and Simmons (2005). The only species is S. petrophilus. Meredith Happold

Sauromys petrophilus ROBERTS’S FLAT-HEADED BAT Fr. Molosse à tête plate de Roberts; Ger. Roberts Flachkopf-Bulldoggfledermaus Sauromys petrophilus (Roberts, 1917). Ann. Transvaal. Mus. 6: 4. Bleskop, near Rustenburg, Northwest Province, South Africa.

Taxonomy Originally Platymops (Sauromys) petrophilus. Synonyms: erongensis, fitzsimonsi, haagneri, umbratus. Subspecies: five of uncertain validity. Chromosome number (Namibia, South Africa): 2n = 48; aFN = 62. X = medium-sized submetacentric;Y = small acrocentric (Rautenbach et al. 1993).

and their validity is uncertain (Smithers 1983, Meester et al. 1986). No subspecies are recognized by Jacobs & Fenton (2002).

Description Very small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; medium-small for an African molossid; head and body extremely flattened; ears separated and without complex folds; dorsal pelage without stripes; forearms without wart-like granulations; anterior upper premolar well developed. Sexes similar in colour; "" on average slightly larger than !!. Pelage soft and dense; mid-dorsal hairs 5–6 mm. Dorsal pelage dark brown, medium yellowish-brown, greyish-brown, dark grey to pale grey; hairs brown with the various shades of brown or grey at tip. Ventral pelage fairly dark greyish-brown, pale grey to whitish; flanks sometimes darker than throat and abdomen; chin naked. Head extremely dorsoventrally flattened. Upper lip smooth and with comparatively few coarse bristles (not spoon-hairs). Ears moderately large but not extending beyond muzzle when laid forward, broad, without complex folds; inner margins separated by narrow gap (Figure 97e) (cf. Tadarida pumila with which it has been confused [Jacobs & Fenton 2002]). Antitragus indistinct. Tragus very small. No interaural crest. No gular gland. Wings long, narrow, blackish-brown, brown to semi-translucent greyish-brown. Skull (Figure 110) extremely dorsoventrally flattened (maximum height 32 [24–37]% of GLS, n = 22). Zygomatic arches slender. Anterior of braincase not elevated above plane of rostrum. Sagittal crest indistinct and depressed in shallow parietal groove; lambdoid crest moderate. Lachrymal ridges indistinct and forming a comparatively small tubercle on each side of rostrum (cf. Platymops setiger). Anterior palatal emargination wide; upper incisors well separated. Basisphenoid pits shallow (indistinct), large, their width much greater than their distance apart. Upper incisors with small secondary cusp on lateral side, just below gum. Anterior upper premolar about same height as cingulum of posterior premolar, within toothrow; canine and posterior premolar well separated (Figure 103c). M3 with second and third ridges equal in length. Two lower incisors on each side. Lower canines with cingula not greatly enlarged and not in contact. Dental formula: 1123/2123 = 30.

Platymops setiger. Ears well separated. Forearm with warty granulations. Upper lip with many wrinkles. Gular sac (gland) present, albeit small, in both sexes. Skull with lachrymal ridges very well developed and prominent. Anterior upper premolar absent (but sometimes a minute spicule, probably a milk tooth, is present in younger animals); canine and posterior premolar in contact or almost so. Mormopterus spp. Inner margins of ears widely separated in M. acetabulosus (but almost touching and joined by a slender band of skin in M. francoismoutoui). Head not extremely flattened. Anterior upper premolar absent. Myopterus daubentonii. Dorsal pelage usually with contrasting lateral bands of paler colour. Wings with arm-wing white, hand-wing transparent. M3 with third ridge absent. North of Equator. M. whitleyi. Inner margins of ears widely separated. Head not extremely flattened. Arm-wing yellowish-white; hand-wing transparent. North of 10° S.

Similar Species complex folds:

Four other African molossids have ears without

Distribution Endemic to Africa. Found in the Zambezian Woodland BZ south of Zambezi R., the South-West Arid BZ and the South-West Cape BZ. Recorded marginally in SW Angola (CrawfordCabral 1986), and in Botswana, Zimbabwe, W Mozambique and South Africa (Smithers 1983). In Zimbabwe, widespread across the S, E and NE regions below 1300 m, but inexplicably absent from large areas of semi-arid savanna woodland in N and W Zimbabwe where suitable day-roosts in granitic and Karoo sandstone outcrops appear abundant (F. P. D. Cotterill unpubl.). Meester et al. (1986) reported an immature specimen from Ghana (USNM) tentatively assigned to S. petrophilus, but subsequently this could not be traced (Jacobs & Fenton 2002). Specimens from Cederberg in South-West Cape BZ listed as Tadarida pumila by Meester et al. (1986) have been re-identified as S. petrophilus (Jacobs & Fenton 2001). Habitat Wetter and drier woodland savannas (including miombo, mopane and undifferentiated woodlands), shrublands, Acaciawooded grasslands and deserts; always in areas with rocky outcrops and hills, principally of Karoo sandstones and granitic intrusions.

GeographicVariation Five subspecies (which were provisionally retained by Peterson 1965) are recognized by Koopman (1994) and Simmons (2005): S. p. petrophilus, S. p. erongensis, S. p. fitzsimonsi, S. p. Abundance Locally abundant in areas where roosts are abundant, haagneri, S. p. umbratus. However, they are based on colour differences notably Mutoko district (NE Zimbabwe), Matobo Hills, Limpopo 486

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from 14 individuals indicated that the diet was predominantly hardshelled insects: Coleoptera (43%), Hemiptera (29%), Hymenoptera (18%). Soft-bodied prey (Lepidoptera, Diptera and Neuroptera) comprised ca. 10% of the diet (Jacobs & Fenton 2002). Echolocation Search-phase call-shape: shallow FM (no details available). Start-frequency 31.4–43.5 kHz; end-frequency 26.7– 30.9 kHz; bandwidth 3.9–14.7 kHz; call-duration 5.0–10.2 ms; peakfrequency 28.0–37.5 kHz (eight bats, 30 calls; Jacobs & Fenton 2002). Social and Reproductive Behaviour Mostly found roosting singly, in pairs or in groups of up to four (Smithers 1971, 1983). Group-members huddle closely together, but scatter if disturbed. Reproduction and Population Structure No conclusive information. In NE Zimbabwe, pregnant and lactating !! have been mist-netted in mid-Nov; !! examined during cool-dry season were neither visibly pregnant nor lactating. Predators, Parasites and Diseases No information. Conservation

Sauromys petrophilus

valley (SW Zimbabwe) and Cederberg (South-West Cape, South Africa). Elsewhere scarce. Adaptations Aspect ratio high; wing-loading medium (Jacobs & Fenton 2002). Based on wing morphology, predicted to fly fast with poor manoeuvrability. Scuttles and climbs very efficiently. By day, roosts under slabs of exfoliated granite or sandstone, and in narrow fissures; roosts located near ground level in precipices. One roost was shared with 18 lizards Platysaurus sp. (Rautenbach 1982). The extreme flattening of the head, and the ability to folddown the ears, enables S. petrophilus to squeeze into very narrow cracks and crevices. In Namibia, captive bats did not need to drink regularly (Roer 1970). Foraging and Food Based on wing morphology, predicted to forage by fast-hawking. Reported flying as high as 37 m (Irwin & O’Donnelly 1962 in Freeman 1981). Most often captured over open water (Jacobs & Fenton 2002), but possibly because individuals come down to drink. Based on craniodental morphology, Freeman (1981) predicted that this species would feed mainly on small, soft-bodied insects, but the stomachs of 11 individuals contained both moths, beetles and other insects (Freeman 1981). Analysis of faecal pellets

IUCN Category: Least Concern.

Measurements Sauromys petrophilus FA: 40.8 (37–50) mm, n = 76 WS (d): 251–275 mm, n = 4* TL: 104.2 (89–131) mm, n = 58 T: 38.4 (29–49) mm, n = 66 E: 16.8 (13–22) mm, n = 64 Tr: 3.3 (2–6) mm, n = 10 Tib: 11.6 (10–12) mm, n = 11 HF: 7.8 (6–10) mm, n = 24 WT: 10.7 (6–22) g, n = 55 GLS: 17.1 (16.0–17.8) mm, n = 53 GWS: 11.2 (10.5–11.7) mm, n = 33 C–M3: 6.3 (5.9–7.9) mm, n = 44 Throughout geographic range (NMZB, TM, ZFMK, Smithers 1971, Smithers & Wilson 1979, Jacobs & Fenton 2001) *Jacobs & Fenton 2002 Key References

Jacobs & Fenton 2002; Smithers 1983. F. P. D. Cotterill

GENUS Tadarida Tadarine Free-tailed Bats Tadarida Rafinesque, 1814. Precis. Som., p. 55. Type species: Cephalotes teniotis Rafinesque, 1814.

Tadarida (sensu lato) contains 43 species (Simmons 2005), of which ca. 27 species occur in Africa; the remainder are found in Madagascar, Mauritius, Aldabra, São Tomé, Príncipe, S Europe, India to Japan, Malay Archipeligo to Philippines, Australia, some Pacific islands and the New World. Some African species recognized as distinct by

Simmons (2005) are considered here to have subspecific status, and some species listed by Simmons (2005), and profiled here, probably contain more than one species. Since this profile was submitted, a new species has been described (as Mops [Xiphonycteris] bakarii) by Stanley (2008) (see Tadarida brachyptera for more information). This species is 487

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not included in tables and maps. African Tadarida are distinguished from other genera of molossids in Africa by the following combination of characters: head, body and skull not extremely flattened although low in some species; ears complexly folded, never whitish or translucent, inner margins meeting (or almost meeting) to form V-shaped valley or joined by band of skin (never widely separated or separated by a narrow gap); upper lip thick, usually with prominent vertical wrinkles and many spoon-hairs; forearms never with warty granulations; dental formula: 1113/2123 = 28, 1123/2123 = 30, or 1123/3123 = 32. The length of the third ridge of posterior M3 relative to the second ridge is highly variable within Tadarida. Of the 28 species found in Africa, one is a Palaearctic species found in North Africa, five are found mainly in forests, 11 mainly in savannas, seven in both forest and savanna habitats, three in savanna and arid habitats, and one only on Mafia I. Five species, T. brachyptera, T. condylura,

T. nigeriae, T. pumila and T. thersites often roost under the corrugated iron roofs of houses occupied by humans, and may well have become more abundant as a result of this adaptation. Although sometimes noisy and smelly, they probably play an important role in controlling insects including some that damage crops. The taxonomy and systematics of Tadarida is controversial. Many authors, mainly following Freeman (1981), consider the forms Chaerephon and Mops to be distinct genera, and Xiphonycteris a subgenus of Mops (e.g. Koopman 1993, 1994, Simmons 2005). Many others, including Legendre (1984), Meester et al. (1986), Corbet & Hill (1992) and Peterson et al. (1995) recognize these forms as subgenera of Tadarida. It is currently considered probable that Chaerephon, Mops and Tadarida (sensu stricto) are not monophyletic. Furthermore, the affinities of some species are unclear and their placement in these taxa depends on what diagnostic characters are

Table 20.  Key to the African species in the genus Tadarida (sensu lato). The traditional subgenera to which the species have been allocated are given, but the monophyly of these taxa, and their taxonomic status is uncertain. Ventral flank-stripe in contrasting colour = band of ventral pelage adjacent to wing membrane (sometimes extending onto wing) the colour of which contrasts with that of the flanks adjacent to the flank-stripe. Ears can be small (extending about half-way along muzzle when laid forward) or large (extending well beyond snout when laid forward). Plantar pad = raised pad on sole of foot. Depth of basisphenoid pits in Tadarida may be shallow, moderate or deep, and the relative size of the pits (i.e. width relative to width of septum between pits) may be small (pit noticeably narrower than septum), medium (pit about as wide as septum) or large (pit noticeably wider than septum). For additional characters, see similar species listed in species profiles. Inner margins of ears

Upper M3: size of ridge 3 in relation to ridge 2

Number of lower incisors on each side

Anterior palatal emargination

Ventral flank-stripe in contrasting colour (colour)

Almost meeting to form V-shaped valley

3 = 2

2

Wide

Present or absent (Whitish)

Meeting to form V-shaped valley.

3 = 2

3

Wide

Absent

Meeting to form V-shaped valley

3 > half 2

2

Wide

Present (White)

Meeting to form V-shaped valley

3 > half 2

2

Wide

Present (White, cream or yellowish)

Meeting to form V-shaped valley

3 > half 2

2

Wide

Absent

Separated by a large bulbous lobe projecting well over the forehead

3 > half 2

2

Closed

Absent

Separated by a flat lappet over a small, backward-opening pocket

3 ≥ half 2

2

Closed in adults

Joined by band

3 > half 2

2

Narrow

Joined by band

3 > half 2

2

Narrow

Absent

Joined by band

3 > half 2

2

Narrow

Absent

Joined by band

3 > half 2

1 or 2

Closed or narrow

Present or absent (White or whitish)

Joined by band

3 > half 2

1 or 2

Closed or narrow

Present or absent (White or whitish)

Present (White or whitish) Present (Whitish)

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given precedence. Pending revision and redefining of these taxa, we present all of the African species as members of Tadarida but give their traditional ‘subgeneric’ affinities in each profile. African members of the ‘subgenera’, as traditionally defined, can usually be distinguished by the following combinations of characters: subgenus Tadarida: inner margins of ears meeting or almost meeting to form a V-shaped valley; third ridge of posterior upper molar (M3) more than half length of second or equal in length; anterior palatal emargination wide. subgenus Chaerephon: inner margins of ears joined by band of skin (except in T. (C.) gallagheri and T. (C.) major); third ridge of posterior upper molar (M3) more than half length of second or equal in length; anterior palatal emargination closed (except T. (C.) ansorgei, T. (C.) bemmelini and T. (C.) bivittata in which it is narrow). subgenus Mops: inner margins of ears joined by band of skin (except perhaps T. (M) niangarae); third ridge of posterior upper molar (M3) less than half length of second or absent (except sometimes equal in T. (M). condylura); anterior palatal emargination closed. subgenus Xiphonycteris: inner margins of ears joined by band of skin; third ridge of posterior upper molar (M3) less than half length of second or absent; anterior palatal emargination narrow.

The African species of Tadarida (sensu lato) can be distinguished by the combination of characters in Table 20, but additional information under Similar Species in all profiles should be consulted in conjunction with this table-key. Many African molossids are very difficult to distinguish, and it is important to use cranial and dental characters as well as external characters. Many of the relevant diagnostic characters are illustrated in the profile of the family Molossidae. In the text, the species of Tadarida (sensu lato) are presented in alphabetical order irrespective of the traditional subgenera to which they have been allocated by some authors. Meredith Happold

Tadarida condylura.

FA (mm)

Wing membrane dorsal side (ventral side if different)

Depth of basisphenoid pits (relative size of pits)

Miscellaneous

Species

42–55

Brown or blackish-brown

Moderate (Medium)

Ears small; inner margins meeting on forehead No white spot between shoulder-blades Plantar pad present

T. (T.) aegyptiaca

56–64

Blackish

Moderate (Large)

Three lower incisors on each side is unique

T. (T.) teniotis

55–62

Greyish (Whitish)

Shallow to moderate (Medium)

56–61

Brown (Whitish)

Deep (Large)

60–67

Dark brown to almost black

Deep (Medium)

38

Greyish-black

39–46

Dark brown to blackish

41–48

Pale grey

43–48 46–51 34–40 32–39

Light reddish-brown or light grey Dark brown or light reddish-brown Usually white, sometimes pale brown or greyishbrown White or blackish-brown

Deep (Large) Shallow to moderate (Small to medium) Shallow (Medium) Moderate (Small) Moderate (Small to medium)

Ears large; inner margins meeting near snout White spot between shoulder-blades of fully mature adults Plantar pad present Ears small; inner margins meeting at base of forehead No white spot between shoulder-blades Plantar pad present Ears small, inner margins meeting on forehead No white spot between shoulder-blades Plantar pad absent Ears large Rostrum uniquely with prominent nasal swellings

T. (T.) lobata

T. (T.) fulminans

T. (T.) ventralis T. (C.) gallagheri T. (C.) major

Uniquely with pair of tail-glands opening ventrally as a slit on each side of base of tail

T. (C.) bemmeleni

Dorsal pelage somewhat frosted, occasionally with white spots

T. (C.) ansorgei

White or whitish stripes and/or rows of spots on crown of most individuals, and sometimes on shoulders and flanks

T. (C.) bivittata

Moderate (Large)

Dorsal pelage pale grey, pale rusty-brown or medium greyish-brown !! with long bicoloured interaural crest

T. (C.) chapini

Moderate or shallow (Medium or small)

Dorsal pelage almost black, brown, greyish-brown or reddish-brown !! with short unicoloured interaural crest

T. (C.) pumila

continued overleaf 489

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Table 20. continued. Inner margins of ears

Upper M3: size of ridge 3 in relation to ridge 2

Number of lower incisors on each side

Anterior palatal emargination

Ventral flank-stripe in contrasting colour (colour)

Joined by band

3 > half 2

2

Closed

Absent

Joined by band

3 > half 2

2

Closed

Absent

Joined by band

3 > half 2

2

Closed

Present (White)

Joined by band

3 ˜ half 2

2

Closed

Present or absent (White)

Joined by band

3 < half 2

2

Narrow

Absent

Joined by band

3 < half 2

2

Narrow

Absent

Joined by band

3 < half 2

2

Narrow

Present but contrast slight (Black)

Joined by band

3 < half 2

2

Closed

Absent

Joined by band

3 absent or vestigial

2

Closed

Absent

Joined by band

3 absent

Usually 1, rarely 2

Narrow

Absent

Joined by band

3 absent

2

Narrow

Absent

Joined by band

3 absent

2

Closed

Absent

Joined by band

3 absent

2

Closed

Present but faint (Whitish or pale grey)

Joined by band

3 absent

2

Closed

Present (White or pale)

Uncertain, ? well separated

3 < half 2

2

Closed

Probably absent

Tadarida aegyptiaca EGYPTIAN FREE-TAILED BAT Fr. Tadaride d’Égypte; Ger. Ägyptische Bulldoggfledermaus Tadarida aegyptiaca (E. Geoffroy, 1818). Descrip. de L’Egypte 2: 128. Giza, Egypt.

Taxonomy Originally Nyctinomus aegyptiacus. Subgenus Tadarida. Synonyms in Africa: anchietae, bocagei, brunneus, geoffroyi, talpinus, tongaensis. Subspecies in Africa: two. Chromosome number (Kenya): 2n = 48; aFN = 54 (Nagorsen et al. 1976); (South Africa) 2n = 68 (Rautenbach et al. 1993). Description Small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral

membrane; medium-sized for an African molossid; ears of medium relative size, with inner margins almost meeting to form V-shaped valley; M3 with second and third ridges equal in length, anterior palatal emargination wide; wings dark; ventral flank-stripe in contrasting colour sometimes present; two lower incisors on each side. Sexes similar. Pelage short (mid-dorsal hairs 5–6 mm), sometimes glossy. Dorsal pelage grey, greyish-brown, reddish-brown, blackish-brown or black, often darker on head and flanks; no spots; hairs mostly

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FA (mm)

Wing membrane dorsal side (ventral side if different)

48–53

Blackish-brown

Depth of basisphenoid pits (relative size of pits)

Mostly dark brown, lighter towards tip Whitish or blackish (Whitish)

Deep (Large) Moderate (Large) Shallow to moderate (Small to medium)

45–51

Pale greyish-brown, paler towards tip

Shallow (Small)

32–35

Blackish-brown

Moderate (Small)

34–41

Various shades of brown

Moderate or shallow (Medium to large)

35–42

Blackish

Moderate (Small to medium)

51–55

Dark brown

54–58

Blackish-brown

27–30

Blackish

27–31

Blackish-brown, pale brown to whitish (Paler)

Shallow (Small)

44–48

Blackish-brown to pale grey

Moderate (Medium to large)

41–46

Medium brown to dark grey

Deep (Large)

59–67

Dark brown

ca. 52

Dark brown

42–46 41–51

Deep (Large) Deep (Large) Shallow (Small)

Moderate (Medium to large) Deep (Large)

Miscellaneous

Species

Sometimes with greyish spots dorsally

T. (C.) aloysiisabaudiae

No spots or stripes

T. (C.) russata T. (C.) nigeriae

Males without scent-glands between penis and anus Ventral pelage with little or no white Crown same colour as back Phalanges of 3rd and 4th fingers long (see profile) Ventral pelage pale Occipital helmet slight Ventral pelage pale Occipital helmet moderately prominent Wing insertion lower on body Ventral pelage dark Occipital helmet prominent Wing insertion higher on body Dorsal pelage medium sepia or yellowish-brown, or pale rusty-brown Crown same colour as back

T. (M.) condylura

T. (X.) petersoni T. (X.) brachyptera T. (X.) thersites T. (M.) trevori

Dorsal pelage dark brown to almost black; not frosted or flecked

T. (M.) congica

Lower canines with greatly enlarged cingula, especially in !!

T. (X.) spurrelli

Lower canines without greatly enlarged cingula Upper incisors procumbent

T. (X.) nanula

Males without scent-glands between penis and anus Ventral pelage predominantly white or cream Crown darker than back Phalanges of 3rd and 4th fingers of medium length (see profile) Males with paired glands between penis and anus Ventral pelage white or very pale Crown darker than back Phalanges of 3rd and 4th fingers short (see profile)

T. (M.) niveiventer

T. (M.) demonstrator

Dorsal pelage colour variable; frosted and flecked

T. (M.) midas

This combination of characters appears to be unique

T. (M.) niangarae

unicoloured but some have a slightly paler tip giving pelage a silvered or frosted appearance in some individuals. Ventral pelage slightly paler mid-ventrally, becoming same as dorsal colour on flanks, sometimes grizzled; no mid-ventral markings; ventral flank-stripe same as dorsal colour or distinctly paler or, exceptionally, white. Orange-phase not known. Head not extremely flattened. Upper lip with ca. five welldefined wrinkles on each side and comparatively few spoon-hairs. Ears blackish-brown, of medium relative length (just reaching tip of snout when laid forward); inner margins almost meeting on muzzle to form V-shaped valley.Tragus large, subrectangular, not concealed by antitragus. Antitragus ca. twice size of tragus, roughly semi-circular. No interaural crest. Gular gland present in both sexes. Wings and interfemoral membrane brown and semi-translucent, or blackish-

brown.Ventral sides of forearms and legs naked and whitish. Foot with raised pad on sole (plantar pad). Skull low but not extremely dorsoventrally flattened. Anterior of braincase not elevated (or only very slightly elevated) above plane of rostrum (as in Figure 99h). Sagittal crest absent or poorly developed; lambdoid crest poorly developed. Anterior palatal emargination wide. Interdental palate tapering slightly but evenly towards incisors. Basisphenoid pits moderate in depth, their width ca. equal to their distance apart. Anterior upper premolar ca. same height as cingulum of posterior premolar or lower; within toothrow. M3 with second and third ridges equal in length. Two lower incisors on each side, bicuspid, crowded. Lower canines with cingula not greatly enlarged, not in contact. Dental formula: 1123/2123 = 30. 491

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Geographic Variation Two subspecies occur in Africa (Koopman 1994): T. a. aegyptiaca: African range excluding W Zambia, Angola, Namibia. Slightly larger; paler. T. a. bocagei:W Zambia to Angola and Namibia. Slightly smaller; darker. Similar Species Four other Tadarida in Africa have ears with inner margins meeting, or almost meeting, to form a V-shaped valley: Tadarida fulminans. Larger (FA: 56–61 mm; GLS: 21.6–24.4 mm). Wings ventrally whitish. Ventral pelage with a white, cream or yellowish mid-ventral band and ventral flank-stripe. T. lobata. Usually larger (FA: 55–62 mm; GLS: 22.3–23.6 mm). Inner margins of ears meeting close to snout; ears extending well beyond snout when laid forward. White spot between shoulderblades of adults. T. teniotis. Three lower incisors on each side. Larger (FA: 58–64 mm; GLS: 22.5–24.8 mm). North Africa. T. ventralis. Much larger (FA: 60–67 mm; GLS: 23.4–26.1 mm). Distribution In Africa, recorded disjunctly from Nile Delta, Red Sea coast and scattered inland localities in the Sahara Arid BZ, from a few scattered localities in the Sudan Savanna BZ, and from one locality in the Rainforest BZ, but mostly recorded from Somalia– Masai Bushland, Coastal Forest Mosaic, Zambezian Woodland, Highveld, South-West Arid, South-West Cape and Afromontane– Afroalpine BZs. Recorded from Morocco, Algeria, Egypt, Niger, Nigeria, Sudan, Ethiopia, DR Congo and Kenya, and all countries southwards except, as yet, Malawi. Extralimitally: Saudi Arabia, Yemen and Oman and beyond to Afghanistan, India and Sri Lanka. Mapped from country checklists (see order Chiroptera), other literature and museum records. Subsequently recorded from E Uganda (Thorn & Kerbis Peterhans 2009): not mapped.

Habitat In Africa, mostly recorded from open woodland and bushland savannas (including undifferentiated woodlands, Acacia– Commiphora bushland, miombo and mopane woodlands), arid scrublands, open grasslands and forest-dominated valley bushveld (Smithers 1983, Taylor, P. 1998). Also found in some very arid areas but probably only where drinking water, insects and suitable day-roosts are available; e.g. in arid Botswana, associated with waterholes, bore-holes, reservoirs and temporary pools (Smithers 1983). Frequently occupies built-up areas (Taylor, P. 1998, R. T. F. Bernard pers. obs.). The record in rainforest is probably that of a vagrant. Abundance Abundant in some parts of geographic range, particularly in southern Africa where day-roosts may contain thousands of individuals (R. T. F. Bernard pers. obs.). Apparently restricted to a very limited range of localities in Angola and Zambia (Smithers 1983). Adaptations Aspect ratio very high; wing-loading medium, lower than in most molossids (Norberg & Rayner 1987). Predicted to fly more slowly than most molossids (Norberg & Rayner 1987). Scuttles and climbs very adeptly over horizontal to vertical surfaces. Day-roosts include narrow horizontal and vertical crevices in rockfaces and buildings, cracks under exfoliating rocks, behind bark of Acacia trees, cracks in tree-trunks, hollow trees, roofs of houses and churches (often between tiles and insulation or between tiles and rafters), crevices and crannies within caves (Shortridge 1934, Rautenbach 1982, Smithers 1983, Taylor, P. 1998). Clings in contact with substrate, usually tucked into narrow cracks and crevices. Occasionally reported roosting with T. pumila, once with both T. pumila and Otomops martiensseni, and once with Pipistrellus capensis (Herselman & Norton 1985, Taylor, P. 1998). Can tolerate extreme roost temperatures, e.g. under corrugated iron roof in South Africa, below zero in winter to above 45 °C in summer (R. T. F. Bernard unpubl.). There is no evidence, from observations in laboratory and field, that this species enters torpor or hibernation (Bernard unpubl.). Specimens from Western Cape, South Africa, captured late Jan to early Feb, had yellow fat at base of ribs and hips, suggesting an increase in body fats before winter (Seamark 2005). Foraging and Food Forages by fast-hawking. Observed flying high, and also skimming over water to pick up water-beetles and other aquatic insects or to sip water (Shortridge 1934, Smithers 1971). Based on remains in faeces, diet includes beetles and moths (Fenton & Thomas 1980, Fenton 1985).

Tadarida aegyptiaca

Echolocation Call-shape variable. Search-phase call-shape: quasi-linear shallow FM. Search-phase (Zimbabwe): start-frequency 26 kHz; end-frequency 15 kHz; peak-frequency 18 kHz; maximum call-duration 15 ms; intensity high (five bats, flying in open; Fenton & Bell 1981). Search-phase (South Africa): start-frequency (mean ± S.D.) 23.2 ± 1.9 kHz; end-frequency 18.7 ± 1.3 kHz; callduration 7.0 ± 2.6 ms, sometimes up to 12 ms; energy mainly in fundamental harmonic but up to three harmonics may be present (three bats, flying in open; Taylor 1999a, 2000).When scuttling, callshape similar but steeper; start-frequency ca. 29 kHz; end-frequency ca. 18.6 kHz; duration ca. 5.5 ms (Taylor 1999a).

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Tadarida aloysiisabaudiae

Social and Reproductive Behaviour Occasionally roosts singly or in pairs, more often in groups of up to 50 or more, and sometimes in colonies of several hundreds (Shortridge 1934, Rautenbach 1982, Taylor, P. 1998) and even thousands (R. T. F. Bernard pers. obs.). Much noisy squeaking and chirping (audible to humans) occurs in the roosts prior to emergence of these bats at dusk. Maternity colonies are formed in summer in South Africa (Taylor 2000, R. T. F. Bernard pers. obs.).

S. namru (Hemiptera: Cimicidae); fleas Araeopsylla gestroi, A. scitula, A. wassifi, Chiropteropsylla brockmani (Siphonaptera: Ischnopsyllidae); ticks Carios vespertilionis, C. boueti, C. confusus (Acari: Argasidae); and mites Parasteatonyssus nyctinomi, P. cornutus (Acari: Macronyssidae), Microtrombicula kanyei (Acari: Trombiculidae) (Anciaux de Faveaux 1984).

Reproduction and Population Structure Litter-size: one. At ca. 33° S in South Africa, the reproductive chronology is restricted seasonal monoestry with copulation in Aug and parturition in early summer (Nov–Dec) after 4-month gestation (Bernard & Tsita 1995). Right ovary and right uterine horn significantly larger than on left; corpora lutea in right ovary only; pregnancies in right uterine horn only. Climate of study area is strongly seasonal, with a single hot and typically wetter season from Oct–Apr, and a cool, drier season from May–Sep. Monoestry at this high latitude is probably explained by the comparatively long gestation and the relatively short period during which temperatures are high enough to ensure abundant insects. In contrast, at 24° S where temperatures are higher for longer, T. pumila and T. condylura have shorter gestations and are polyoestrous (see profiles). There appear to be no conclusive data for T. aegyptiaca elsewhere in Africa. In Zimbabwe, pregnant "" were caught in Nov and lactating "" in Dec (no data given for other months) (Smithers & Wilson 1979). In South Africa, !! apparently reach sexual maturity in second year; "" in first year.

Measurements Tadarida aegyptiaca FA: 47.6 (42–55) mm, n = 97 WS: n. d. TL: 109.8 (91–131) mm, n = 85 T: 39.7 (30–50) mm, n = 83 E: 19.4 (16–26) mm, n = 78 Tr: n. d. Tib: 13, 14 mm, n = 2 HF: 9.5 (7–12) mm, n = 77 WT: 14.7 (9–22) g, n = 53 GLS: 18.7 (17.1–21.9) mm, n = 68 GWS: 11.4 (10.7–13.5) mm, n = 60 C–M3: 6.7 (6.0–8.2) mm, n = 62 Egypt, Sudan, Angola, Namibia, Botswana, Zimbabwe, Swaziland, South Africa (AM, TM, ZFMK and literature)

Predators, Parasites and Diseases Remains have been found in pellets of Barn Owls Tyto alba in the West Coast N. P., South Africa (Avery 1992). Ectoparasites include bed-bugs Stricticimex transversus,

Conservation

Key References P. 1998.

IUCN Category: Least Concern.

Bernard & Tsita 1995; Smithers 1971; Taylor, Ric T. F. Bernard & Meredith Happold

Tadarida aloysiisabaudiae DUKE OF ABRUZZI’S FREE-TAILED BAT Fr. Tadaride d’Aloys Sabaud; Ger. Fürst von Abruzzen Bulldoggfledermaus Tadarida aloysiisabaudiae (Festa, 1907). Boll. Mus. Zool. Anat. Comp. Univ. Torino 22 (546): 1. Toro, W Uganda.

Taxonomy Originally Nyctinomus Aloysii-Sabaudiae. Subgenus Chaerephon. Synonyms: cyclotis. Subspecies: none recognized here (see Geographic Variation). Chromosome number (Cameroon): 2n = 48; aFN = 66 (Smith et al. 1986). The name aloysiisabaudiae is a latinization of one of the names and titles (Luigi di Savoia) of the Duke of the Abruzzi who led the Italian Rwenzori expedition during which this bat was first collected. Savoy was known to the Romans as Sabaudia (Hayman & Hill 1971). Considered to be closely related to T. russata by Peterson (1967, 1969, 1971a) and Fenton & Peterson (1972). Possibly also closely related to T. bivittata and T. ansorgei (Freeman 1981). Description Small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; medium-sized for an African molossid; ears joined by interaural band; M3 with third ridge > half length of second ridge; anterior palatal emargination closed; wings blackish-brown; ventral flank-stripe not in contrasting colour; no tail-glands; rostrum uniquely narrow and elongated; FA: 48–53 mm. Sexes similar.

Pelage short (mid-dorsal hairs ca. 4 mm), sparse on nape; dorsal pelage extending over rump and onto interfemoral membrane. Dorsal pelage chocolate brown to dark rusty-brown (orange-phase: bright rusty-orange), sometimes with small, diffuse whitish spots or scattered whitish hairs; all dorsal hairs pale at base. Ventral pelage with flanks dark brown, belly similar to dorsal pelage but paler and suffused with greyish-fawn; no mid-ventral markings; ventral flankstripe same colour as flanks. Head not extremely flattened; dorsal surface of snout inconspicuously covered with short, stiff bristle-like hairs. Upper lip with 8–9 well-defined wrinkles on each side and many spoon-hairs. Ears blackish-brown, broad, not quite reaching snout when laid forward; inner margins joined by interaural band across forehead, which projects forward as a knob-like protuberance (containing a backward-opening interaural pocket) that almost covers the snout. Tragus small, tip bluntly pointed, concealed by antitragus. Antitragus well developed, roughly semi-circular. Interaural crest of dark brown hairs arising from interaural pocket in both sexes. No gular gland. Wings and interfemoral membrane blackish-brown. For additional information, see Lanza & Harrison (1963). 493

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Skull large and elongated with a long, narrow rostrum; not extremely dorsoventrally flattened. Anterior of braincase slightly but distinctly elevated above plane of rostrum. Sagittal and lambdoid crests weakly developed or absent. Anterior palatal emargination closed; two foramina, and usually also a third, median foramen behind the incisors. Basisphenoid pits deep, large, their width more than twice their distance apart. Anterior upper premolar distinctly taller than cingulum of posterior premolar, within toothrow; canine and posterior upper premolar well separated. M3 with third ridge almost as long as second ridge. Two lower incisors on each side; bicuspid, crowded. Lower canines with cingula not greatly enlarged, not in contact. Anterior lower premolar ca. equal in size to posterior premolar in !!, but smaller than posterior premolar in "". Dental formula: 1123/2123 = 30. Geographic Variation None recorded but data are limited. Grubb et al. (1998) provisionally listed cyclotis Brosset, 1966 as a subspecies, but this view is not held by Hayman & Hill (1971), Fenton & Peterson (1972) and Simmons (2005), and is not held here. Similar Species Four other African Tadarida have the following combination of characters: ears joined by interaural band; M3 with third ridge > half length of second; wings neither white nor whitish; no ventral flank-stripe in contrasting colour; no tail-glands (Table 20, p. 488): Tadarida ansorgei. Smaller (FA: 43–48 mm; GLS: 18.6–20.5 mm). Wings semi-translucent light reddish-brown to light grey. Interaural band comparatively less prominent and with V-shaped fold in middle. Skull with anterior palatal emargination narrow (bulb-shaped). Basisphenoid pits moderate in depth, small, their width less than their distance apart. T. bivittata. Often with white spots and sometimes short stripes on head and neck; dorsal pelage not extending on to interfemoral membrane. Ears on average smaller; not reaching snout when laid forward. Interaural band comparatively less prominent and with V-shaped fold in middle. Skull with anterior palatal emargination narrow (bulb-shaped). Basisphenoid pits moderate in depth, small to medium-sized, their width ≤ their distance apart. T. russata. Smaller (FA: 42–46 mm). Skull shorter (GLS: 17.8– 19.4 mm) and narrower (GWS: 10.5–11.4 mm). Otherwise very similar in pelage pattern and colour as well as in external characters. T. pumila (sometimes). Much smaller (FA: 32–39 mm; GLS: 14.2– 17.6 mm). Distribution Endemic to Africa. Recorded mainly from Rainforest BZ (Western,West Central and East Central Regions) and adjacent Northern and Eastern Rainforest–Savanna Mosaics, with one record very marginally in Sudan Savanna BZ on the upper Mountain Nile R. (Bahr al Jabal) in S Sudan. Known from 16 localities in Côte d’Ivoire, Ghana, Cameroon, Gabon, Central African Republic (Boukoko, MNHN), DR Congo, Sudan (Juba, SMNS) and Uganda (Brosset 1966, Peterson 1967, 1969, 1972, Fenton & Peterson 1972, Smith et al. 1986, Beaucournu & Fahr 2003, J. Fahr unpubl.). Habitat Mainly recorded from lowland rainforest, montane forest flanking the Albertine Rift Valley, swamp forest, and rainforest–

Tadarida aloysiisabaudiae

savanna mosaic adjacent to rainforest; also from one locality in flooded grassland (Juba, Sudan). One specimen was taken in a clearing within forest in Gabon, one from cutover rainforest in Ghana, others from ‘Guinea woodland’ in Ghana and Uganda, and others appear to have been taken in forested areas but without further details (Fenton & Peterson 1972). Fenton & Peterson (1972) suggested that this species ‘may be associated with a forest-edge or semi-open forest habitat rather than with either dense high forest or with open forest or savannas’, but this needs confirmation because it has now been captured within closed primary rainforest (J. Fahr unpubl.). Mainly recorded from lower altitudes, with 1200 m as the known maximum (Fenton & Peterson 1972). Individuals were mostly captured with mist-nets set over water or in forest clearings (Brosset 1966, Fenton & Peterson 1972, J. Fahr unpubl.). Abundance Not known. Appears to be quite rare but surveys with elevated mist-nets are needed to determine accurately both the abundance and distribution of this species. Remarks In Garamba N. P., NE DR Congo, one ! was taken from a fissure in the trunk of an Isoberlinia doka tree that was also inhabited by a colony of ca. 150 T. condylura (Verschuren 1957 as T. (Mops) trevori but re-identified by Peterson 1972). Like most other molossids, mainly forages by fast-hawking in open spaces above the vegetation. In forest-savanna vegetation in Comoé N. P., Côte d’Ivoire, the median height of captures (in mist-nets set 0–25 m above ground) was 17.0 (12.4–19.6) m (n = 5). In rainforest in Taï N. P., Côte d’Ivoire, two individuals were mist-netted between 1–3 m over small creeks, but they had probably descended to drink. Predators, Parasites and Diseases Ectoparasites include a flea Lagaropsylla senckenbergiana (Siphonaptera: Ischnopsyllidae) (Beaucournu & Fahr 2003).

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Tadarida ansorgei

Conservation IUCN Category: Least Concern. Only 16 localities; seems comparatively rare. Major threats: loss and degradation of habitat mainly because of logging and agriculture. Population trend inferred to be declining. Measurements Tadarida aloysiisabaudiae FA: 51.5 (48–53) mm, n = 18 WS (c): 383 (343–401) mm, n = 6 TL: 118.8 (108–132) mm, n = 19 T: 40.1 (34–46) mm, n = 19 E: 22.0 (19–25) mm, n = 17 Tr: 1.5, 1.8 mm, n = 2 Tib: 18.1 (16–20) mm, n = 13

HF: 12.3 (10–14) mm, n = 17 WT: 26.1 (18–38) g, n = 13 GLS: 21.4 (20.0–22.3) mm, n = 18 GWS: 12.5 (11.7–13.1) mm, n = 18 C–M3: 7.8 (7.5–8.2) mm, n = 17 Côte d’Ivoire, Ghana, Gabon, DR Congo, Uganda (AMNH, FC, IRSN, LACM, MNHN [holotype cyclotis], MZUT [holotype alyosiisabaudiae], ROM, SMF, USNM) Key References Brosset 1966; Fenton & Peterson 1972; Lanza & Harrison 1963; Peterson 1967, 1969. Jakob Fahr

Tadarida ansorgei ANSORGE’S FREE-TAILED BAT Fr. Tadaride d’Ansorge; Ger. Ansorges Bulldoggfledermaus Tadarida ansorgei (Thomas, 1913). Ann. Mag. Nat. Hist., ser. 8, 11: 318. Malanje [= Malange], Angola.

Taxonomy Originally Nyctinomus ansorgei. Subgenus: placed in Chaerephon (e.g. by Koopman 1994 and Simmons 2005) but previously placed in subgenus Tadarida by some authors (e.g. Hayman & Hill 1971). Synonyms: rhodesiae. Subspecies: none. Appears closely related to the often sympatric T. bivittata (Eger & Peterson 1979), but the ansorgei/bivittata complex needs further revision (see profile of T. bivittata). Tadarida bivittata currently appears to contain a form that resembles T. ansorgei, and some information in the literature attributed to T. bivittata probably refers to T. ansorgei, and vice versa. Furthermore, museum collections probably contain misidentified specimens. Chromosome number (Cameroon): 2n = 48; aFN = 66. X = medium-sized subtelocentric, Y = small acrocentric (Smith et al. 1986). In South Africa: 2n = 48; aFN = 68. X = medium metacentric, Y = small acrocentric (Rautenbach et al. 1993). Description Small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; medium-sized for an African molossid; ears joined by interaural band; M3 with third ridge > half length of second ridge; anterior palatal emargination narrow; dorsal pelage dark greyish-brown to reddish-brown (occasionally spotted); wings light grey or reddish-brown; no ventral flank-stripe in contrasting colour; no tail-glands. Not easily distinguished from T. bivittata. Sexes similar in colour; !! slightly larger on average than "". Pelage short (mid-dorsal hairs ca. 4 mm), soft, dense, sparse on crown behind junction of ears and sometimes sparse on nape; terminates abruptly on rump. Dorsal pelage dark greyish-brown to chocolate brown or reddish-brown, somewhat frosted and occasionally with white spots; hairs unicoloured or with pale grey at tip. Based on museum specimens, greyish-brown individuals more common (cf. T. bivittata). Ventral pelage paler than dorsal pelage except on throat, which is blackish to brown, conspicuously darker and denser than elsewhere, especially in !!. Chin naked, in marked contrast with dense pelage on throat. No mid-ventral markings; no ventral flank-stripe in contrasting colour; pelage not extending onto wing-

membrane. Head not extremely flattened. Upper lip with 6–7 welldefined wrinkles on each side and many spoon-hairs. Ears pale-grey to reddish-brown, reaching ca. half-way along muzzle when laid forward; inner margins joined by interaural band with V-shaped fold in middle. Tragus very small, subquadrangular; antitragus much larger and roughly trapezoid (widest at base). Interaural crest of dark hairs (ca. 2 mm) in shallow pouch, in both sexes; denser and more conspicuous in !!. Gular gland: no information.Wings semitranslucent light reddish-brown to light grey dorsally and ventrally; interfemoral membrane slightly darker. Skull not extremely dorsoventrally flattened. Anterior of braincase only slightly elevated above plane of rostrum (frontals inflated dorsally into a very shallow dome, Figure 99e) (cf. T. bivittata). Mastoid width: 10.8 (10.2–11.2) mm, n = 139 (sexes alike) (cf. T. bivittata) (Eger & Peterson 1979). Sagittal crest distinct although very low, and usually complete (extending unbroken across whole braincase), but in 7% of 60 specimens from DR Congo (RMCA) the crest does not extend across the posterior third of the cranium, and in 3%, it is very indistinct over this section. Lambdoid crests moderate. Anterior palatal emargination narrow, bulb-shaped (narrowest between incisors). Basisphenoid pits moderate in depth, small, their width less than their distance apart. Anterior upper premolar ca. same height as cingulum of posterior premolar, within toothrow; canine and posterior premolar separated. M3 with third ridge almost as long as second ridge.Two lower incisors on each side, bicuspid, crowded. Lower canines with cingula not greatly enlarged, usually almost in contact, occasionally in contact. Dental formula: 1123 /2123 = 30. Geographic Variation

None recorded.

Similar Species Four other African Tadarida have the following combination of characters: ears joined by interaural band; M3 with third ridge > half length of second; wings neither white nor whitish, no ventral flank-stripe in contrasting colour; no tail-glands (Table 20, p. 488): 495

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Family MOLOSSIDAE

woodland savanna in the vicinity of rugged hills and mountain ranges with rock cliffs and precipices. Abundance Locally common in Zimbabwe; some roosts accommodate hundreds of individuals.

Tadarida ansorgei

Tadarida aloysiisabaudiae. Larger (FA: 48–53 mm; GLS: 20.0– 22.3 mm). Wings blackish-brown. Interaural band comparatively prominent, with knob-like projection almost covering snout. Skull with anterior palatal emargination closed. Basisphenoid pits deep, large, their width more than twice their distance apart. T. russata. Dorsal pelage uniformly dark rusty-brown to sepia brown; no grizzling, no white flecks or spots; hairs with paler bases. Skull with anterior of braincase moderately elevated above plane of rostrum. Anterior palatal emargination closed. Basisphenoid pits moderate in depth, separated by narrow bony ridge (< half breadth of one pit). T. bivittata. Dorsal pelage often (but not always) with white stripes or rows of spots on head and neck. Wings dark brown or reddishbrown. Ears broader and dorsally browner. More robustly built (see weights). Skull with anterior of braincase moderately elevated above plane of rostrum. Mastoid width greater. Sagittal crest sometimes incomplete. A classification coefficient for distinguishing T. bivittata is given by Eger & Peterson (1979). T. pumila (sometimes). Much smaller (FA: 32–39 mm; GLS: 14.2– 17.6 mm). Distribution Endemic to Africa. Mainly recorded from Eastern Rainforest–Savanna Mosaic, Somalia–Masai Bushland BZ and Afromontane–Afroalpine BZs in NE DR Congo, S Sudan, Ethiopia and East Africa, and from the Zambezian Woodland BZ from SE DR Congo through Zambia and Zimbabwe to KwaZulu–Natal (South Africa). There is also an apparently isolated record from Guinea Savanna BZ in Côte d’Ivoire and another in Southern Rainforest– Savanna Mosaic in N Angola (Eger & Peterson 1979, Hill 1983, Nikolaus & Dowsett 1989, Bouchard 2001). Habitat Mainly woodland savannas (including Isoberlinia woodland, Acacia–Commiphora bushland and miombo woodland), and montane habitats (no details available). In Zimbabwe, recorded from dry

Adaptations Aspect ratio high to very high; wing-loading medium. Flight fast and agile. Scuttles rapidly; scrambles backwards into small crevices. Most often roosts by day in small crevices that are high up in cracks and clefts in cliffs and rocky hills; also roosts in crevices inside caves, mine-adits and occasionally buildings, and in expansion joints high up in concrete bridges. Near Faradje, NE DR Congo, a colony roosted, in total darkness, in a crack in a hill rising ca. 60 m above the surrounding bushveld (Lang & Chapin 1917b). The accumulation of 30 cm of guano in this day-roost indicated long-term residence. However, day-roosts in Zimbabwe are sometimes temporarily abandoned; departures do not appear related to seasonal changes, and it is not known if the bats move into nearby day-roosts, or migrate further afield (F. P. D. Cotterill unpubl.). In NE DR Congo, sometimes shares roosts with T.major (Verschuren 1957). In Zimbabwe, sometimes shares roosts with T. bivittata (flocks of both species emerge together) and T. fulminans (Cotterill & Fergusson 1993a). Foraging and Food Forages by fast-hawking. Flocks were observed leaving roost at dusk and flying fast to a considerable altitude (R. A. Fergusson pers. comm.). Stomachs of two individuals from DR Congo contained legs of beetles, wings of Formicidae (?) and antennae of Hymenoptera (?) (Verschuren 1957). Echolocation Search-phase call-shape (Zimbabwe): shallow FM. Start-frequency 28 kHz; end-frequency 16 kHz; bandwidth 16– 28 kHz; peak-frequency 17.8 kHz; maximum call-duration 15 ms (five bats flying in open; Fenton & Bell 1981). Social and Reproductive Behaviour In NE DR Congo, found roosting in groups (colonies) of no more than 30–40 individuals; !! and "" in approximately equal numbers; no segregation of sexes observed (Verschuren 1957). In Zimbabwe, usually roosts in colonies of up to many hundreds. Within roosts, groups of bats huddle tightly together in nooks and crannies, but are often restless and can be heard squeaking and jostling. Samples of bats taken from roosts in Zimbabwe contained more !! than "". Reproduction and Population Structure Litter-size: one (n = 10). At 16–18° S in Zimbabwe, the reproductive chronology appears to be seasonal bimodal polyoestry without postpartum oestrus, with births at beginning of hot-wet season (Nov) and at end of hot-wet season (Apr), but this needs confirmation.This is based on the following observations: ten "" pregnant and near term in Oct (crown–rump length of embryos 25 [21–29] mm); lactating "" found in Nov–Dec and also in Apr (no data for other months) (F. P. D. Cotterill unpubl.). Polyoestry is indicated by lactating "" collected in late Dec and mid-Jan, each of which had a fully developed Graafian follicle in the right ovary. Predators, Parasites and Diseases Skulls have been recovered from pellets of Barn Owls Tyto alba, and Mackinder’s Eagle-owls Bubo

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Tadarida bemmeleni

capensis have been observed hawking molossids emerging from their roosts in Zimbabwe (Cotterill 1992). Tadarida ansorgei is collected from its day-roosts and eaten by people in DR Congo (Lang & Chapin 1917b). Ectoparasites include a flea Lagaropsylla anciauxi (Siphonaptera: Ischnopsyllidae) and mites Chelanyssus spiniferus (Acari: Macronyssidae), Alabidocarpus molossicola (Acari: Chirodiscidae) (Anciaux de Faveaux 1984). Conservation

IUCN Category: Least Concern.

Measurements Tadarida ansorgei FA: 45.9 (43–48) mm, n = 199* WS (d): 342 (318–364) mm, n = 82 TL: 106 (99–119) mm, n = 87

T: 37 (32–46) mm, n = 86 E: 20.5 (15–23) mm, n = 87 Tr: n. d. Tib: 16.2 (15–17) mm, n = 12 HF: n. d. WT: 14.6 (10.5–24.5) g, n = 87 GLS: 19.6 (18.6–20.5) mm, n = 139* GWS: 11.6 (11.0–12.2) mm, n = 139* C–M3: 7.7 (7.3–8.2) mm, n = 139* Throughout geographic range (ROM) *Eger & Peterson 1979 Key References

Bouchard 2001; Eger & Peterson 1979. F. P. D. Cotterill

Tadarida bemmeleni GLAND-TAILED FREE-TAILED BAT Fr. Tadaride à glande caudale; Ger. Schwanzdrüsen-Bulldoggfledermaus Tadarida bemmeleni (Jentink, 1879). Notes Leiden Mus. 1: 125. Liberia (no specified locality).

Taxonomy Originally Nyctinomus bemmeleni. Sometimes incorrectly spelled bemmelini. Subgenus Chaerephon. Synonyms: cistura. Subspecies: two currently recognized (but see Geographic Variation). Possibly includes more than one species. The species was described from a specimen from Liberia. Later, Jentink (1888) stated that the specimen had been received in 1875 through the Zoological Garden Rotterdam, which led subsequent authors to question Liberia as the type locality. However, recent specimens from Liberia, Sierra Leone and Côte d’Ivoire support the Liberian origin of the holotype. According to Freeman (1981), T. bemmeleni is rather distinct and basal within Chaerephon and, together with T. ansorgei and T. bivittata, connects this subgenus with the subgenus Tadarida. Seems to be most closely related to T. bivittata (Peterson 1971b, Freeman 1981). Chromosome number (Kenya): 2n = 48; aFN = 54 (Nagorsen et al. 1976). Description Small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; medium-sized for an African molossid; ears joined by low interaural band, M3 with third ridge > half length of second; anterior palatal emargination narrow; wings pale grey, ventral flankstripe whitish; distinguished from other molossids by conspicuous tail-glands that open ventrally, one on each side of the base of the tail. Sexual dimorphism slight: on average, !! with longer second phalanx of third and fourth finger, longer skull and longer canines. Pelage short (mid-dorsal hairs ca. 4 mm); covering rump but not extending onto interfemoral membrane, which is clad only with very short, thin hairs. Dorsal pelage dark brown with slight frosting but no spots or stripes; hairs bicoloured with pale tip. Head same colour as dorsal pelage. Ventral pelage paler, flanks darker than belly; hairs pale brown with whitish or greyish tips; no mid-ventral markings; ventral flank-stripe whitish, contrasting with dark flanks. No black or darker pelage on throat. Head not extremely flattened. Upper lip with 6–8 well-defined wrinkles on each side and comparatively few spoon-hairs. Ears blackish-brown, very broad and squarish in

outline, reaching snout when laid forward; inner margins joined by low interaural band with V-shaped fold in middle. Tragus small and concealed by antitragus. Antitragus large. Interaural band without distinct crest, sometimes with dense hairs, sometimes almost naked. No gular gland. Wings and interfemoral membrane pale grey. Forearm and third metacarpal dorsally clad with very short, fine hairs, giving a velvety appearance. Both sexes with longitudinal sac-like tail-glands that open ventrally on either side of the tail just posterior to its junction with the body (Figure 111). Sometimes these glands are also dorsally visible as paired swellings, one on each side of the base of the tail. Skull not extremely dorsoventrally flattened. Anterior of braincase slightly to moderately raised above plane of rostrum. Sagittal crest weakly developed; lambdoid crest moderate. Anterior palatal emargination narrow. Basisphenoid pits shallow, their width ca. equal to their distance apart. Anterior upper premolar slightly higher than cingulum of posterior premolar, within toothrow. M3 with third ridge ca. three-quarters to almost same length as second. Two lower incisors on each side, slightly to clearly crowded. Lower canines with cingula not greatly enlarged, not in contact. Dental formula: 1123/2123 = 30. a

b

Figure 111. The unique tail-glands of Tadarida bemmeleni. (a) Dorsal view showing paired swellings associated with the glands, one on each side of the base of the tail. (b) Ventral view showing the two longitudinal openings of the glands. Based on Peterson (1971b).

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Geographic Variation Two subspecies are currently recognized (e.g. Peterson 1971b, Koopman 1975, Simmons 2005): T. b. bemmeleni: Guinea, Sierra Leone, Liberia, Côte d’Ivoire and Cameroon. On average smaller (FA: 43.8 [41–46] mm, n = 34; Tib: 13.3 [12–15] mm, n = 34; GLS: 17.3 [16.6–18.1] mm, n = 24). T. b. cistura: E DR Congo, S Sudan, SW Kenya and N Tanzania. On average larger (FA: 46.0 [43–48] mm, n = 9; Tib: 13.9 [13– 15] mm, n = 6; GLS: 18.3 [17.8–18.9] mm, n = 8). The apparent disjunction between the distributions of these taxa, together with differences in size (especially in FA, tibia and craniodental measurements), skull morphology (braincase higher, lachrymal breadth of rostrum greater and width across upper canines greater in T. b. cistura; see Peterson 1971b) and habitat preferences, suggest that these taxa might be distinct species, but further material, especially from West Africa, is needed to resolve this question. Similar Species No other African molossid has conspicuous saclike tail-glands that open ventrally, one on each side of the base of the tail (sometimes also visible dorsally as paired swellings). Distribution Endemic to Africa. Tadarida b. bemmeleni is known from only five localities in Rainforest BZ (Western and West Central Regions), Afromontane–Afroalpine BZ and Northern Rainforest– Savanna Mosaic. Recorded from Sierra Leone, Liberia, Côte d’Ivoire (Duékoué; Taï N. P.) and Cameroon (Peterson 1971b, J. Fahr unpubl.). A record from Marshall Territory near Harbel, Liberia, published by Bray (1958) as T. bemmelini, was later referred to T. brachyptera by Koopman (1989). Tadarida b. cistura is more widely known from ca. 17 localities in Eastern Rainforest–Savanna Mosaic and Somalia–Masai Bushland BZ in S Sudan, E DR Congo, Uganda, SW Kenya and N Tanzania (Peterson 1971b, J. Fahr unpubl.). Habitat Tadarida b. bemmeleni has been found in lowland rainforest and semi-deciduous forest (Liberia, Côte d’Ivoire), forest–savanna mosaic (Cameroon) and montane grassland (Mt Nimba). Recorded from lowlands to ca. 1600 m (Mt Nimba, Liberia; SMF). In Cameroon, individuals were mist-netted over a fast-running stream where it emerged from a forested area into a cleared area adjacent to a banana plantation (Peterson 1971b). One ! and one " were captured on consecutive nights, at 13.6 and 10.8 m above the ground, in elevated mist-nets set over the clearing of a research station within Taï N. P., Côte d’Ivoire (S. Pettersson pers. comm.). Tadarida b. cistura has been recorded from drier vegetation zones such as Acacia–Commiphora bushland, Isoberlinia woodland and forest– savanna mosaic. Most specimens have been caught in mist-nets over rivers and streams. One from Budongo Forest, Uganda, was mistnetted over a dam at the edge of a forest; another was taken from a hole in some rocks on the side of a gorge at 1600 m near Kampala, Uganda, and another from the slopes of Mt Kilimanjaro, Tanzania. Possibly rocky outcrops are required for roosting (Peterson 1971b). Abundance Uncertain. Apparently localized and probably fairly rare (especially the western subspecies), but data are not sufficient for accurate assessment.

Tadarida bemmeleni

Remarks Body mass appears to be relatively low for the size of this bat, possibly indicating a particular foraging strategy. Start (1969) found four individuals exhausted in a swimming pool: probably they became trapped after descending to drink from the pool. No other molossid in Africa has tail-glands: the cellular structure and function of these glands is not known. In Tanzania, one specimen was recovered from pellets of a Barn Owl Tyto alba (SMF). Ectoparasites include a mite Ewingana bispinosa (Acari: Myobiidae) (Anciaux de Faveaux 1984). Conservation IUCN Category: Least Concern. If T. b. bemmeleni and T. b. cistura are found to be distinct species (see Geographic Variation), the appropriate IUCN Category for T. bemmeleni would be ‘Near Threatened’ or ‘Data Deficient’ based on its limited area of occupancy, which is very fragmented and degraded. For T. cistura, ‘Least Concern’. Measurements Tadarida bemmeleni FA: 44.3 (41–48) mm, n = 43 WS (d): 330.3 (320–338) mm, n = 23* TL: 103.1 (94–110) mm, n = 39 T: 36.3 (34–40) mm, n = 41 E: 18.7 (14–20) mm, n = 41 Tr: n. d. Tib: 13.4 (12–15) mm, n = 40 HF: 11.1 (9–13) mm, n = 40 WT: 12.7 (11–14) g, n = 27 GLS: 17.5 (16.6–18.9) mm, n = 32 GWS: 10.4 (9.8–11.3) mm, n = 32 C–M3: 6.3 (6.0–6.8) mm, n = 36 Sierra Leone, Liberia, Côte d’Ivoire, Cameroon, Sudan, Kenya, Tanzania, Uganda (AMNH, BMNH, FC, HZM, MHNG, RMNH [holotype T. b. bemmeleni], ROM, SMF, USNM, Peterson 1971b)

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Tadarida bivittata

Samples include only 6–9 specimens of T. b. cistura (see also Geographic Variation) *T. b. bemmeleni only Key Reference

Peterson 1971b. Jakob Fahr

Tadarida bivittata SPOTTED FREE-TAILED BAT Fr. Tadaride tachetée; Ger. Gefleckte Bulldoggfledermaus Tadarida bivittata (Heuglin, 1861). Nouv. Act. Acad. Caes. Leop.-Carol., Halle 29 (8): 4, 13. Keren, Ethiopia.

Taxonomy Originally Nyctinomus bivittatus. Subgenus Chaerephon. Synonyms: none. Appears closely related to T. ansorgei (Eger & Peterson 1979). Contains more than one form. Heuglin described T. bivittata (on p. 13) from three umber brown specimens that have white markings (spots and/or stripes on head and neck) and dark brown wings and dark throats. As Heuglin did not designate a holotype, these specimens are considered to be syntypes (Hayman & Harrison 1966). Later (on p. 18), Heuglin referred to further specimens some of which are reddish-brown with almost no white markings (Hayman & Harrison 1966). In Zimbabwe, there appears to be a smaller, darkthroated form with lightly pigmented wings, and a larger, browner form, with more white markings and darker wings, which is not dark-throated; both are found in the same day-roosts, and both are sympatric with T. ansorgei in some areas (F. P. D. Cotterill pers. obs.). Although the relationship between T. bivittata and T. ansorgei was examined by Eger & Peterson (1979), it seems that the ansorgei/ bivittata complex is in need of further revision to clarify the diagnostic characters of T. ansorgei and T. bivittata, and to ascertain if T. bivittata is polymorphic and/or contains cryptic species. Pending revision, some of the information below needs confirmation. Furthermore, museum collections probably contain some misidentified material, and the literature probably contains references to misidentified material. Chromosome number: 2n = 48; aFN = 54. X = subtelocentric;Y = acrocentric (Peterson & Nagorsen 1975), but these authors reported difficulty in distinguishing between subtelocentric and acrocentric chromosomes. Description Information not applicable to the syntypes is in brackets and marked with *. A small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; medium-sized for an African molossid; ears joined by interaural band; M3 with third ridge > length of second ridge; anterior palatal emargination narrow; dorsal pelage brownish (or reddish*), with white spots or stripes on head and neck (sometimes absent*); wings dark brown (or light reddish-brown*); no ventral flank-stripe in contrasting colour; no tail-glands. Not easily distinguished from T. ansorgei. Males slightly larger on average than "". Pelage short (mid-dorsal hairs 3–4 mm), soft, dense; sparse on crown; dorsal pelage covering rump but not extending on to interfemoral membrane (cf. T. aloysiisabaudiae). Dorsal pelage umber brown (or dark reddish-brown, greyish-brown to blackish-brown*), with rows of white spots and sometimes short stripes on crown, neck and back (or with almost no such markings*). Based on museum

specimens, reddish-brown individuals appear to be more common (cf. T. ansorgei) (Eger & Peterson 1979). It is not yet clear if these variations in colour and markings indicate polymorphism or cryptic species or perhaps both (see Taxonomy). Ventral pelage brown, often slightly frosted (especially on belly) and with or without pale speckling; no mid-ventral markings; ventral flank-stripe same colour as flanks and not extending onto the wing-membrane. Throat much darker than chest and belly (or similar in colour*). (Dark-throated specimens sometimes have, and sometimes lack, white spots and/ or stripes*.) Head not extremely flattened. (Upper lip with 5–8 well-defined wrinkles on each side and many spoon-hairs [Freeman 1981, BMNH]*). (Ears light reddish-brown, reaching about half-way along muzzle; inner margins joined by interaural band with V-shaped fold in middle*.) (Tragus very small, concealed by antitragus*.) (Antitragus large, roughly rectangular*.) Males with interaural crest of very short (ca. 2 mm) dark hairs. (No gular gland*.) Wings and interfemoral membrane dark brown (or wings light reddish-brown and interfemoral membrane slightly darker*). Skull not extremely dorsoventrally flattened. Anterior of braincase moderately elevated above plane of rostrum (frontals inflated dorsally into a shallow dome, Figure 99d) (cf. T. ansorgei). (Mastoid width !!: 11.7 [11.3–12.2] mm, n = 71; mastoid width "": 11.5 [11.0–11.9] mm, n = 57 [cf. T. ansorgei] [Eger & Peterson 1979]*.) Sagittal crest very low (and sometimes incomplete [i.e. with gap near middle of braincase]*). (Lambdoid crests moderate*.) Palate with small anterior vacuities (Hayman & Harrison 1966). (Anterior palatal emargination narrow, bulb-shaped [narrowest between incisors]*.) Basisphenoid pits moderate in depth, small to medium-sized, their width ≤ their distance apart. Anterior upper premolar small (usually ca. same height as cingulum of posterior premolar but sometimes rising higher*), within toothrow; canine and posterior premolar well separated. M3 with third ridge almost as long as second ridge. (Two lower incisors on each side, bicuspid, crowded*.) (Lower canines with cingula not greatly enlarged, well separated to almost in contact*.) Dental formula: 1123/2123 = 30. Geographic Variation

None reported.

Similar Species Four other African Tadarida have the following combination of characters: ears joined by interaural band; M3 with third ridge > half length of second; wings neither white nor whitish; no ventral flank-stripe in contrasting colour; no tail-glands (Table 20, p. 488): 499

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Tadarida aloysiisabaudiae. Dorsal pelage only sometimes spotted; extending on to interfemoral membrane. Ears on average larger; almost reaching snout when laid forward. Interaural band comparatively prominent, with knob-like projection almost covering snout. Skull with anterior palatal emargination closed. Basisphenoid pits deep, large, their width more than twice their distance apart. T. russata. Smaller (FA: 42–46 mm; GLS: 17.8–19.4 mm). Skull with anterior palatal emargination closed, pair of incisive foramina visible. Basisphenoid pits moderate in depth, separated by narrow bony ridge (< half breadth of one pit). T. ansorgei. Dorsal pelage occasionally spotted, sometimes frosted, but not with rows of spots or stripes on head and neck. Wings semitranslucent light reddish-brown to light grey. Ears narrower and pale grey to off-white. Less robustly built (see weights). Anterior of braincase less obviously elevated above plain of rostrum. Mastoid width smaller. Sagittal crest complete. A classification coefficient for distinguishing T. ansorgei is given by Eger & Peterson (1979). T. pumila (sometimes). Much smaller (FA: 32–39 mm; GLS: 14.2– 17.6 mm). Distribution Endemic to Africa. Mainly recorded from Afromontane–Afroalpine, Somalia–Masai Bushland, Coastal Forest Mosaic and Zambezian Woodland BZs, with marginal records in Sahel Savanna and Sudan Savanna BZs and in Eastern Rainforest–Savanna Mosaic. Known from scattered localities on eastern side of Africa, from Eritrea and Ethiopia to Zimbabwe (contra Hutton 1986) and SW Mozambique (Koopman 1975, Eger & Peterson 1979, Smithers 1983, F. P. D. Cotterill unpubl.). Habitat Mainly savanna woodlands (including dry Acacia– Commiphora bushland, and Acacia woodland in East Africa, and

miombo woodland in the south), and montane habitats in Eritrea, Ethiopia and Kenya. Occurs peripherally to the Congo Basin, but evidently not within it. At least in Zimbabwe, typically found near exposed rocky outcrops of granite, basalt and sandstone, which provide day-roosts. Abundance Uncertain. In Zimbabwe, locally common in vicinity of day-roosts. At roosts shared with T. ansorgei and T. bivittata, T. bivittata is always captured in smaller numbers. Remarks Very little is known, and some information in literature refers to T. ansorgei. Wings long and narrow. Flight and cursorial locomotion as in other Tadarida. By day, roosts high up in rock crevices and occasionally in buildings. Not known to roost in mines (contra Smithers 1983 whose record refers to T. ansorgei). Roosts in small groups or colonies; individuals huddle together in nooks and crannies. In Zimbabwe, shares roosts with T. ansorgei, but samples captured as these bats emerged always contained fewer T. bivittata. In these samples, !! outnumbered "" but the reason for this is not known – perhaps !! emerge earlier than "". Litter-size: no information. In Zimbabwe, pregnant "" have been reported in Oct and Nov, and reproductively inactive parous "" have been reported in late Mar. Skulls have been recovered from pellets of Barn Owls Tyto alba found below the day-roosts of the bats, and Mackinder’s Eagleowls Bubo capensis have been observed hawking molossids emerging from these day-roosts in Zimbabwe (Cotterill 1992). Ectoparasites include a flea (Lagaropsylla anciauxi [Siphonaptera: Ischnopsyllidae]) (Beaucournu & Kock 1996). Conservation

IUCN Category: Least Concern.

Measurements Tadarida bivittata FA: 49.4 (46–51) mm, n = 131* WS (d): 363.2 (316–392) mm, n = 116 TL: 114.7 (106–129) mm, n = 136* T: 40 (32–48) mm, n = 134 E: 19.0 (15–22) mm, n = 130 Tr: n. d. Tib: n. d. HF: 10.6 (9–13) mm, n = 39† WT: 18.1 (15–32) g, n = 104 GLS: 20.5 (19.5–21.3) mm, n = 128* GWS: 12.4 (11.7–13.1) mm, n = 117* C–M3: 7.5 (7.0–8.0) mm, n = 128* Throughout geographic range (ROM) *Derived from Eger & Peterson 1979 †Hayman & Harrison 1966, Smithers & Wilson 1979 Key References 1966.

Eger & Peterson 1979; Hayman & Harrison F. P. D. Cotterill

Tadarida bivittata

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Tadarida brachyptera

Tadarida brachyptera SHORT-WINGED FREE-TAILED BAT Fr. Tadaride à ailes courtes; Ger. Kurzflügel-Bulldoggfledermaus Tadarida brachyptera (Peters, 1852). Reise nach Mossambique, Säugeth., p. 59. Mozambique I. (15° S, 40° E), Mozambique.

Taxonomy Originally Dysopes brachypterus. Subgenus: Xiphonycteris. Synonyms: leonis, ochraceus. Subspecies: two of uncertain validity. Apparently closely related to T. thersites and some authors suggest that these forms might be conspecific (e.g. Rosevear 1965, Hayman & Hill 1971). Some authors (e.g. Freeman 1981) consider leonis (including ochraceus) to be a distinct species, but El-Rayah (1980, 1981) and Simmons (2005) treat leonis as conspecific. Chromosome number (Cameroon): 2n = 48; aFN = 54; one pair large and three pairs medium metacentric and 19 pairs medium-small acrocentric autosomes (Smith et al. 1986). Since this profile was submitted, a molossid from Pemba I. Tanzania, was described as Mops (Xiphonycteris) bakarii (Stanley 2008). This new species resembles T. b. brachyptera in size (FA: 35.9 [34–38] mm, n = 21; GSL: 19.3 [18.0–21.1] mm, n = 13), but differs mainly in that the basisphenoid pits are essentially absent and the anterior upper premolar barely extends above the cingulum of the canine. The new species is larger than T. nanulus, T. petersoni and T. spurrelli. It differs from T. thersites in lacking basisphenoid pits and having a much more reduced anterior upper premolar. There is no profile for this species and it is not mentioned in other profiles or tables. Further details in Appendix. Description Very small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; small for an African molossid (FA: 34– 41 mm); ears joined by interaural band; M3 with third ridge present but < half length of second; anterior palatal emargination narrow, usually bulb-shaped; wings dark brown to black; ventral pelage pale, no ventral flank-stripe in contrasting colour; canines without greatly enlarged cingula. Not easily distinguished from T. thersites. Sexes similar. Pelage short (mid-dorsal hairs 3–4 mm), sometimes sparse on nape; dorsally there is either no band of naked skin, or only an inconspicuous narrow band of naked skin, adjacent to the flight-membranes (cf. wide band in T. thersites). Rump with two tufts of long hairs (as in several other species of Tadarida). Dorsal pelage blackish-brown, dark rustybrown, sepia brown, or greyish-brown; no grizzling, no white flecks or spots; hairs dark brown with paler base. Orange-phase: bright orangered specimens known from Cameroon. Ventral pelage pale brownishgrey, very pale grey, orange, yellowish-cream or white; mid-ventral marking inconspicuous or absent; ventral flank-stripe sepia brown or brownish-orange not contrasting with colour of flanks. Head not extremely flattened. Upper lip with 5–7 well-defined wrinkles on each side and many spoon-hairs. Ears blackish-brown, relatively small (not extending as far as snout when laid forward); inner margins joined by interaural band with V-shaped fold in middle. Tragus small. Antitragus large, subrectangular sometimes with corners very rounded. Interaural crest probably absent, but the posterior base of the interaural band often supports a tuft of dark brown, longer hairs. Gular gland: no information. Wings slightly translucent with heavy to light amounts of dark brown or black pigmentation. Wings inserting lower on body than in T. thersites. Interfemoral membrane dark brown.

Skull not extremely dorsoventrally flattened. Anterior of braincase elevated above plane of rostrum (more so in T. b. leonis than in T. b. brachyptera; Freeman 1981). Sagittal crest absent, weakly developed or moderate depending on age. Lambdoid crest very well developed in both subspecies; helmet not quite so prominent as in T. thersites but more prominent than in T. nanula, T. petersoni and T. spurrelli. Anterior palatal emargination narrow, usually bulb-shaped (narrowest between incisors). Basisphenoid pits of moderate depth (T. b. brachyptera) or shallow (T. b. leonis) (Freeman 1981); their width slightly greater to much greater than their distance apart. Upper incisor not procumbent. Upper canines with cingula not enlarged; viewed laterally, incisor not obscured by canine (cf. T. spurrelli). Anterior upper premolar distinctly taller than cingulum of posterior premolar, within toothrow or slightly displaced labially. M3 with third ridge much less than half length of second. Two lower incisors on each side, strongly bicuspid, crowded between canines. Lower canines long and robust; cingula not greatly enlarged, well separated. Dental formula: 1123/2123 = 30. Geographic Variation Two subspecies are recognized by Koopman (1994) and Simmons (2005). Koopman (1994) proposed the following distributions for these subspecies: T. b. leonis: Sierra Leone to E DR Congo. T. b. brachyptera: Uganda to Mozambique; Zanzibar I. The two widest gaps in the distribution of the species (see Map), especially that between the rainforest and coastal populations which is unlikely to be an artefact, do not seem to support the subspecific distributions proposed by Koopman (1994). If environmental factors (such as climate changes) have caused these two gaps, and if the gaps have resulted in subspeciation, it is more likely that there are three subspecies; leonis in the west, ochraceus in NE DR Congo and Uganda, and brachyptera in the Coastal Forest Mosaic BZ. However, this proposal needs further investigation. Similar Species Five other African Tadarida have the following combination of characters: ears joined by interaural band; M3 with third ridge absent or present but < half length of second; palatal emargination narrow; FA almost always < 40 mm (Table 20, p. 488): Tadarida nanula. Forearm shorter (27–31 mm). M3 with third ridge absent. Upper incisors procumbent (projecting in front of anterior faces of the canine cingula) (Figure 121). T. petersoni. Usually smaller (FA: 32–35 mm; GLS: 15.8–17.4 mm). Skull without well-developed lambdoid crest. Canines shorter. T. spurrelli. Forearm shorter (27–30 mm). M3 with third ridge absent. Canines with greatly enlarged cingula; in lateral view, upper incisors obscured by upper canines (Figures 121 and 122). T. thersites. Ventral pelage medium brown becoming very dark brown on flanks; ventral flank-stripe black. Dorsally, there 501

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the Congolian rainforest, although a series from Budongo Forest, Uganda was collected when this was primary forest (ROM, J. Eger pers. comm.). Also occurs in the East African coastal forest mosaic, but was not found in any of the coastal forests investigated by Cockle et al. (1998) except Kiwengoma Forest. Abundance Uncertain, but apparently common in some parts of geographic range.

Tadarida brachyptera

is a conspicuous, wide band of naked skin adjacent to flightmembranes. Wing insertion higher on body. Also, T. bakarii mentioned above in Taxonomy. Distribution Endemic to Africa. Recorded, disjunctly, from Rainforest BZ and marginally from the Northern Rainforest–Savanna Mosaic, from Sierra Leone to SW Central African Republic (except in Dahomey Gap) and from NE DR Congo to Uganda (Budongo and Bugala I.: not mapped). An apparently separate population occurs in the Coastal Forest Mosaic BZ from Kenya to Mozambique, including Zanzibar I. (Neumann 1900) and Mozambique I. Said to occur on Mafia I. by Simmons (2005), but not recorded there by Moreau & Pakenham (1940), Cockle et al. (1998) or Kock & Stanley (2009). Said to occur in Gambia by Koopman (1993), but not recorded west of Sierra Leone according to Grubb et al. (1998). Reports of this species on Bioko I. (e.g. Koopman 1994) are considered erroneous by Eisentraut (1973) and J. Juste (pers. comm.). Mapped from country checklists (see order Chiroptera), other literature and museum records. Habitat Considered mainly a lowland rainforest species in West Africa (Rosevear 1965, Koopman et al. 1995), but also occurs along the rainforest–savanna ecotone and in the mosaic of lowland rainforest and secondary grassland. Not observed in primary rainforest in Gabon (Brosset 1966) and possibly occurs mainly in clearings and areas of invasive woodland savanna in and around

Remarks Roosts by day in large hollow tree-trunks and hollow branches, deserted holes of barbets (Capitonidae) in dead trees, cracks in fabric of buildings and, very commonly, under corrugated iron roofs of houses. Large colonies are noisy and (to humans) pungently smelly; consequently regarded as pests. Sometimes roosts with T. thersites; less often with T. condylura. Recorded flying over water (Jones 1971). Known to eat winged termites (Lang & Chapin 1917b) but no other data available. Roosts in groups of 10–20 and in much larger colonies. Colonies reported to contain more "" than !! (Lang & Chapin 1917b). At 2–3° N in DR Congo, 14 of 18 "" had one medium-sized embryo (implanted in right uterine horn) in March (Lang & Chapin 1917b). Predators include Bat Hawks Macheiramphus alcinus (Chapin 1932). Conservation IUCN Category: Least Concern. If leonis is a distinct species, and even if it is retained as a subspecies, its conservation status should be assessed separately from that of T. brachyptera as it is probably much less common and widespread. Measurements Tadarida brachyptera FA: 37.3 (34–41) mm, n = 69 WS (d): 250 mm, n = 1 TL: 88.1 (78–100) mm, n = 64 T: 28.9 (25–35) mm, n = 66 E: 17.0 (14–19) mm, n = 65 Tr: n. d. Tib: n. d. HF: 11, 11 mm, n = 2 WT: 15.3 (12–18) g, n = 3 GLS: 17.9 (16.2–19.9) mm, n = 41 GWS: 11.3 (10.1–12.3) mm, n = 44 C–M3: 6.6 (6.1–7.7) mm, n = 49 Throughout geographic range (BMNH, HZM, RMCA, SMF, SMNS and literature) Key References Allen 1917a; El-Rayah 1980, 1981; Lang & Chapin 1917b; Rosevear 1965. Meredith Happold

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Tadarida chapini

Tadarida chapini PALE FREE-TAILED BAT (CHAPIN’S FREE-TAILED BAT, LONG-CRESTED FREE-TAILED BAT) Fr. Tadaride de Chapin; Ger. Chapins Bulldoggfledermaus Tadarida chapini (J. A. Allen, 1917). Bull. Amer. Mus. Nat. Hist. 37: 461. Faradje, Orientale, DR Congo.

Taxonomy Originally Chaerephon (Lophomops) chapini. Subgenus Chaerephon. Synonyms: lancasteri, shortridgei. Subspecies: three. Some authors consider shortridgei to be a distinct species (e.g. Peterson et al. 1995, Simmons 2005) but this view is not followed by Fenton & Eger (2002) and is not followed here. Chromosome number (Namibia, Zimbabwe): 2n = 48; aFN = 64 (Rautenbach et al. 1993). Description Very small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; small for an African molossid (FA: 34–40 mm); ears joined by interaural band; M3 with third ridge > half length of second ridge; anterior palatal emargination closed or slightly open; wings white (tinged with yellow) with minute black spots; ventral pelage pure white or pale greyish-brown; ventral flank-stripe white; !! with a long, bicoloured interaural crest. Sexes similar except for crest. Pelage short (mid-dorsal hairs 4–5 mm), soft; covering rump but not extending onto interfemoral membrane. Dorsal pelage pale grey, pale rusty-brown or medium greyish-brown; no grizzling, no white spots; patch of whitish hairs on crown; mid-dorsal hairs pale rusty-brown with off-white at base. Ventral pelage pure white or pale greyish-brown; mid-ventral markings absent or white, usually a broad band; ventral flank-stripe white, only sometimes contrasting with ventral pelage. Head not extremely flattened; forehead slightly elevated. Upper lip with 5–6 wrinkles on each side and many spoon-hairs. Ears brown; of

medium relative size (just reaching snout when laid forward); inner margins joined by interaural band with deep, backward-opening, pocket-like invagination in middle.Tragus minute, partly concealed by antitragus. Antitragus moderately large, roughly trapezoid with front-side and base widest. Adult !! with conspicuous long bicoloured interaural crest arising from interaural pocket (Figure 112); hairs 12–15 mm with basal half rusty-red or grey, terminal half white (Allen 1917a). Adult "" with short tuft of white hairs. No gular gland. Wings usually white becoming yellowish near body or almost transparent with little white pigmentation; armwing with many minute black specks; dorsal skin over bones dark brown. A few specimens from Kenya have pale brown to greyishbrown wings (Fenton & Eger 2002). Interfemoral membrane medium brown. Skull not extremely dorsoventrally flattened. Anterior of braincase slightly elevated above plane of rostrum. Sagittal crest weakly developed. Lambdoid crests weakly developed or absent dorsally, moderately developed laterally. Anterior palatal emargination variable, can be closed or slightly open and narrow. Basisphenoid pits, moderate in depth, large, their width ca. twice their distance apart. Anterior upper premolar distinctly taller than cingulum of posterior premolar, within toothrow; canine and posterior premolar separated. M3 with third ridge from just over half to ca. three-quarters the length of the second ridge. M3–M3: 43.1 (41–44)% of GLS, n = 11 (cf. T. pumila). One or two lower incisors on each side (depending on locality); bicuspid, sometimes crowded. Lower canines slender and relatively short, with cingula not greatly enlarged and not in contact. Dental formula: 1123/2123 = 30 or (rarely) 1123/1123 = 28. For additional information see Fenton & Eger (2002). Geographic Variation Koopman (1994):

a

Three subspecies are recognized by

T. c. chapini: Ethiopia to DR Congo. Ventral pelage greyish-brown. T c. lancasteri: NE Angola to Botswana and Zimbabwe. Ventral pelage pure white. T. c. shortridgei: SW Angola, NW Namibia.Ventral pelage greyish-brown. The West African material has not been allocated. b

c

Figure 112. The long, bicoloured interaural crest of Tadarida chapini. (a) Lateral view based on photo in which details of face and ear are obscure. (b) Dorsal view (BMNH 52.1505, T. c. lancasteri, Zambia). (c) Dorsal view showing crest erected (BMNH 66.6036, T. c. lancasteri incorrectly labelled T. pumila, Zambia).

Similar Species Only one other African Tadarida has the following combination of characters: ears joined by interaural band; M3 with third ridge > half length of second; no paired tail-glands at base of tail; FA: half length of second ridge. Sexes different in colour, similar in size. Pelage velvety; mid-dorsal hairs 6–7 mm. Dorsal pelage ("") dark chocolate brown; no spots or flecking; hairs pale greyishbrown with dark chocolate brown tips. Ventral pelage ("") dark chocolate brown on flanks, paler on abdomen with white or cream mid-ventral band; ventral flank-stripe white or cream. Dorsal pelage (adult !!) reddish-brown; no spots or flecking; hairs with basal half pale greyish-brown or creamy-fawn. Ventral pelage (adult !!) reddish-brown on flanks, paler on abdomen with cream or yellowish mid-ventral band; ventral flank-stripe yellowish. Subadults same as adult "". Head not extremely flattened. Upper lip without well-defined wrinkles; comparatively few spoon-hairs. Ears dark brown, comparatively small (not reaching snout when laid forward); with inner margins meeting at base of forehead to form a V-shaped valley. Tragus large, roughly rectangular, not concealed by antitragus. Antitragus low, triangular, only slightly larger than tragus. No interaural crest. Gular gland naked, conspicuous especially in !! in which it is often ringed by greenish hairs.Wingmembranes brown dorsally (paler over phalanges), whitish ventrally becoming browner towards tips. Interfemoral membrane dorsally brown, ventrally whitish, darkening towards margins. Ventral sides of forearms and legs naked and white. Foot with raised pad on sole (plantar pad). Skull not extremely dorsoventrally flattened. Anterior of braincase elevated above plane of rostrum, but sometimes only slightly. Sagittal crest very weakly developed in !!, absent in ""; lambdoid crests

well developed in both sexes. Anterior palatal emargination wide. Interdental palate broad posteriorly, narrowing smoothly towards canines with only a slight constriction at level of posterior premolars (cf. T. lobata). Basisphenoid pits deep, large, their width greater than their distance apart. Upper incisors short and close together. Anterior upper premolar ca. same height as cingulum of posterior premolar, slightly displaced labially but canine and posterior premolar not in contact although sometimes nearly so. M3 with third ridge > half second ridge. Two lower incisors on each side, bicuspid and crowded. Lower canines well developed; cingula not greatly enlarged but nearly in contact. Dental formula: 1123/2123 = 30. Geographic Variation

None recorded.

Similar Species Four other Tadarida in Africa have ears with inner margins meeting, or almost meeting, to form a V-shaped valley: Tadarida aegyptiaca. Almost always smaller (FA: 42–55 mm; GLS: 17.1–21.9 mm). Wings brown or blackish ventrally and dorsally. Ventral pelage with no mid-ventral markings and usually no ventral flank-stripe in contrasting colour. T. lobata. Inner margins of ears meeting close to snout; ears extending well beyond snout when laid forward. White spot between shoulder-blades of adults. Skull with interorbital constriction slight; interdental palate markedly constricted at level of posterior premolar. Lower canines with cingula separated by the incisors. T. teniotis. Three lower incisors on each side. Wings blackish. North Africa. T. ventralis. No ventral flank-stripe in contrasting colour. Foot usually without plantar pad. Skull usually larger and more robust (GLS: 23.4–26.1 mm). Upper incisor longer. Distribution In Africa, recorded mainly from Somalia–Masai Bushland, Zambezian Woodland and Afromontane–Afroalpine BZs, but probably extends into Eastern Rainforest–Savanna Mosaic near mountainous areas. Recorded from E DR Congo, Uganda, Rwanda, Kenya and Tanzania and southwards to NE South Africa 511

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Family MOLOSSIDAE

backwards and sideways. Cannot take off from ground, and needs to dive to gain sufficient speed for flight. Roosts by day in narrow, horizontal rock-crevices located either high up in vertical cliffs or under overhangings. In Zimbabwe, most known roosts are in granite inselbergs; one colony roosted in cracks in the ceiling of a dolomite cave. Scuttles around roost-sites; scuttles backwards into crevices, using tip of tail to feel the way. In Zimbabwe, sometimes shares roosts with T. ansorgei and T. bivittata (Cotterill & Fergusson 1993a). Often mist-netted over water, suggesting dependence on accessible water for drinking. Foraging and Food Forages by fast-hawking. Emerges at dusk, remains active all night and does not return to the day-roost until just before first light (Smithers 1983, F. P. D. Cotterill pers. obs.). Feeds on flying insects; details not known.

Tadarida fulminans

(Harrison 1971, Hayman & Hill 1971, Ansell 1978, Baeten et al. 1984, Happold et al. 1987, Cotterill 1996b, 2001c). Extralimitally: Madagascar (two specimens – the holotype collected in late nineteenth century and a melanistic topotype [FMNH 166074] collected in 1999). Habitat Apparently prefers woodland savannas where steep rocky terrain provides suitable day-roosts. No details for the northern records except many appear to be near mountains. In Malawi, recorded in miombo woodland at base of Mulanje Mountain, which is a great, granite-syenite massif with precipitous, bare sides. In SW Zambia and NW Zimbabwe, recorded from the precipitous basalt gorges of the Upper Zambezi R. (Victoria Falls and Kalomo). In S and E Zimbabwe, recorded from a range of habitats including miombo and mopane woodlands, but mostly recorded in miombo woodland on the granitic shield, especially in rugged granite country where there are high, steep-sided inselbergs (Cotterill 2001c). Also recorded from day-roosts in a dolomite cave and in Karoo sandstone in the Sebungwe and Limpopo basins.

Echolocation Search-phase call-shape shallow FM. In Zimbabwe, start-frequency 27 kHz; end-frequency 14 kHz; peak-frequency 17 kHz; maximum duration 20 ms (1 bat, flying free; Fenton & Bell 1981). In South Africa, start frequency 27 kHz; end-frequency 14 kHz; intensity high (one bat in flight-cage; Aldridge & Rautenbach 1987). Social and Reproductive Behaviour Roosts singly or in groups. No more than 20 individuals have been taken from any one day-roost but, in some cases, not all group-members were taken (Smithers 1983, F. P. D. Cotterill pers. obs.). Four captive adult !! roosted separately, suggesting that they are territorial; adult !! have also been observed roosting singly in rock-crevices. In contrast, captive "" huddled tightly together (F. P. D. Cotterill & N. C. Bennett unpubl.). Within day-roosts, there is much jostling and squeaking (audible to humans). Group-members emerge from the roost together, and make shrill squeaks as they dive away.

Reproduction and Population Structure Litter-size: one (n = 10); only right ovary is functional. Based on small samples of "" collected at 17° 30' S in NE Zimbabwe at ca. monthly intervals from Jul 1988 to Sep 1989 (Cotterill & Fergusson 1993b), the reproductive chronology appears to be seasonal polyoestry, but further data are needed to elucidate the details. Births followed by postpartum oestrus are inferred to have occurred in ca. Mar– May at beginning of dry season (evidence: in late Jul, 3 of 5 "" were simultaneously lactating and in very early pregnancy and two Abundance Locally common, but patchily distributed (based on were lactating with sperm in uteri and in vaginae; and in late Aug, a series collected in Rwanda, and observations and captures at roosts 4 of 4 "" were post-lactating and in early pregnancy). In midin NE and S Zimbabwe). Distribution and abundance is probably Oct, 3 of 3 "" were pregnant. In mid-Nov, 5 of 5 "" were determined by the availability of suitable day-roosts (Cotterill lactating and also pregnant with full-term foetuses; this indicates 2001c). Rarely collected because it flies high and roosts in inaccessible births from ca. late Oct to Nov (beginning of hot-wet season) and places. suggests either that the lactation period after the Mar–May births is exceptionally long, or that these bats had had a previous litter in ca. Adaptations Aspect ratio very high; wing-loading very high to Aug at some other locality. There was no evidence of postpartum exceptionally high (F. P. D. Cotterill unpubl.); this species, and T. oestrus after the Oct–Nov births (8 of 8 "" in Dec, 5 of 5 in ventralis, have the narrowest wing-tips found in any African Tadarida Feb and 2 of 2 in Apr were lactating but not pregnant), therefore (Freeman 1981). Flight fast, agile, usually direct (Smithers 1983); this study did not indicate when the "" that gave birth in Mar– manoeuvrability very poor; the agile swooping flight as a bat returns May became pregnant. Perhaps only some "" have a postpartum to its day-roost, with air rushing over its wings, is very reminiscent oestrus following births in Oct–Nov, or perhaps births in May– of the flight of swifts (Apodidae). Adept at rapid scuttling forwards, Jun do not occur every year. Perhaps this species migrates, and 512

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Tadarida gallagheri

its reproductive chronology cannot be elucidated from studies at only one locality. Considered overall, the unusual reproductive chronology of T. fulminans suggests that the pattern of annual fluctuation in the availability of its prey differs from those of the prey exploited by other Chiroptera. Predators, Parasites and Diseases Predators include Mackinder’s Eagle-owl Bubo capensis, which has been observed capturing this bat in NE Zimbabwe (Cotterill 1992). Ectoparasites include a bed-bug Crassicimex sexualis (Hemiptera: Cimicidae), another bug Hypoctenes faini (Hemiptera: Polyctenidae) and mites Microtrombicula intranasalis (Acari:Trombiculidae), Notoedres spp. (Acari: Sarcoptidae) (Anciaux de Faveaux 1984). All examined T. fulminans carried one or more interstitial nematodes (ca. 10 mm overall length), clearly visible under skin on underside of wing-membrane along anterior edge of the forearm. (Note: internal parasites are not mentioned in other profiles.) Conservation

IUCN Category: Least Concern.

Measurements Tadarida fulminans FA: 58.5 (56–61) mm, n = 50 WS (d): 428 (400–510) mm, n = 21 TL: 144 (132–161) mm, n = 24 T: 57.5 (53–66) mm, n = 24 E: 23.5 (19–25) mm, n = 23 Tr: n. d. Tib: 21.2 (20–22) mm, n = 18 HF: 13.0 (11–15) g, n = 14* WT: 34.6 (23–49) g, n = 58 GLS: 23.0 (21.6–24.4) mm, n = 39 GWS: 13.6 (12.9–14.2) mm, n = 36 C–M3: 8.5 (7.8–9.2) mm, n = 40 Throughout African geographic range (BMNH, FMNH, HZM, NMZB, RMCA, ROM, USNM) *Zimbabwe (Smithers & Wilson 1979) Key References 1993b.

Cotterill 1996b, 2001c; Cotterill & Fergusson F. P. D. Cotterill

Tadarida gallagheri GALLAGHER’S FREE-TAILED BAT Fr. Tadaride de Gallagher; Ger. Gallaghers Bulldoggfledermaus Tadarida gallagheri Harrison, 1975. Mammalia 39: 313. Scierie Forest, 30 km south-west of Kindu, DR Congo.

Taxonomy Currently in subgenus Chaerephon. Synonyms: none. At the time of its discovery, the uniqueness of this little molossid, especially its nasal swellings and interaural membrane morphology, suggested that it might represent a distinct genus (Harrison 1975). Subsequently, T. tomensis, another species with very similar nasal swellings and interaural membrane morphology, was described from São Tomé (Juste & Ibáñez 1993b). Both of these species bear some resemblance to Tadarida (Chaerephon) johorensis fromAsia. Chromosome number: not known.

and extends beyond the snout. This lobe is penetrated by a deep, backward-opening pocket. The pocket is bordered by two vertical hairless folds of skin, which extend backwards out of the pocket and unite on the crown of the head. Arising from the smooth floor of the pocket is an interaural crest of dark brown hairs, ca. 9 mm long, and presumably the pocket can be partly everted to display this crest. Tragus small but not concealed by antitragus. Antitragus well developed, roughly rectangular. No gular gland. Wings and interfemoral membrane greyish-black.

Description Very small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; small for an African molossid; with inner margins of ears joined by an interaural membrane and a large, bulbous, interaural lobe, which projects well forward over the muzzle and dominates the lateral profile of the head (Figure 97g): no other mainland species resembles this bat. Sexual dimorphism: no information (only one ! specimen known). Pelage short (middorsal hairs 5–6 mm). Dorsal pelage uniformly umber brown; no spots or flecking. Ventral pelage uniformly umber brown; no mid-ventral markings; ventral flank-stripe umber brown. Head not extremely flattened; muzzle blunt with transverse ridge above snout. Upper lip with five well-defined wrinkles on each side and few spoon-hairs except in front of the snout. Ears relatively large, extending beyond snout when laid forward; inner margins joined by a naked interaural membrane that rises centrally to form a flattopped, truncated triangle with convex sides. Below this, there is a large bulbous lobe, which projects forward over the muzzle

Figure 114. Tadarida gallagheri. Skull (holotype; based on Harrison 1975).

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Skull (Figure 114) not extremely dorsoventrally flattened. Rostrum relatively very long for a Tadarida, with prominent paired nasal swellings surrounding the nasal aperture. Anterior of braincase prominantly elevated above plane of rostrum (Figure 99a). Sagittal crest absent; lambdoid crest weakly developed. Anterior palatal emargination closed, incisive foramina not visible in holotype. Basisphenoid pits deep, large, much wider than the narrow septum separating them. Dentition not robust. Upper incisors comparatively large, well separated. Anterior upper premolar ca. same height as cingulum of posterior premolar, within toothrow but lateral to cingulum of the canine; canine and posterior premolar well separated. M3 with third ridge > half length of second. Two lower incisors on each side; not crowded; inner incisor larger and faintly bicuspid; outer incisor narrower and unicuspid. Lower canines slender; cingula not greatly enlarged and not in contact. Dental formula: 1123/2123 = 30. Geographic Variation No information. Similar Species Only one other Tadarida in the African region (and T. johorensis from Asia) has a bulbous lobe containing an interaural pocket projecting over the muzzle, and a rostrum with nasal swellings: Tadarida tomensis. Known only from São Tomé I. in Gulf of Guinea (and therefore not profiled), but occurrence on mainland is possible. Bulbous lobe similar but not projecting quite as far forward. No folds of skin extending backwards out of the pocket and uniting on crown of head. Swellings surrounding nasal aperture smaller; nasal aperture longer (Juste & Ibáñez 1993b). Distribution Known only from type locality in Rainforest BZ (South Central Region).

Habitat The only known specimen was collected in the vicinity of the village of Scierie (03° 10' S; 25° 49' E) near the west bank of the Congo R. It was mist-netted in a clearing within the thick, evergreen undergrowth of a partly deciduous forest, in which the secondary regrowth was well advanced. The forest had been heavily logged and only a few giant trees remained (Gallagher & Harrison 1977). Remarks Wings long and narrow, therefore flight predictably fast and agile, but with poor manoeuvrability, as in other molossids. The extraordinary nasal swellings, which are reminiscent of those of Rhinopoma, are so distinctive that generic status should be considered for this bat (Harrison 1975). In most features, however, it represents a highly specialized Chaerephon, a swift-flying species that has developed some special modification of the nasal region for echolocation (Harrison 1975). The discovery of a " is awaited with great interest as the structure of its ears may differ from that of the !. It is not inconceivable that the three female bats from São Tomé, described as Tadarida tomensis by Juste & Ibáñez (1993b), represent T. gallagheri but, if so, the species exhibits a marked and very unmolossid-like degree of sexual dimorphism, especially in the extent to which the interaural pouch and the nasal swellings are developed. Furthermore, the skulls and teeth of the two forms differ in other ways, for example the extent of the interorbital constriction, the sizes and shapes of the upper incisors and canines, and the size and depth of the basisphenoid pits (Juste & Ibáñez 1993b), and these differences are more likely to occur in different species than in different sexes of the same species. Conservation IUCN Category: Data Deficient. Known from only one location, and population reduction is predicted to occur based on decline in area of occupancy, extent of occurrence or quality of habitat (Hutson et al. 2001). Other collections in area have not found this species. Major threat: loss of habitat because of logging, clear-cutting and farming. However, possibly more widespread than presently believed. Measurements Tadarida gallagheri FA: 38 mm WS: n. d. TL: 77 mm T: 28 mm E: 19 mm Tr: n. d. Tib: 13 mm HF: 8.4 mm WT: n. d. GLS: 16.0 mm GWS: 8.9 mm C–M3: 5.8 mm DR Congo (holotype, adult !, Harrison 1975, Juste & Ibáñez 1993b) Key References

Tadarida gallagheri

Gallagher & Harrison 1977; Harrison 1975. F. P. D. Cotterill

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Tadarida lobata

Tadarida lobata BIG-EARED FREE-TAILED BAT Fr. Tadaride du Kenya; Ger. Großohr-Bulldoggfledermaus Tadarida lobata (Thomas, 1891). Ann. Mag. Nat. Hist., ser. 6, 7: 303. Turkwell Gorge, West Pokot, Kenya.

Taxonomy Originally Nyctinomus lobatus. Subgenus Tadarida. Synonyms: none. Chromosome number: not known. Description Medium-small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; large for an African molossid; ears very large, meeting to form a V-shaped valley; wings greyish; ventral flank-stripe white; two lower incisors on each side; M3 with third ridge > half length of second ridge. Sexes almost similar; !! larger in some skull dimensions including C–M3. Pelage short, velvety. Dorsal pelage dark brown (various shades) with conspicuous white spot between shoulder-blades in all known fully mature adults, and variable amounts of white or pale pelage behind the junction of the ears; hairs bicoloured, base white, tip dark brown. Ventral pelage brown on flanks, very much paler (sometimes white) mid-ventrally; no mid-ventral markings; ventral flank-stripe fluffy, white, contrasting with flanks. Orange-phase: not known. Head not extremely flattened. Upper lip without well-defined wrinkles; comparatively few spoonhairs (Freeman 1981). Ears semi-translucent pale greyish-brown; relatively large (extending well beyond snout when laid forward); inner margins meeting to form a V-shaped valley. Uniquely, the bases of the inner margins meet well forward on the muzzle where they join to form a membrane extending along muzzle almost as far as the snout (Figure 115). Tragus large, roughly rectangular, not concealed by antitragus. Antitragus moderate, roughly twice as big as tragus. No interaural crest. Gular gland present, equally developed in both sexes. Wings and interfemoral membrane semi-translucent, greyish dorsally, whitish ventrally. Ventral sides of forearms and legs naked and white. Foot with raised pad on sole (plantar pad). Skull not extremely dorsoventrally flattened. Anterior of braincase only slightly elevated above plane of rostrum (Freeman 1981). Sagittal crest weakly developed in !!, inconspicuous in ""; lambdoid crests distinct in both sexes. Anterior palatal emargination wide. Interdental palate broad posteriorly, narrowing towards canines but not smoothly

– there is a marked constriction at level of posterior premolars (cf. T. fulminans). Basisphenoid pits shallow to moderate in depth, mediumsized, their width slightly greater to slightly less than their distance apart. Upper incisors moderate in length and close together. Anterior upper premolar distinctly taller than cingulum of posterior premolar, within toothrow; canine and posterior premolar well separated. M3 with third ridge > half length of second ridge. Two lower incisors on each side; minute. Lower canines slender; cingula not enlarged, separated by the incisors. Dental formula: 1123/2123 = 30. Geographic Variation

None recorded.

Similar Species Four other Tadarida in Africa have ears with inner margins meeting, or almost meeting, to form a V-shaped valley (Table 20, p. 488): Tadarida aegyptiaca. Usually smaller (FA: 42–55 mm; GLS: 17.1– 21.9 mm). Inner margins of ears almost meeting on forehead; ears not extending beyond snout when laid forward. No white spot between shoulder-blades. T. fulminans. Inner margins of ears meeting on forehead; ears not extending beyond snout when laid forward. No white spot between shoulder-blades. Skull with interorbital constriction moderate; interdental palate only slightly constricted at level of posterior premolar. Lower canines with cingula nearly in contact. T. teniotis. Three lower incisors on each side. Inner margins of ears meeting on forehead; ears not extending beyond snout when laid forward. Interdental palate not markedly constricted at level of posterior premolar. North Africa. T. ventralis. Inner margins of ears meeting on forehead; ears not extending beyond snout when laid forward. No white spot between shoulder-blades. Skull with interorbital constriction pronounced. Anterior upper premolar minute; canine and posterior premolar nearly in contact. Lower canines well developed; cingula in contact. Distribution Endemic to Africa. As yet, only recorded from Afromontane–Afroalpine and Somalia–Masai Bushland BZs in Kenya, and Zambezian Woodland BZ in Zimbabwe. In Kenya, recorded from four localities (including type locality) in the northwest, and Maungu Hill in the south-east. In Zimbabwe, recorded from the Sengwa Gorge, Siabuwa District in the north-west, from two localities (Borrowdale Brook & Hatfield) on outskirts of Harare (Cotterill 1996b), and recently from the Mutoko District, where specimens were mist-netted in 1997 and 1999 (Cotterill 2001c).

Figure 115. Left ear of Tadarida lobata (based on Peterson & Harrison 1970).

Habitat Woodland savannas with suitable sites for day-roosts. In Kenya, 19 individuals were taken on a large flat plain of open thorn scrub with scattered rocky hills including Maungu Hill, a high rock outcrop with deep crevices along its cliff face (A.Williams in Peterson 1974). In Zimbabwe, one was mist-netted on the wall of a dam near 515

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Remarks Wings very long and very narrow, as in T. fulminans and T. ventralis. Flight fast, agile, with poor manoeuvrability. Flies fast and straight, or circles more slowly (Peterson 1974). Bats converging on the areas around two water tanks (? dams) in Kenya, uttered a distinctive single- or double-noted loud squeak, audible to humans; no more than 2–3 individuals were seen or heard at a time. Although obviously attracted to the water, none of the bats was seen to descend to drink, and none was caught in mist-nets set over the tank. Two specimens from Harare (Zimbabwe) were found dead in suburban gardens; eight others were mist-netted near open water in the Mutoko District (Zimbabwe) in association with Sauromys petrophilus, T. ansorgei and T. fulminans. Three of these were mist-netted within the first hour of darkness in Nov 1997 and Nov 1999. All eight were lactating "", suggesting parturitions had occurred in late Oct or early Nov. Conservation IUCN Category: Least Concern. New records in Zimbabwe suggest that T. lobata is more widespread and perhaps more abundant than estimated by Hutson et al. (2001). Tadarida lobata

precipitous sandstone cliffs of the 10 km-long Sengwa Gorge; the vegetation is semi-arid, mainly Combretum/Commiphora scrub and mopane woodland (Cotterill 2001c). The Harare specimens were taken in miombo woodland within 20 km of small granite outcrops, and the Mutoko specimens from a mosaic of farmlands with miombo woodland in terrain dominated by large granitic inselbergs, many with precipitous sides. The Mutoko specimens were mist-netted over, or very near, open water. Abundance Uncertain. Tadarida lobata is one of Africa’s rarest and most enigmatic molossids. Despite its description in 1891, it remained known only by the holotype until single specimens were collected from Kenya and Zimbabwe in 1964 and 1969, respectively (Peterson & Harrison 1970). Two larger series of specimens were collected from these two countries subsequently. This species appears to be locally common (based on a series of 19 specimens shot at Maungu Hill, Kenya, by Peterson [1974]; and eight specimens mist-netted in the Mutoko District of Zimbabwe in 1997 and 1998 [Cotterill 2001c]).

Measurements Tadarida lobata FA: 58.0 (55–62) mm, n = 38 WS (d): 407 (370–425) mm, n = 38 TL: 131.8 (124–146) mm, n = 40 T: 52.2 (46–60) mm, n = 40 E: 28.2 (25–32) mm, n = 40 Tr: n. d. Tib: 19.5 (19–21) mm, n = 13* HF: 12.9 (12–14) mm, n = 13* WT: 25.0 (20–33) g, n = 30 GLS: 22.8 (22.3–23.6) mm, n = 20 GWS: 13.4 (13.0–14.0) mm, n = 20 C–M3 (""): 8.7 (8.6–8.8) mm, n = 8* C–M3 (!!): 9.0 (8.9–9.1) mm, n = 5* Kenya and Zimbabwe (BMNH, HZM, ROM, NMZB, Peterson 1974) *Kenya only (Peterson 1974) Key References

Cotterill 1996b, 2001c; Peterson 1974 F. P. D. Cotterill

Tadarida major LAPPET-EARED FREE-TAILED BAT Fr. Tadaride à oreillettes; Ger. Stirnlappen-Bulldoggfledermaus Tadarida major (Trouessart, 1897). Cat. Mamm. Viv. Foss. 1: 146. 5th Cataract of the Nile, N Sudan.

Taxonomy Originally Nyctinomus pumilus var. major. Subgenus Chaerephon. Synonyms: abae, emini. Subspecies: none. Similarities between the comparatively simple lappet between the ears of T. major, and the highly developed, forward-projecting lobe between the ears of T. gallagheri from DR Congo and T. tomensis from São Tomé, indicate a need to re-examine the relationships between these species, and between these species and other Tadarida. Chromosome number: not known.

Description Small to very small microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; medium-small for an African molossid; ears separated by a distinctive lappet over an interaural pocket, which, in adult !!, contains an interaural crest; M3 with third ridge > half second ridge; wings usually dark (rarely white or pale), ventral flank-stripe white or whitish. Sexes almost similar. Pelage short (mid-dorsal hairs ca. 3 mm), covering rump but not extending

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onto interfemoral membrane. Dorsal pelage dark sepia brown, yellowish-brown, greyish-brown or dull rusty-brown; sometimes slightly grizzled but no white flecks or spots; hairs unicoloured, sometimes with white at tip. Ventral pelage similar becoming paler mid-ventrally; mid-ventral markings white, highly varied in size and shape; ventral flank-stripe white or whitish. Head not extremely flattened. Upper lip with 6–7 well-defined wrinkles on each side and many spoon-hairs. Ears dark brown or blackish; just reaching snout when laid forward; inner margins meeting to form V-shaped valley but, unlike other African Tadarida, there is a conspicuous, flat, subtriangular or roughly semi-circular flap (= lappet or lobe) projecting backwards between the inner margins of the ears: this forms a lid over a small pocket that opens posteriorly (Figure 97). In adult !!, the pocket under the lappet is lined by short (7 mm) rusty-brown hairs, which form an interaural crest. Tragus small, squarish, concealed by antitragus. Antitragus large, tall, almost rectangular but with top corners rounded. No gular gland. Wings semi-translucent, with varying degrees of dark brown or black pigmentation in most individuals – the holotype, however, has white wings (Rosevear 1965). Interfemoral membrane dark brown. Skull not extremely dorsoventrally flattened. Anterior of braincase not noticeably elevated above plane of rostrum. Sagittal crest low or absent; lambdoid crest moderate. Anterior palatal emargination slightly open (in young adults) or closed. Basisphenoid pits shallow to moderately deep, variable in size – sometimes elongated and ca. half as wide as their distance apart, sometimes oval and as wide as their distance apart. Anterior upper premolar ca. same height as cingulum of posterior premolar or lower or higher, within toothrow; canine and posterior premolar separated. M3 with third ridge equal to, or just over, half length of second. Two lower incisors on each side; bicuspid, somewhat crowded. Lower canines with cingula not greatly enlarged and either in contact or not in contact. Dental formula: 1123/2123 = 30. Geographic Variation None recorded. Similar Species No other African Tadarida has a flat lappet over an interaural pocket between the ears. Distribution Endemic to Africa. Distribution appears disjunct. Recorded from Sahel Savanna, Sudan Savanna and Guinea Savanna BZs and the Northern Rainforest–Savanna Mosaic, and from one locality in the Rainforest BZ (Western Region), from W Liberia to S Niger and Nigeria. Also occurs in these biotic zones, and also in the Sahara Arid BZ, in the valleys of the Nile R. and its tributaries in Sudan, in the Rainforest–Savanna Mosaics in NE DR Congo, Uganda, Kenya and Tanzania, and in the Coastal Forest Mosaic BZ in Kenya and Tanzania. It is not known if the apparent gap between ca. 10° E and 33° E reflects insufficient collecting or the absence of the species from this area for some unknown reason. Simmons (2005) indicates the occurrence of T. major in Senegal but without details. Mapped from country checklists (see order Chiroptera), other literature and museum records. Subsequently recorded widely in the south-western half of Uganda (Thorn & Kerbis Peterhans 2009): not mapped. Habitat Recorded mainly from woodland and grassland savanna vegetation zones including undifferentiated woodland, Isoberlinia

Tadarida major

woodland, Sahel Acacia wooded grassland and deciduous bushland, and Acacia–Commiphora deciduous bushland and thicket, and from riverine habitats along the Nile R. and its tributaries (terminology: White 1983). Also recorded from areas of invasive grassland in the mosaic of rainforest and secondary grassland in West Africa and NE DR Congo, and from the Coastal Forest Mosaic BZ (but not within the coastal forests). Abundance

Uncertain.

Adaptations The wings are long and narrow implying flight characteristics similar to those of other species of Tadarida. By day, roosts in dark crannies within rocky clefts in hills, in hollows formed by aggregations of boulders, in hollow trees, lofts of houses and holes in walls of buildings (Lang & Chapin 1917b, Rosevear 1965). Roosts are sometimes revealed by their musty smell and the incessant squeaking of their occupants. Lang & Chapin (1917b) noted that disturbed bats seemed intent on returning to their habitual roosts, suggesting that roost-fidelity is high. Emergence occurs well before nightfall (Rosevear 1965). The triangular lappet between the ears is usually laid backwards and, in the ", covers a hairless spot. In the !, it can be erected to expose the interaural crest. The hairs of the crest are thicker than those of the body, and structurally resemble scent-dispersing hairs (osmetrichia) (Hickey & Fenton 1987), which suggests that they might arise from glandular tissue, but the contexts in which the crest is erected are not yet known. Foraging and Food hawking.

No information. Predictably forages by fast-

Social and Reproductive Behaviour Roosts in groups of ca. four to more than 100. Rosevear (1965) suggests size of groups depends on available accommodation. Only two of many more than 30 individuals captured together were !! (Lang & Chapin 1917b), 517

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but the reproductive condition of these bats was not published and the social organization of this species is not known. Reproduction and Population Structure No information. Predators, Parasites and Diseases Ectoparasites include fleas Lagaropsylla anciauxi, L. taeniolae (Siphonaptera: Ischnopsyllidae); a tick Carios confusus (Acari: Argasidae); and a mite Chelanyssus aethiopicus (Acari: Macronyssidae) (Anciaux de Faveaux 1984, Beaucournu & Kock 1996). Conservation

IUCN Category: Least Concern.

TL: 100.1 (85–113) mm, n = 107 T: 33.8 (27–42) mm, n = 108 E: 16.8 (12–21) mm, n = 105 Tr: 4.3 (3.2–5.3) mm, n = 12 Tib: 14.2 (13–15) mm, n = 11 HF: 8.0 (7–11) mm, n = 43 WT: 15.3 (10–28) g, n = 55 GLS: 18.0 (17.0–19.3) mm, n = 45 GWS: 11.4 (10.6–12.3) mm, n = 50 C–M3: 6.9 (6.3–7.3) mm, n = 47 Throughout geographic range (BMNH, NMK, SMNS, ZFMK and literature) Key Reference

Measurements Tadarida major FA: 42.4 (39–46) mm, n = 120 WS (d): 323.3 (311–335) mm, n = 11

Rosevear 1965. Meredith Happold

Tadarida midas MIDAS FREE-TAILED BAT Fr. Tadaride midas; Ger. Midas-Bulldoggfledermaus Tadarida midas (Sundevall, 1843). Kongl. Svenska Vet.-Akad. Handl. Stockholm 1842: 207 [publ. 1843]. Jebel el Funj, White Nile River, West Bank, Blue Nile [= Bahr-el-Abiad Prov.], Sudan.

Taxonomy Originally Dysopes midas. Subgenus Mops. Synonyms: unicolor, miarensis. Subspecies: two; only one in Africa (but see Geographic Variation). Chromosome number (Somalia, Namibia, South Africa): 2n = 48; aFN = 66 (Smith et al. 1986; Rautenbach et al. 1993). Description Medium-sized microbat without noseleaf and with terminal portion of tail projecting freely from posterior margin of interfemoral membrane; large, robustly built and particularly heavy for an African molossid (FA: 59–67 mm; WT: 38–69 g); ears joined by interaural band; M3 with third ridge absent; anterior palatal emargination closed; wings dark brown; ventral pelage greyishbrown to silvery-grey; ventral flank-stripe white or pale. Sexes almost similar. Pelage short (mid-dorsal hairs 5–6 mm), silky, sparse; nape almost naked. Dorsal pelage (grey-phase) dark brown, medium sepia brown, pale brown or pale grey with pale grey or white frosting and flecking; hairs with paler base and mostly with white or pale grey tip. Crown same colour as back. Ventral pelage greyishbrown, pinkish-brown to silvery-grey, frosted (subadults almost white); no mid-ventral markings; ventral flank-stripe white or paler than ventral pelage. Orange-phase: dorsal pelage reddish to almost orange. Pelage colouration apparently uniform within colonies, but Verschuren (1957) found two colonies roosting in the same tree, whose members had distinctly different colours. Head not extremely flattened. Upper lip with 5–6 well-defined wrinkles on each side and many spoon-hairs. Ears blackish-brown, relatively large (extending just beyond snout when laid forward); inner margins joined over muzzle by interaural band with a V-shaped fold and a forwardprojecting pocket-like invagination, which protrudes ca. half-way along muzzle. Tragus small, squarish or hatchet-shaped; concealed by antitragus. Antitragus large, semi-circular. Males and "" with interaural crest of longer, brown hairs arising from the interaural

pocket. No gular gland. Wings and interfemoral membrane dark brown. Skull not extremely dorsoventrally flattened. Anterior of braincase moderately to prominantly elevated above plane of rostrum (Figure 99c). Sagittal crest moderately to well developed in both sexes; lambdoid crests well developed and forming a prominent helmet in adult !!, weakly to moderately developed in "". Anterior palatal emargination closed; incisive foramina visible. Basisphenoid pits moderately deep, medium-large, their width varying from equal to their distance apart to almost twice their distance apart. Anterior upper premolar variable in size (lower to taller than the cingulum of the posterior premolar), within toothrow. M3 with third ridge absent. Two lower incisors on each side, bicuspid, crowded or not crowded. Lower canines with cingula not greatly enlarged, sometimes well separated, sometimes almost in contact. Dental formula: 1123/2123 = 30. Geographic Variation Koopman (1994):

Two subspecies are recognized by

T. m. midas. Throughout African geographic range. T. m. miarensis. Madagascar. However, based on molecular genetics, the Madagascan and African populations are not distinct (Goodman 2011). Similar Species Five other African Tadarida have the following combination of characters: ears joined by interaural band; M3 with third ridge present (but usually* < half length of second ridge) or absent; anterior palatal emargination closed; FA: >40 mm (Table 20, p. 488). *In T. condylura, the length of the third ridge is variable and can be up to half length of second ridge.

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T. condylura. Much smaller and lighter (FA: 45–51 mm;WT: 16–39 g). Tadarida congica. Forearm shorter (54–58 mm). Dorsal pelage less variable, dark brown to black; not frosted and without whitish flecking. Skull of similar length but less robust and often narrower (GWS: 14.8–16.3 mm). T. demonstrator. Much smaller and lighter (FA: 41–46 mm; WT: 20– 24 g). T. niveiventer. Much smaller and lighter (FA: 44–48; WT: 20–30 g). T. trevori. Smaller (FA: 51–55 mm; GLS: 23.1–25.1 mm). M3 with third ridge present (but < half length of second ridge). Distribution In Africa, recorded from very scattered localities mainly in Sudan Savanna, Guinea Savanna and Zambezian Woodland BZs, with some records marginally in Sahel Savanna, Somalia–Masai Bushland and South-West Arid BZs, and the Rainforest–Savanna Mosaic in NE DR Congo. Recorded from Senegal to Eritrea and Ethiopia (but not from all countries) and southwards through S Sudan, NE DR Congo and W Kenya, and further southwards in N Namibia, SW Zambia, Botswana, Zimbabwe, S Malawi and NE South Africa. Distribution often follows major rivers and rift valleys including the Blue Nile R. in Sudan, the upper reaches of the White Nile R. and the East African Rift Valley, the Shire Valley in Malawi, and the valleys of the Zambezi R. and its tributaries, and the Limpopo R. in southern Africa. Gaps in distribution probably reflect insufficient sampling. Map based only on published records including Dorst (1959), Hayman et al. (1966), Smithers (1971), Vielliard (1974), Koopman (1975), Bergmans (1977b), Koopman et al. (1978), Ansell (1978), Cotterill (1996b) and others, and some museum records. Kingdon (1974), Smithers (1983) and Dunlop (1999) mapped vast areas (especially north of Equator) for which there appear to be no records. Extralimitally: Saudi Arabia, Madagascar. Habitat Open woodland savannas (including Sudanian undifferentiated woodland, Isoberlinia woodland and drier miombo

woodland), grassland savannas and, in the Kalahari Desert, the transition zone from undifferentiated woodland to Acacia bushland and wooded grassland. Almost always found near major rivers and wetlands including the Sudd in Sudan, the Okavango Swamp in Botswana and the Shire R. and its marshes in S Malawi. Evidently avoids closed forests. Not dependent on rocky, hilly or mountainous terrain for its day-roosts. Abundance Uncertain. Not very plentiful in collections (Rosevear 1965). Rarely recorded in Malawi (Happold & Happold 1997). Common, locally, in NE Botswana and Limpopo Valley (Zimbabwe) (Cotterill 1996b). Elsewhere: no information. Adaptations Wings long and narrow. Flight fast, agile with poor manoeuvrability. Roosts by day in pitch-dark attics, in hollow cement bricks of the walls of buildings, in dimly lit expansion joints of bridges, under corrugated-iron roofs and, in DR Congo, in very long, narrow fissures penetrating deeply into the trunks or large branches of trees (particularly Vitex and Parinari) (Verschuren 1957, Smithers 1971, Smithers & Wilson 1979, Rautenbach 1982, Fenton & Rautenbach 1986). In DR Congo, not found roosting in rockcrevices that were available (Verschuren 1957). Entrances must be accessible by direct flight, and high enough to allow the bats to dive when they leave the roost. Apparently not reported to share roosts with other species of bats except in Kruger N. P., South Africa, where a roost was shared with T. condylura and T. pumila. At Faradje, NE DR Congo, only present for a short period at end of dry season: small beetles (prey) were common at that time (Lang & Chapin 1917b). Foraging and Food Forages by fast-hawking in open spaces including over canopies, open water (rivers, swamps, flooded pans) and open plains. In DR Congo, individuals emerged at dusk and did not return to the day-roost until just before sunrise (Verschuren 1957). In Mar, ca. 12 were observed arriving about dusk to forage 15–30 m above the Dungu R.; they flew at high speed, and dodged and dived after small hard-shelled beetles, which were then common; stomachs of some of these bats contained remains of these beetles that had been masticated in flight (Lang & Chapin 1917b). In Botswana, observed flying 30–40 m above ground, and swooping down to a waterhole to take insects (Smithers 1971, 1983). In Kruger N. P., South Africa, radio-tracked individuals seemed to move at least 10 km from their day-roost; of 15 foraging flights observed, five (in immediate vicinity of roost) lasted half length of second ridge; palatal emargination wide; wings dark brown; no ventral flank-stripe in contrasting colour; two lower incisors on each side. Sexes similar except !! on average heavier than "". Pelage velvety, sometimes glossy; mid-dorsal hairs 4–5 mm. Dorsal pelage (grey-phase) chocolate brown, rusty-brown or dark brown without spots or flecking; hairs with beige or cream at base. Ventral pelage paler with white or cream mid-ventral stripe; ventral flank-stripe same colour as flank but with longer hairs. Orange-phase: dorsal pelage orangebrown, ventral pelage yellowish. Head not extremely flattened. Upper lip without well-defined wrinkles; comparatively few spoonhairs. Ears brown, relatively small (extending to ca. middle of muzzle when laid forward); inner margins meeting on forehead to form a V-shaped valley. Tragus large, roughly rectangular, not concealed by antitragus. Antitragus low, triangular, not much larger than tragus. No interaural crest. Gular gland naked, conspicuous (especially in !!), and often ringed by grey hairs. Wings and interfemoral membrane semi-translucent, dark brown to almost black. Ventral sides of forearms naked and whitish; ventral sides of legs naked and brownish. Foot without raised plantar pad on sole. Skull not extremely dorsoventrally flattened. Anterior of braincase not noticeably elevated above plane of rostrum. Sagittal crest very weakly developed; lambdoid crest well developed laterally but absent dorsally. Anterior palatal emargination wide. Interdental palate broad, narrowing smoothly (but slightly) to canines. Basisphenoid pits deep, medium-sized, their width slightly greater than their distance apart to almost twice their distance apart. Upper incisors long and close together. Anterior upper premolar variable (it can be lower, level with, or taller than the cingulum of posterior premolar); in toothrow or slightly displaced labially; canine and posterior premolar nearly in contact, sometimes in contact. M3 with third ridge > half length of second ridge. Two lower incisors on each side, bicuspid and crowded. Lower canines well developed; cingula enlarged and in contact or almost so, or separated. Dental formula: 1123/2123 = 30. Geographical Variation No information. Similar Species Four other Tadarida in Africa have ears with inner margins meeting, or almost meeting, to form a V-shaped valley: Tadarida aegyptiaca. Much smaller (FA: 42–55 mm; GLS: 17.1– 21.9 mm).

Tadarida ventralis

T. fulminans.Ventral flank-stripe white, cream or yellowish. Foot with plantar pad. Skull usually smaller and less robust (GLS: 21.6– 24.4 mm). Upper incisor shorter. T. lobata. Inner margins of ears meeting close to snout; ears extending well beyond snout when laid forward. White spot between shoulder-blades. Skull with interorbital constriction slight. Anterior upper premolar large; canine and posterior premolar well separated. Lower canines slender; cingula separated by incisors. T. teniotis. Three lower incisors on each side. Upper lip with five well-defined wrinkles. Anterior upper premolar large; canine and posterior premolar well separated. North Africa. Distribution Endemic to Africa. Recorded from scattered localities in Afromontane–Afroalpine, Somalia–Masai Bushland and Zambezian Woodland BZs, and Eastern Rainforest–Savanna Mosaic. Recorded from Eritrea and Ethiopia, S Sudan, E DR Congo, Kenya, Tanzania, Malawi, Zimbabwe, Mozambique and from somewhere unspecified in the former Transvaal (South Africa) (Harrison 1971, Hill & Morris 1971, Largen et al. 1974, Koopman 1975, Cotterill 1996b, 2001c). Most records are from Kenya (Cotterill 2001c). Voucher specimens (USNM) for two localities in Mozambique (Chiuta and Vila Gouveia) mapped by Smithers & Lobão Tello (1976) as T. africana actually represent T. fulminans. Habitat Mostly recorded from dry woodland savannas (including Acacia–Commiphora bushland, drier types of miombo woodland and mopane woodland), but also recorded from montane habitats (including Keren, Eritrea, at 1400 m, Fatam R. [= Blue Nile Gorge] 539

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Measurements Tadarida ventralis FA: 63.7 (60–67) mm, n = 49 WS (d): 459.7 (440–485) mm, n = 12 TL: 153.3 (142–168) mm, n = 36 T: 58.3 (51–66) mm, n = 58.3 Abundance Not known. E: 24.7 (18–29) mm, n = 36 Tr: n. d. Remarks Only five specimens were known until 1971 (Hayman Tib: n. d. & Hill 1971) but more have been collected since, especially from HF: n. d. Kenya (Cotterill 2001c). Nevertheless, all aspects of the biology of WT (!!): 45 (38–55) g, n = 23 T. ventralis remain barely known. A pregnant " with one foetus WT (""): 39 (31–46) g, n = 10 (CR = 30 mm) was collected from a rock crevice in E Zambia in mid- GLS: 23.8 (23.4–26.1) mm, n = 51 Nov (Ansell 1986b), and a skull was recovered from an owl pellet GWS: 15.2 (14.0–15.8) mm, n = 50 underneath crevices in a large granitic overhang in S Zimbabwe. This C–M3: 9.5 (8.9–10.0) mm, n = 53 site was occupied as day-roosts by T. fulminans and large numbers of T. Throughout geographic range (BMNH, FMNH, HZM, LACM, ansorgei (Cotterill 1996b).Two specimens from Kenya were collected NMZB, ROM) as they emerged from the roofs of houses (Start 1966, Cotterill 1996b). Tadarida ventralis occurs sympatrically with T. lobata at Maungu Hill, Key References Cotterill 1996b, 2001c; Harrison 1971. Kenya (Cotterill 2001c). F. P. D. Cotterill Conservation IUCN Category: Data Deficient. Remains poorly known despite intensive sampling effort. Most records are historical. Has a wide geographic range, but is inferred to be threatened by pesticides and loss of habitat. at Great Abbai, Ethiopia at 1900 m, and the Kenyan Highlands), and from semi-desert grassland and shrubland near L. Turkana, Kenya. The availability of rock crevices in which to roost appears to be a major determinant of their occurrence in these vegetation zones (Cotterill 2001c).

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Family VESPERTILIONIDAE

Family VESPERTILIONIDAE VESPER BATS Vespertilionidae Gray, 1821. London Med. Repos., 15: 299. Subfamily Vespertilioninae Barbastella (2 species) Eptesicus (5 species) Glauconycteris (12 species) Laephotis (4 species) Mimetillus (1 species) Nyctalus (2 species) Nycticeinops (1 species) Otonycteris (1 species) Pipistrellus sensu lato (28 species) Plecotus (3 species) Scotoecus (2 species) Scotophilus (6 species) Subfamily Myotinae Cistugo (2 species)* Myotis (11 species) Subfamily Miniopterinae* Miniopterus (5 species) Subfamily Kerivoulinae Kerivoula (7 species) Phoniscus (? 1 species)

Barbastelles Serotines Butterfly Bats African Long-eared Bats Mimic Bat Noctules Twilight Bat Desert Bat Pipistrelles Long-eared Bats Lesser House Bats House Bats Wing-gland Bats Myotises Long-fingered Bats Woolly Bats Trumpet-eared Bats

p. 545 p. 546 p. 550 p. 560 p. 578 p. 585 p. 589 p. 594 p. 597 p. 600 p. 660 p. 666 p. 672 p. 684 p. 685 p. 688 p. 710 p. 711 p. 723 p. 724 p. 734

fawn with yellowish wings patterned with fine brown ‘veins’ so that they resemble dry leaves, and the Rufous Myotis Myotis bocagii has a subspecies that is bright rufous dorsally and cream ventrally with very dark brown ears, snout and wings. African vespertilionids include Pipistrellus musciculus (FA: 25, WT: 3–4 g), which is one of the smallest of all mammals, and Scotophilus nigrita (FA: 77–88 mm, WT: 88–91 g), which is one of the largest microbats. Most, however, are small to very small; some medium-sized. Most have pelage that is shortish and dense, or woolly, and most have slightly elongated bodies. In other characters, vespertilionids are very varied. The head can be rounded to very flat; the muzzle conical, and short to long. The ears (Figure 116) are comparatively simple, usually widely separated but joined in two genera, often relatively short (means 23–48% of FA), sometimes relatively long (means 72–97% of FA).The tragus is well developed and its shape is often used to identify genera and species (Figure 116). The eyes are small and sometimes hidden by pelage.The wings are very varied in shape and colour; the second finger has a metacarpal and one short phalanx (Figure 117).The hindlimbs vary in relative length and, in some species, are modified for climbing over slippery leaves.The toes (except the first [hallux]) have three phalanges. The interfemoral membrane is broad and long. Usually it fully encloses

* Since this profile was submitted, it has been indicated that two of these taxa should be raised to family status: the genus Cistugo to Cistugidae (Lack et al. 2010), and the subfamily Miniopterinae to Miniopteridae (Miller-Butterworth et al. 2007). As major updating was not possible, both are still presented here as vespertilionids.

This is a very large polytypic family that is distributed throughout the world wherever there are enough insects to eat and day-roosts of some sort: only the polar regions, the highest mountains and the most arid deserts are uninhabited by vespertilionids. Having 48 extant genera and 407 extant species (Simmons 2005), this is the largest family of bats and, among mammals, it is second in size only to the rodent family Muridae. The family Vespertilionidae is also one of the most widely dispersed families of mammals. In Africa, there are ca. 93 species (some ‘species’ almost certainly contain two or more cryptic species, and two species, Pipistrellus permixtus and Phoniscus aerosa, are unlikely to be genuine African species). Simmons (2005) recognizes 19 genera which have African representatives: here, partly because their species contents are uncertain, two genera (Neoromicia and Hypsugo) are included in Pipistrellus sensu lato. Vespertilionids have a simple muzzle without a noseleaf (Figure 32i), and the tail is completely enclosed by the interfemoral membrane (except for part of the terminal vertebra in some species) (Figure 33i). Most are brown, grey or blackish-brown with paler bellies and are often dubbed ‘little brown bats’. However, some vespertilionids are unexpectedly colourful, with delicate to spectacular markings, and are among the most beautiful of all mammals. For example, the Pied Butterfly Bat Glauconycteris superba has a striking pattern of white patches and lines on a black background, and black wings. Welwitsch’s Bat Myotis welwitschii is rufous and cream with spectacular wings that are black with bright orange-red markings. The Variegated Butterfly Bat Glauconycteris variegata is often creamy-

a

b

d

h

e

i

c

f

g

j

k

Figure 116. Ears and tragi of African bats in the family Vespertilionidae to show differences between subfamilies. (a) Ears joined (e.g. Barbastella, Vespertilioninae). (b) Ear not funnel-shaped; widely separated from other ear (e.g. Miniopterus, Miniopterinae). (c) Ear funnel-shaped; widely separated from other ear (e.g. Kerivoula, Kerivoulinae). Tragus short and blunt as in (d) Scotoecus, Vespertilioninae; (e) Laephotis, Vespertilioninae; (f) Glauconycteris, Vespertilioninae and (g) Miniopterus, Miniopterinae. Tragus long and narrow but not sharply pointed as in (h) Otonycteris, Vespertilioninae and (i) Myotis, Myotinae. Tragus long, narrow and sharply pointed as in (j) and (k) Kerivoula spp., Kerivoulinae.

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2nd 1st

a

b

c

Figure 117. Variations in the length of the second phalanx of the third finger in relation to length of the first phalanx of third finger, in African Vespertilionidae. (a) Second phalanx ca. as long as first (most Vespertilioninae). (b) Second phalanx ca. twice as long as first (Glauconycteris, Vespertilioninae). (c) Second phalanx about three times as long as first (Miniopterus, Miniopterinae).

the tail, which extends to the membrane’s V-shaped posterior border although, in some species, the terminal vertebra protrudes beyond the interfemoral membrane. Relative tail length varies from short to long (27–54% of TL). Females usually have one pair of nipples in the pectoral region; some species (but only Otonycteris hemprichii in Africa) have two pairs. There are no pubic nipples. The skull has a rostrum that varies from short to moderately long, and the cranial region varies from flat to highly domed.There are no postorbital processes and no enlarged supraorbital ridges.The sagittal and lambdoid crests are usually low but may combine to form a helmet in the largest members of some genera. The premaxillae have nasal branches that are usually widely separated; the palatal branches are absent and consequently the palate has a deep to very deep, wide anterior emargination. Within this family, there is a trend towards shortening of the jaws and a concomitant loss of incisor and premolar teeth: there may be a total of 38 teeth, or as few as 28. Consequently the dental formula varies from genus to genus and is used diagnostically (Figure 119). There is greater diversity in the adaptations of the vespertilionids than in any other family of bats in Africa. In African species (excluding Cistugo and Miniopterus), aspect ratios range from low to medium, and wingloadings range from very low to extremely high (the majority have low aspect ratios and low or very low wing-loadings).Tip shapes range from short and rounded to long and pointed. In Miniopterus, aspect ratios range from medium to very high, and wing-loadings from low to medium. Most vespertilionids, including all African species, are insectivorous (a few species from elsewhere eat fish), but Nyctalus lasiopterus also preys on small birds migrating during the night (see profile). The flying abilities, foraging strategies and diets of vespertilionids are as varied as their wing morphology. Foraging behaviour includes fast-hawking (e.g. Nyctalus, Pipistrellus savii), moderately fast-hawking (e.g. Miniopterus, Pipistrellus tenuipinnis, Scotophilus, Scotoecus and at least one Glauconycteris), slowhawking (most other species), gleaning (e.g. Eptesicus serotinus, Kerivoula spp., some Myotis spp., Otonycteris hemprichii and Plecotus christii) and trawling (e.g. Myotis bocagii, M. capaccinii). Some species exemplify more than one strategy. There are species of vespertilionids in almost all environments, from open to densely cluttered. Many vespertilionids use their wings to flick a flying insect into a temporary pouch created between the legs by the interfemoral membrane, and then tuck the head down to remove the insect and eat it. Their aerial acrobatics, associated with the pursuit, capture and manipulation of prey, are spectacular to watch and, as their name implies, vesper bats often forage in the evening and can be watched with ease.They often forage in groups.The adaptive radiation within this family has been so extensive that there are many examples of convergent evolution. In Africa, for example, the wing morphology and foraging habits of vespertilionids in the genus Kerivoula are similar to those of nycterids while, at the other end of the continuum, vespertilionids such as Scotophilus and Scotoecus have converged towards the emballonurids and molossids. This convergence also embraces echolocation. The majority of African vespertilionids for which data are available, emit high intensity steep FM/QCF calls (Figure 118), but

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Family VESPERTILIONIDAE 110

b

100

p

h

90

s r

j

Frequency (kHz)

80 70

c a

d

n g

e

m

60 50

f

q 0

l i k

40 30 20 10 Time (ms)

Figure 118. Sonograms of individual echolocation calls emitted by 13 species of vespertilionids from Malawi to indicate differences between species and, in some cases, differences associated with echolocating in different situations: all bats flying < 3 m above ground except where indicated (M. Happold unpubl.). (a) Laephotis botswanae; (b) Pipistrellus nanus; (c) P. zuluensis; (d) P. capensis; (e) P. cf. melckorum; (f), (g) and (h) P. rueppellii; (i) Nycticeinops schlieffeni (well above ground); (j) N. schlieffeni (< 3 m above ground); (k) Scotophilus dinganii (search-phase); (l) and (m) S. dinganii (approach-phase); (n) S. viridis; (o) and (p) Miniopterus fraterculus; (q) Myotis bocagii (flying near ground); (r) Myotis tricolor (flying near ground) and (s) Myotis welwitschii (flying near ground). See also Figure 27.

there are ‘whispering’ bats in the vespertilionid genus Kerivoula as there are in the Nycteridae and Megadermatidae, and some of the highflying vespertilionids such as Scotophilus and Scotoecus, emit shallow to very shallow FM search-phase calls that are remarkably similar to those emitted by high-flying Molossidae (Figure 27). A less obvious example concerns the use of Doppler shift to distinguish targets that are fluttering from those that are not. Rhinolophids and hipposiderids are well known to do this by perceiving Doppler-shifted changes in echoes of the CF-components of their calls when they are reflected from fluttering targets but, recently, it has been demonstrated that bats emitting FM calls may also be able to do this by detecting Dopplershifted ‘frequency smears’ in echoes reflected from fluttering targets (Taylor 1990). Because many vespertilionids forage in groups, their high-intensity echolocation calls, when made audible to our ears by a bat-detector, make an almost deafening racket of harsh staccato noises not unlike machine-gunfire. These bats emit their echolocation calls through the mouth, so individuals with bared teeth and threatening grimaces are usually just having a good ‘look’ at you with no intention to bite at all! It is unfortunate that so many artists make bats look savage and repulsive when, in fact, they are usually gentle and attractive. Most vespertilionids make vocalizations, in the contexts of communication and the expression of pain and/or fear, which are audible to humans as sounds such as squeaking, twittering, buzzing and nickering. Vespertilionids are moderately adapted for cursorial locomotion and are able to move over horizontal, sloping and vertical surfaces. However, with the exception of Miniopterus, they cannot furl their flight-membranes as effectively as emballonurids and molossids, and they do not normally scuttle or climb except when moving very short distances between a landing place and a roost-site. Some species have pads on the wrists and heels to facilitate climbing slippery banana leaves. Vespertilionids, as a family, exploit the full range of dayroosts known to be used by bats (but with limitations on where they roost within some of these day-roosts), and some utilize day-roosts that are not used or seldom used by bats of other families in Africa.

These include inside the furled leaves of banana plants (exploited by Pipistrellus nanus and Myotis bocagii), weaver-bird and sunbird nests (Kerivoula), holes in trees made by nesting woodpeckers and barbets (Scotophilus but also some molossids), and crevices in bark and spaces under loose sheets of bark (Barbastella, Mimetillus, and some species of Laephotis, Myotis, Nyctalus and Pipistrellus [and also some molossids]). With few exceptions, vespertilionids cling in contact with the sloping or horizontal surfaces of their day-roosts: very few ever roost hanging freely. Consequently, vespertilionids rarely compete with bats of other families that hang freely from ceilings. Vespertilionids roost singly or in small to very large groups, and group-members typically roost in close contact with each other. Many African vespertilionids go into torpor during the day, even in the tropics, but true hibernation is rare. Some exceptions include Barbastella barbastellus, Nyctalus leisleri, Pipistrellus pipistrellus, Myotis spp. and Miniopterus schreibersii, which hibernate in North Africa, and Myotis tricolor, Miniopterus fraterculus and M. natalensis, which hibernate in South Africa.The social behaviour and mating systems of African vespertilionids have received little attention: exceptions are Myotis bocagii and Pipistrellus nanus. Some appear to have solitary organizations, some establish pairs and harems and many live in small to very large groups of various compositions. Maternity colonies are established by many species.The reproductive strategies of African vespertilionids are not as diverse as might be expected, except in litter size. Most vespertilionids are monotocous but there are many species in Africa (and elsewhere) that have litters of 1–2, usually two, or 2–3 (and one non-African species, Lasiurus borealis, regularly has four, and occasionally five, young). All African vespertilionids (and the vast majority elsewhere) for which data are available, are seasonally monoestrous with the exceptions in Africa of Myotis bocagii, which is polyoestrus in Gabon, and Pipistrellus nanus, which is perhaps aseasonally monoestrous in some equatorial localities (assuming the data are not from several cryptic or misidentified species). Polyoestrous individuals have also been reported for Mimetillus molonyi and Pipistrellus rendalli (see species profiles), although the reproductive chronologies of 543

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these species are not yet fully understood. The advantages of timing spermatogenesis and lactation to coincide with seasons of maximum abundance of insects and/or optimal temperatures has led to the evolution of delays between copulation and parturition. These delays, which usually occur during the cold dry season or winter, include sperm storage and delayed fertilization (Myotis tricolor, Pipistrellus capensis, P. nanus), delayed implantation (Scotophilus leucogaster, Miniopterus fraterculus, M. natalensis, M. schreibersii and probably M. minor) and retarded embryonic development (M. schreibersii, Scotophilus viridis and perhaps P. rusticus) (see species profiles). The !! of some species carry their neonates while foraging but leave them in the roost when they become larger and heavier. The geological range of the family Vespertilionidae is middle Eocene to Recent in Europe, late Oligocene to Recent in North America, middle Miocene to Recent in Africa and Asia, Pleistocene to Recent in the West Indies, South America and Australasia, and Recent over the remainder of the present geographic range (Koopman 1984). Of the 91 species definitely known to occur in Africa, 48 species are only or mainly found in the tropics, 26 species occur in both tropical and temperate regions and 17 species (of which 16 are Palaearctic species) occur only in temperate regions. Of the 90 species for which data are available, 23 are only or mainly found in savannas, 22 only or mainly in forests, 20 in both forests and savannas, 15 in Mediterranean habitats, six in arid regions, and four in both savanna and arid regions. Thirty-eight species (42%) are found in montane habitats, but only five of these species have been found only in montane habitats. Because vespertilionids are very diverse in their wing morphology and flight, and in their feeding and roosting habits, it is not surprising that there are considerably more than one species at most localities (except localities in some arid regions), for example: four species along the Nile R. at or near Khartoum, six at Kikwit, DR Congo (rainforest), six at Luluabourg, DR Congo (rainforest–savanna mosaic), six on Zomba Plateau, Malawi (montane forest and grassland, 1500–1800 m), 12 in Taï N. P., Côte d’Ivoire (rainforest), 13 in Liwonde N. P., Malawi (miombo woodland) and 17 in Comoe N. P., Côte d’Ivoire (forest– savanna mosaic) (refs. Hayman et al. 1966, Koopman 1975, Happold & Happold 1997, Van Cakenberghe et al. 1999, Fahr & Kalko 2011). The family Vespertilionidae was placed by itself in the superfamily Vespertilionoidae by Simmons (1998) although some authors, including Teeling et al. (2005), also include the Molossidae in this superfamily. However, recent studies based on molecular data contradict some of the groupings based on morphological data and, pending resolution of the controversies, no chiropteran superfamilies are recognized by Simmons (2005). There is also some uncertainty about the monophyly of the Vespertilionidae: Miniopterus (currently in subfamily Miniopterinae) may represent a distinct family. Indeed, since this profile was submitted, Miller-Butterworth et al. (2007) have confirmed that Miniopterus does represent a distinct family (Miniopteridae) and, furthermore, Lack et al. (2010) concluded that Cistugo (placed here in the Vespertilionidae, subfamily Myotinae) also represents a distinct family (Cistugidae). The Vespertilionidae are divided into subfamilies but the number, composition and affinities of the subfamilies is perhaps not yet fully understood. However, Simmons (2005) recognizes six subfamilies of which four are represented in Africa, and they are profiled here because it was not possible to incorporate the recent recognition of the families Cistugidae and Miniopteridae (see above).

a

b

c

d

Figure 119. Variations in the number of upper incisors and cheekteeth on each side in African Vespertilionidae. Left: occlusal views of teeth on left side of upper jaw. Right: lateral views of labial sides of the same teeth. (a) Two incisors and six cheekteeth (e.g. Myotis). (b) Two incisors and five cheekteeth (e.g. Plecotus). (c) Two incisors and four cheekteeth (e.g. Glauconycteris). (d) One incisor and four cheekteeth (e.g. Scotophilus).

The characters by which the four subfamilies (sensu Simmons 2005) can be distinguished in Africa, and the African genera in each subfamily, are listed below and illustrated in Figures 116, 117 and 119. Vespertilioninae. Second phalanx of third finger ca. as long as the first phalanx or longer but not more than twice as long as the first; ears not funnel-shaped; tragus short and blunt or long and narrow but not sharply pointed; pelage not frizzled; four or five upper cheekteeth; baculum present. All genera except those listed below. Myotinae. Second phalanx of third finger ca. as long as the first; ears not funnel-shaped; tragus long and narrow but not sharply pointed; pelage not frizzled; six upper cheekteeth; baculum present. Cistugo, Myotis. Miniopterinae. Second phalanx of third finger ca. three times longer than the first; braincase high and rounded; ears not funnel-shaped, tragus short and blunt; pelage not frizzled; five upper cheekteeth; baculum absent. Miniopterus. Kerivoulinae. Second phalanx of third finger ca. as long as the first; braincase high and rounded; ears funnel-shaped with deep emargination below tip; tragus long, narrow and sharply pointed; pelage frizzled; six upper cheekteeth; baculum: no information. Kerivoula and (very dubiously) Phoniscus. Following Simmons (2005), the subfamilies are presented in the above order. The African genera in each subfamily are presented in alphabetical order irrespective of any relationships between them. Meredith Happold

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Subfamily VESPERTILIONINAE – Barbastelles, Serotines, Butterfly Bats, Long-eared Bats, Noctules, Pipistrelles, House Bats and others Vespertilioninae Gray, 1821. London Med. Repos. 15: 229.

Diagnostic combination of characters: second phalanx of third finger ca. as long the first or longer but not more than twice as long as the first (Figure 117); ears not funnel-shaped (Figure 116); tragus short and blunt, or long and narrow, but not sharply pointed (Figure 116); less than six upper cheekteeth (Figure 119).This subfamily includes 37 genera (Simmons 2005) of which 14 occur in Africa (Simmons 2005) but, of the latter, only 12 are presented here (Hypsugo and Neoromicia

being provisionally treated as Pipistrellus [sensu lato] for reasons given below – see genus Pipistrellus). The phylogenetic relationships of the genera in this subfamily have been investigated many times, numerous hypotheses have been advanced, and various tribes have been described (e.g. Hill & Harrison 1987, Menu 1987, Koopman 1994, Volleth & Heller 1994, Hoofer & Van Den Bussche 2001 and many others). Simmons (2005) recognizes seven tribes of which five are represented

Table 21.  Key to the African genera in the subfamily Vespertilioninae. Ears are said to be relatively short if 23–48% of FA, medium if 49–72% of FA and long if 73–97% of FA. Measurements and distributions pertain only to Africa. Ear length (mm) (relative length)

Number of lower cheekteeth

Number of upper incisors

Number of upper cheekteeth

Ears joined at base

6

2

5

Yes

5

2

5

Yes

5

2

5

No

5

2

4 or 5

No

5–16 (Short)

23–39

5

2

4

No

15–25 (Medium)

30–41

5

2

4

No

9–14 (Short)

27–31

5

2

4

No

6–16 (Short)

32–48

5

2

4

No

10–20 (Short)

34–54

5

1

4

No

30–41) (Long

50–65

5

1

4

No

11–24 (Short)

41–88

5

1

4

No

9–13 (Short)

28–35

5

1

4 or 5

No

9–15 (Short)

28–40

32–40 (Long) 13–18 (Short) 13–23 (Short)

FA (mm)

Miscellaneous

36–42

North Africa, Sudan, Eritrea, Ethiopia

36–44

North Africa

41–64

Fifth finger shortened (about = 4th metacarpal) North Africa Fifth finger not shortened (> 4th metacarpal) Head not flattened No body pattern; wings not reticulated Outer margin of ear not reaching mouth Head slightly flattened No body pattern; wings not reticulated Outer margin of ear not reaching mouth Head conspicuously flattened No body pattern; wings not reticulated Outer margin of ear not reaching mouth Head not flattened Most with body pattern of spots and/or lines, or reticulated wings Outer margin of ear reaching mouth Lower lip with lobe at posterior angle Head not flattened No body pattern; wings not reticulated Outer margin of ear not reaching mouth Lower lip without lobe at posterior angle

Genus Plecotus (3 species) Barbastella (2 species) Nyctalus (2 species) Pipistrellus (28 species) Laephotis (4 species) Mimetillus (1 species)

Glauconycteris (12 species)

Eptesicus (5 species) Otonycteris (1 species)

Penis relatively short Tragus relatively moderately long Posterior upper molar with two ridges Penis short (5.6–6.2 mm) Upper canine with anterior surface rounded not grooved Posterior upper molar with three ridges Anterior lower premolar ca. half height of posterior premolar Penis extremely long (ca. 9–16 mm) Upper canine with anterior surface flat and grooved Posterior upper molar with three ridges Anterior lower premolar at least two-thirds height of posterior premolar, usually subequal

Scotophilus (6 species)

Nycticeinops (1 species)

Scotoecus (2 species)

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in Africa: the total number of genera in these tribes is given in brackets, and the named genera are those represented in Africa. Eptesicini (3 genera): Eptesicus. Nycticeiini (8 genera): Nycticeinops, Scotoecus, Scotophilus. Pipistrellini (4 genera): Nyctalus, Pipistrellus (sensu stricto). Plectotini (6 genera): Barbastella, Otonycteris, Plecotus. Vespertilionini (13 genera): Glauconycteris, Hypsugo, Laephotis, Mimetillus, Neoromicia. However, because it is unlikely that relationships between genera are fully understood, tribal affinities are not included in most species profiles here. The status of some genera is also controversial and there have been many changes of opinion since Koopman (1993), including the following examples. (a) Following Hill & Harrison (1987) and others, Glauconycteris is recognized as a distinct genus instead of a subgenus of Chalinolobus. (b) Following Horáček & Hanák (1986), Hill & Harrison (1987) and others, Nycticeinops is recognized as a distinct genus instead of a subgenus of Nycticeus. (c) Following Hill & Harrison (1987), Menu (1987),Volleth & Tidemann (1991),Volleth et al. (2001) and others, many species traditionally in Eptesicus have been removed, with the result that, in Africa, Eptesicus is now considered to be represented by only five species (details in Genus Eptesicus). The species which have been removed from Eptesicus (brunneus, capensis,

grandidieri [as flavescens], guineensis, cf. melckorum, rendalli, somalicus, tenuipinnis and zuluensis), have been placed in Neoromicia by several authors, including Simmons (2005), but the placement of some of these species in Neoromicia is controversial (e.g. Hoofer & Van Den Bussche 2001), and so too is the placement of nanus in Neoromicia by (Simmons 2005). (d) Following Horáček & Hanák (1986) and others, savii, which was previously in Pipistrellus, is considered to belong to the genus Hypsugo, and this, and the generic status of Hypsugo, is well supported by morphological, chromosomal and molecular evidence. However, often in the absence of evidence other than morphology, many other species formerly in Pipistrellus have now been placed in Hypsugo (see Simmons 2005) and this also is controversial. Therefore, pending resolution of the species contents of Neoromicia and Hypsugo, and also for convenience, all of the Pipistrellus-like species in Africa are presented here as species of Pipistrellus (sensu lato).With this exception, the African species in the subfamily Vespertilioninae are placed in the genera recognized by Simmons (2005), but some genera (Plecotus, Pipistrellus) contain new species, and some forms given specific status by Simmons (2005) are considered here to be subspecies. The 12 genera of Vespertilioninae represented in Africa can be distinguished by the characters given in Table 21. In the text, they are presented in alphabetical order irrespective of any relationships that might exist between them. Meredith Happold

GENUS Barbastella Barbastelles Barbastella Gray, 1821. London Med. Repos. 15: 300. Type species:Vespertilio barbastellus Schreber, 1774.

support the relationship with Plecotus, but not with Otonycteris (Hoofer & Van Den Bussche 2001). Simmons (2005) places Barbastella in the tribe Plecotini (with both Otonycteris and Plecotus, and three nonAfrican genera). Some authors consider that Barbastella is monotypic (but see profile of B. leucomelas). Barbastella is distributed from North Africa through Europe, SW Asia, N India to W China and also Japan (Koopman 1994). In Africa, both species have been found in semidesert habitats but B. barbastella is also found in cool montane habitats in NW Africa, whereas B. leucomelas has been found only at low altitudes (and probably the only records in Africa are those of vagrants). The two species are: Barbastella barbastellus.

A polytypic genus with two species, both found in North Africa. Diagnostic combination of characters: five upper and five lower cheekteeth and two upper incisors on each side; ears joined at base (as in Plecotus) but short (cf. Plecotus); tympanic bullae small (cf. large in Plecotus). Dental formula 2123/3123 = 34 (as in Nyctalus and most African Pipistrellus). Selected characters of Barbastella are illustrated in Figure 120. Morphologic, bacular and karyotypic data suggest close affinity with Plecotus, Otonycteris and about three other non-African genera (e.g. Hill & Harrison 1987, Frost & Timm 1992, Tumlison & Douglas 1992, Qumsiyeh & Bickham 1993). Mitochondrial ribosomal sequences

B. barbastellus. Pelage, wings and interfemoral membrane pale; ventral pelage pale fawn to off-white; condylobasal length 13–14 mm; tympanic bullae smaller; auditory meati smaller; mastoid region less inflated and without a prominent projecting flange (Figure 121a); outer margin of ear usually with a prominent projecting lobe (Figure 120). B. leucomelas. Pelage, wings and interfemoral membrane dark, ventral pelage dark grey; condylobasal length 14–15 mm; tympanic bullae larger; auditory meati larger; mastoid region noticeably inflated and with a prominent projecting flange (Figure 121b); outer margin of ear normally without projecting lobe. Meredith Happold

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Barbastella barbastellus

Figure 120. African Barbastella. Flight membranes, and bones of wing, hindlimb and tail of B. barbastellus. Left eye, ear and tragus of B. barbastellus: note small lobe projecting from middle of outer margin of ear (absent in B. leucomelas). Skull of B. barbastellus (ZFMK 61.200 from Spain). Occlusal view of upper teeth of B. barbastellus.

BELOW:

Figure 121. The tympanic regions (left side) of (a) Barbastella barbastellus and (b) B. leucomelas (after Harrison & Makin 1988).

a

b

Barbastella barbastellus WESTERN BARBASTELLE Fr. Barbastelle d’Europe; Ger. Mopsfledermaus Barbastella barbastellus (Schreber, 1774). Die Säugethiere 1: 168. Burgundy, France.

Taxonomy Originally Vespertilio barbastellus. Synonyms: barbastelle, communis, daubentonii. Subspecies: none. Possibly includes leucomelas but, pending further information, B. leucomelas is treated as a distinct species here (see profile for B. leucomelas). Chromosome number (Turkey): 2n = 32; aFN = 50 (Volleth 1985).

Description Very small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; five upper and five lower cheekteeth and two upper incisors on each side; dorsal pelage silvery-black; wings blackish-brown to black; ears joined at base, short for a vespertilionid (13–14 mm), normally with projecting lobe on 547

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outer margin. Pelage silky, dense; mid-dorsal hairs ca. 10 mm. Dorsal pelage dark blackish-brown to black with silvery sheen; hairs dark blackish-brown or black with whitish or yellowish tip. Ventral pelage dark grey; hairs dark blackish-brown with greyish-white to beige tip. Chin, throat and chest darker than belly and flanks. Dorsal pelage of young up to 12 months old is black without silvery sheen. Muzzle short, flat and broad; nostrils opening upwards and outwards between two high, lateral glandular swellings and behind prominent median pad. Ears (Figure 120) blackish, relatively short for a vespertilionid (40.1 [36–46]% of FA, n = 12) (cf. relatively long in Plecotus and moderately long in Otonycteris); forward-facing, inner margins joined at base by low frontal band, outer margin conspicuously notched near base and usually with a small lobe projecting from the middle of the margin (cf. B. leucomelas). Ears fold back close to head when bat rests. Tragus reaching just over half height of ear, triangular with long rounded tip, covered with fine hairs.Wings and interfemoral membrane dark brown to black (cf. B. leucomelas). Tail 91.6 (70–110)% of HB (Europe). Skull (Figure 120) with rostrum narrow and weak; braincase elongated and rounded; zygomatic arches slender and without marked dorsal curvature; zygomatic width < mastoid width. Sagittal crest absent or very low; lambdoid crest weakly developed.Tympanic bullae comparatively small (cf. Plecotus); smaller than in B. leucomelas (Harrison & Makin 1988). Auditory meati comparatively small (cf. B. leucomelas); mastoid region less inflated and without a prominent projecting flange (Figure 121a). Inner incisor bicuspid; outer incisor smaller, not reaching level of secondary cusp of inner incisor. Upper canines slender.Two upper premolars on each side; anterior premolar minute (shorter than cingulum of canine), displaced lingually; canine and posterior premolar in contact. Lower incisors (three on each side) tricuspid, arranged in a V-shape with the outer pair the largest. Two lower premolars and three lower molars on each side.

Barbastella barbastellus

Abundance Uncommon throughout much of the African geographic range.

Adaptations Aspect ratio low; wing-loading low (Norberg & Rayner 1987). Based on wing morphology, flight predicted to be slow with great manoeuvrability. In Europe, roosts by day in summer in roof-spaces and cracks in buildings, behind wooden windowshutters, holes in trees and behind bark. Hibernates in winter in caves, mines and fissures. Tolerates low ambient temperatures (ca. Geographic Variation Uncertain. If new evidence shows B. –5 to +5 °C) and low as well as high relative humidities (40–96%) leucomelas is conspecific, it might be considered a subspecies of B. (Bogdanowicz & Urbanczyk 1983, Weidner 2000). Migratory barbastellus. movements as far as 290 km were reported by Kepka (1960). For more information, see Rydell & Bogdanowicz (1997). Similar Species The only other African vespertilionids that have ears with inner margins joined are: Echolocation The echolocation calls of B. barbastellus are unique among Palaearctic bats (Ahlén 1981, 1990). Search-phase calls are of Barbastella leucomelas. Outer margin of ear normally without small two types – one loud and one weak. (a) Call-shape short CF/FM; calls projecting lobe; ear longer (15–18 mm) and narrower. Pelage, beginning with a short (1–1.5 ms) CF or narrow bandwidth component, wings and interfemoral membrane pale; ventral pelage pale and ending with a short downward FM sweep; start-frequency 35 kHz, fawn to off-white. Tympanic bullae and especially the auditory end-frequency 28 kHz; call-duration 4 ms; intensity high. (b) Callmeati larger; mastoid region more inflated and with a prominent shape short CF/FM as above or FM; start-frequency 43 kHz; endprojecting flange; lambdoid crest less evident. frequency 33 kHz; call-duration 5.2 ms; intensity low. Highest call Plecotus spp. Ears much longer (32–40 mm). Six lower cheekteeth intensity at 30–35 kHz or 40–43 kHz (Schober & Grimmberger 1987, on each side. Ahlén 1990). It is suggested that the bat alternates between the loud and weak calls (Jones 1993). Call repetition-rate: 8–9 calls per second. Calls Distribution In Africa, this Palaearctic species occurs only in are emitted through the nose as well as through the mouth. the mountains of the Mediterranean Coastal and Afromontane– Afroalpine BZs in Morocco (Rif, Middle Atlas, High Atlas) (Panouse Foraging and Food No information from Africa. In Europe, 1955, Ibáñez 1988, Fonderflick et al. 1998). Extralimitally: England forages by slow-hawking above the canopy in forested areas and and W Europe to Caucasus, also Canary Is. and larger islands in low over water. Has been reported to glean from foliage but this is Mediterranean (Simmons 2005). doubtful. Small moths make up 70–99% of prey by volume (based on analysis of faeces from Europe and C Asia; Beck 1995, Rydell et al. Habitat In Africa, inhabits cool mountains and semi-desert areas. 1996, Sierro & Arlettaz 1997). Has a small gape and weak dentition, Associated with forested areas in Europe. therefore its diet does not include large, hard-shelled prey. 548

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Barbastella leucomelas

Social and Reproductive Behaviour In Europe, hibernates singly, or in clusters in colonies of up to 1000 individuals. In summer, sexes segregate; !! form maternity colonies of 5–80 individuals; "" roost in small groups (Schober & Grimmberger 1987, Richarz 1989). Mating takes place in autumn, when the bats return to the hibernacula. Reproduction and Population Structure Litter-size: normally one, rarely two. Reproductive chronology in Africa not known; in Europe, it is seasonal monoestry with mating in autumn (and sometimes in hibernacula); births from mid-Jun (Schober & Grimmberger 1987). Maximum life-span 23 years. In many hibernacula, there is a predominance of "" (Schober & Grimmberger 1987). Predators, Parasites and Diseases Predators in Europe include owls Asio otus, Tyto alba, Strix aluco, Bubo bubo and the Beech Marten Martes foina (Abeijentsev et al. 1956, Urbanczyk 1981). Ectoparasites in Europe include fleas (Siphonaptera) and bat-flies (Diptera), both of which are found occasionally, and mites (Acari), which heavily infest this bat. Conservation IUCN Category: Near Threatened. Population fragmented and predicted to decline significantly within next 15 years. Has specific habitat requirements and its habitats are declining.

Measurements Barbastella barbastellus FA: 38.1 (36–40) mm, n = 6 WS: n. d. TL: 102.1 (97–107) mm, n = 8* T: 50.1 (45–56) mm, n = 8* E: 13.5 (13–14) mm, n = 4 E: 15.7 (15–18) mm, n = 8* Tr: n. d. Tib: n. d. HF: n. d. WT: 7.5, 8 g, n = 2 GLS: 15, 15 mm, n = 2 GWS: n. d. ZW: 7.6, 7.6 mm, n = 2 C–M3: 4.6 mm, n = 1 Morocco (Panouse 1955, Ibáñez 1988, Fonderflick et al. 1998) *Spain (ZFMK) Key References Rydell & Bogdanowicz 1997; Sierro 1999; Sierro & Arlettaz 1997. Antoine Sierro

Barbastella leucomelas EASTERN BARBASTELLE Fr. Barbastelle asiatique; Ger. Östliche Mopsfledermaus Barbastella leucomelas (Cretzschmar, 1826). In: Rüppell, Atlas Reise Nordl. Afr., Zool. Säugeth., p. 73. Sinai, Egypt.

Taxonomy Originally Vespertilio leucomelas. Synonyms: blanfordi, caspica, darjelingensis, walteri. Subspecies: two, of which probably only the nominate occurs in Africa. Considered a subspecies of Barbastella barbastellus by Qumsiyeh (1985), but not by Corbet (1978), Harrison & Makin (1988), Harrison & Bates (1991) and Corbet & Hill (1991) and, pending further study, Horáček et al. (2000) and Simmons (2005). Chromosome number (Japan): 2n = 32; aFN = 50 (Andō et al. 1977). Description Very small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; five upper and five lower cheekteeth and two upper incisors on each side; dorsal pelage silvery-fawn over dark brown; wings pale; ears joined at base, medium-short for a vespertilionid (15–18 mm), without projecting lobe on outer margin. Sexes similar. Pelage silky, dense; mid-dorsal hairs ca. 10 mm. Dorsal pelage dark brown with conspicuous silvery-fawn to golden-beige sheen; hairs dark chocolate brown with fawn to beige at tip. Ventral pelage: pale fawn to off-white (throat occasionally appears darker); hairs dark brown with terminal third pale fawn to off-white. Muzzle short, flat and broad; nostrils opening upwards and outwards between two high, lateral glandular swellings and behind prominent median pad. Ears relatively short (40.2 [36– 46]% of FA, n = 4) (cf. relatively long in Plecotus and moderately long in Otonycteris), forward-facing, joined at base by low frontal band; outer margin conspicuously notched near base and without small projecting lobe (cf. B. barbastella). Ears fold back close to head when bat rests. Tragus long with covering of fine hairs; tip rounded.

Wings and interfemoral membrane pale brown, semi-translucent (cf. B. barbastellus). Tibia 47.8 (46–51)% of FA, n = 4. Tail 105 (96– 119)% of HB, n = 4. Baculum (Turkmenistan) lanceolate in dorsal view; proximal part comparatively wide, stout and with well-marked furrow (cf. B. barbastella from Europe) (Strelkov 1987). Skull with rostrum narrow and weak; braincase elongated and rounded; zygomatic arches slender and without marked dorsal curvature; zygomatic width < mastoid width. Supraorbital ridges poorly developed; sagittal crest very low or absent; lambdoid crest very weakly developed and less evident than in B. barbastella. Tympanic bullae comparatively small (cf. Plecotus), but larger than in B. barbastella (Harrison & Makin 1988). Auditory meati comparatively large (cf. B. barbastella); mastoid region inflated and with a prominent projecting flange (Figure 121b). Dentition as in B. barbastella. Geographic Variation

None in Africa.

Similar Species The only other vespertilionids in Africa that have ears with inner margins joined are: Barbastella barbastellus. Outer margin of ear normally with small projecting lobe ca. half-way between base and tip; ear shorter (13– 14 mm) and broader. Pelage, wings and interfemoral membrane dark, ventral pelage dark grey. Tympanic bullae and especially the auditory meati smaller; mastoid region less inflated and without projecting flange; lambdoid crest more evident. 549

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Habitat No details available for the African records other than that Massawa is near the coast and in the semi-desert grassland and shrubland vegetation zone (terminology: White 1983). Abundance Very rarely recorded in Africa. Has not been recorded in Sinai since the species was described (Corbet 1978). Remarks Nothing is known about the biology of this species in Africa and there is very little information from elsewhere. Flight heavy and fluttering; sometimes close to ground, sometimes high (Roberts 1977). In Pakistan, roosts by day in crevices under bark and in tree hollows during summer (Roberts 1977). Also roosts in caves, tunnels, mine-shafts and old deserted houses (Cretzschmar 1826, Ognev 1928, Lay 1967). Roosts and forages singly (Roberts 1977). Litter-size (Eastern Europe and northern Asia): one or two (Ognev 1928). Conservation IUCN Category: Least Concern (assessment based mainly on extralimital data). Barbastella leucomelas

Plecotus spp. Ears much longer (32–40 mm). Six lower cheekteeth on each side. Distribution In Africa, the nominate subspecies has been recorded from Sinai (Egypt) and from the Somalia–Masai Bushland BZ in Eritrea, but the reports are few and far between and it seems probable that they are records of vagrants. Von Heuglin (1862, 1877) listed this species from Massawa, Eritrea, and there is a specimen from ‘Erythrea’ in the MCZ, and the holotype is from Sinai. Barbastella leucomelas was also recorded from Senegal by de Rochebrune (1883), but there have been no further records of any Barbastellus from Senegal (Hayman & Hill 1971): furthermore, this record is more likely to refer to B. barbastella than B. leucomelas. Extralimitally: the nominate subspecies is known from Arabia and Iran, and B. l. darjelingensis is known from S Israel and Iran to N India, Nepal, W China and Japan.

Measurements Barbastella leucomelas FA: 39 (37–44) mm, n = 10 WS: n. d. TL: 100 (92–107) mm, n = 4 T: 46 (33–58) mm, n = 8 E: 16 (15–18) mm, n = 4 Tr: 9.1, 14 mm, n = 2 Tib: 18.2 (17–20) mm, n = 4 HF: 6.6 (6–7) mm, n = 4 WT: n. d. GLS: 15.3 (14.6–16.0) mm, n = 5 GWS: 8.3 (8.0–8.5) mm, n = 5 ZW: 7.7 (7.4–8.1) mm, n = 3 C–M3: 4.6 (4.3–4.9) mm, n = 8 Arabia, Israel, Iran (BMNH, HZM, SMF, Kock 1969a, Harrison & Makin 1988) Key Reference

Harrison & Bates 1991. Meredith Happold

GENUS Eptesicus Serotines Eptesicus Rafinesque, 1820. Ann. Nature, p. 2. Type species: Eptesicus melanops Rafinesque, 1820 (= Vespertilio fuscus Beauvois, 1796).

A polytypic genus, considered by Hill & Harrison (1987) to contain 19 species, and by Simmons (2005) to contain 23 species of which five occur in Africa (see subfamily Vespertilioninae). Diagnostic combination of characters for African members: invariably four upper and five lower cheekteeth and two upper incisors on each side; ears comparatively and relatively short and not joined at base; 5th finger not shortened (considerably longer than metacarpals of 3rd and 4th fingers) (cf. Nyctalus); no body pattern; wings dark coloured, not reticulated; outer margin of ear not reaching mouth;

lower lip without lobe near posterior angle (cf. Glauconycteris); FA: >37.8 mm (except E. floweri); GLS: >15.0 mm. Dental formula: 2113 /3123 = 32. Selected characters of African Eptesicus are illustrated in Figure 122. Species of Pipistrellus (sensu lato) sometimes share most of these characters but African Eptesicus (except E. floweri) are almost always larger (FA: 38–51 mm; GLS: 15.3–21.3 mm) than African Pipistrellus (FA: 23–39 mm; GLS: 10.0–15.4 mm). Furthermore, species of Eptesicus invariably have only one upper premolar on each side whereas, in contrast, 20 of the 28 African species of Pipistrellus

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d a

b

Figure 122. African Eptesicus. (a) Approximate outline of flight membranes, and bones of wing and hindlimb of E. serotinus (traced from dried specimen from Iran; tail vertebrae not distinguishable). (b) Left ear and tragus of E. serotinus. (c) Tragus of E. hottentotus. (d) and (e) Lateral and dorsal views of the skull of E. hottentotus (BMNH 22.12.17.39 from Malawi). (f) Occlusal view of upper teeth of E. hottentotus.

c

have two upper premolars on each side (although the anterior upper premolar is often minute and therefore sometimes not visible above the gum, and can be absent in some individuals). Also, in Eptesicus, the baculum (Figure 123) is more-or-less triangular (cf. variable but not triangular in Pipistrellus), and the chromosome number (2n) is 50 (cf. variable but always less than 50 in Pipistrellus). a

b

c

d

e

Figure 123. Bacula of African Eptesicus: dorsal views on left, lateral views of right side on right, tips uppermost. (a) Eptesicus bottae innesi from Cairo. (b) E. hottentotus from KwaZulu Natal. (c) E. floweri lowei from Sudan. (d) E. serotinus isabellinus from Morocco. All based on Hill & Harrison (1987). All drawn to same size (for scale drawings, see Hill & Harrison 1987).

f

Eptesicus was traditionally considered to contain ca. 33 species world-wide (Koopman 1993) and, traditionally, the generic distinction between Eptesicus and Pipistrellus was based on the absence or presence, respectively, of a second upper premolar (the anterior upper premolar). However, this proved unreliable because tooth reduction appears to have taken place independently in several lineages of the Vespertilionidae. Based on bacular morphology, Hill & Harrison (1987) retained only 19 species in Eptesicus, and placed the remaining species into Neoromicia (nine species; Africa only) and Vespadelus (five species; Australia only), both of which were considered to be subgenera of Pipistrellus. This rearrangement is supported by dental morphology (Menu 1987) and chromosomes (Heller & Volleth 1984, Volleth & Tidemann 1991, Volleth et al. 2001, Kearney et al. 2002). For further details, see genus Pipistrellus. Of the species that remain in Eptesicus, five occur in Africa: E. bottae, E. floweri, E. hottentotus, E. platyops and E. serotinus. The relationships between these taxa are uncertain (Rosevear 1962, Kock 1969a, Ibáñez & Valverde 1985, Schlitter & Aggundey 1986) but, pending revision of the genus, we follow Koopman (1993) and Simmons (2005), and tentatively recognize all of these taxa as distinct species. Eptesicus – as currently conceived – is distributed almost worldwide (with the exception of the islands of South-East Asia and Australia), in a wide variety of tropical, sub-tropical and temperate forests, woodlands, savannas and deserts. In Africa, one species is found mainly in the Rainforest BZ (Western Nigerian sub-region and Bioko I.), two mainly in savannas, one in the Sahara Arid BZ and one mainly in the Mediterranean Coastal BZ. No members of this genus are known from the Congolian rainforest. 551

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Table 22.  Key to the African species in the genus Eptesicus. Relative heights of the braincase are based on height as a percentage of GLS (details in Genus Pipistrellus and in Eptesicus species profiles). Measurements and distributions pertain only to Africa. Inner upper incisor

Ventral hairs

FA (mm)

GSL (mm)

Unicuspid

Bicoloured

48.2 (45–54)

16.9–21.5

Bicuspid

Bicoloured

45, 50

18.8

Bicuspid

Bicoloured

42.2 (38–47)

15.3–18.0

Bicuspid

Unicoloured

48.7 (45–52)

18.1–20.1

Bicuspid

Unicoloured

36.5 (34–38)

12.0, 13.5

Miscellaneous

Species

Braincase relatively low Savannas of southern and eastern Africa Braincase relatively high Nigeria, Bioko I. and ? Senegal Braincase relatively high Egypt (Cairo district) Braincase of medium relative height Morocco, Algeria and Tunisia (except deep Sahara), to NW Libya Unique horny warts on forearms, legs and tail Braincase relatively high Sudan south of Sahara

The species occurring in Africa can be distinguished by the characters and distributions given in Table 22. Further information is given in the Similar Species sections in which comparisons are made only between species of Eptesicus and Pipistrellus that occur in Africa.

E. hottentotus E. platyops E. bottae E. serotinus

E. floweri

For other characters and explanations of terminology, see genus Pipistrellus. Victor Van Cakenberghe & Meredith Happold

Eptesicus bottae  Botta’s Serotine Fr. Sérotine de Botta; Ger. Bottas Breitflügelfledermaus Eptesicus bottae (Peters, 1869). Monatsber. K. Preuss. Akad. Wiss. Berlin 1869: 406.Yemen. Nader & Kock (1990) limit the type locality to the area between Hodeida, Hays, Ta’izz and Al Muhka in SW Yemen.

Taxonomy  Originally Vesperus bottae. Sometimes considered to belong to E. serotinus or E. hottentotus, but Harrison (1964) corrected this misconception. Synonyms: innesi and others extralimitally (Koopman 1993). Subspecies: six of which only one, E. b. innesi, occurs in Africa (Koopman 1994). Chromosome number (Soviet Central Asia, Greece): 2n = 50; aFN = 48 (Zima & Horáček 1981, Volleth et al. 2001). Description  Small to very small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; four upper and five lower cheekteeth and two upper incisors on each side; ears separated and short (13–18 mm); medium-sized for an African Eptesicus (FA: 38–47 mm); ventral hairs bicoloured; inner upper incisor bicuspid. Sexes similar in colour; // on average slightly larger than ??. Pelage soft, dense; mid-dorsal hairs 7–10 mm. Dorsal pelage dark cream; hairs with dark grey at base. Ventral pelage much paler; hairs whitish with pale greyishbrown bases. Ears brown to blackish-brown, relatively short, subtriangular with tip rounded. Tragus length ca. half of E; breadth almost constant over the entire length, tip blunt. Eyes very small. Wings and interfemoral membrane dark brown to blackish, semitranslucent; wings without white hind-border but sometimes with very narrow pale border. Tail protruding ca. 3.5 mm beyond membrane. Postcalcareal lobe well developed. Baculum short, flat, stout, triangular with rounded corners (Figure 123a; Hill & Harrison 1987) and see also Spitzenberger (1994).

Skull medium-sized for an African Eptesicus. Braincase relatively high (43% of GLS, n  = 1) and broad (47% of GLS, n  = 1). Rostrum fairly broad, flattened dorsally. Profile of forehead region (viewed laterally) weakly concave (for definition, see genus Pipistrellus). Occipital helmet present, sagittal and lambdoid crests moderately developed for this genus. Inner upper incisor large, bicuspid. Outer upper incisor very small (only just reaching above cingulum of inner incisor). Anterior upper premolar absent; canine and posterior premolar in contact. Lower molars myotodont (Menu 1987). Geographic Variation  None in Africa. Similar Species  Six other vespertilionids in North Africa have (on each side) two upper incisors, five lower cheekteeth and only one upper premolar or (at least sometimes) only one that is clearly visible above the gum: Eptesicus serotinus. Ventral hairs unicoloured. Usually larger (FA: 48.7 [45–52] mm; GLS: 18.1–20.1 mm). Morocco to NW Libya. Pipistrellus (5 spp.). Smaller (FA: 24–38 mm; GLS: 10.8–14.6 mm). Anterior upper premolar usually present although not always visible above gum. For other African Eptesicus, see Table 22, p. 552.

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Eptesicus bottae

Foraging and Food No information for Africa. Elsewhere, E. bottae forages by slow-hawking, rather high above ground (Harrison & Bates 1991). Begins foraging at dusk in some localities, but at some time after dusk at others. In Kurdistan, has been seen feeding on numerous insects gathered around an electric light (Harrison & Bates 1991). In Jordan, recorded flying over a pool, but it is not clear if the bats were foraging or coming to drink. In Israel, feeds mainly on Hymenoptera and Lepidoptera (Feldman et al. 2000). Echolocation Search-phase call-shape (Israel) QCF. Startfrequency ca. 41 kHz; end-frequency and peak-frequency (mean ± S.D.) 32.5 ± 0.87 kHz; call-duration 6.9 ± 1.32 ms; intercallinterval 155.6 ± 61 ms (134–143 calls, Holderied et al. 2005). The loudest calls emitted had an intensity of 121 ± 7.8 dB at 10 cm, and it was calculated from this that E. bottae would be able to detect large prey and conspecifics up to a distance of 21 m. Social and Reproductive Behaviour

Eptesicus bottae

Distribution In Africa, known only from the Sahara Arid BZ in the Cairo area of Egypt. Extralimitally: the subspecies in Egypt (E. b. innesi) occurs in Israel and Jordan (Harrison & Bates 1991); the other subspecies are recorded from Rhodes (Greece), Turkey, SW and SE Arabia, and from Iraq, Iran and Pakistan north-eastwards to NW China and Mongolia (Gaisler 1970, Harrison & Bates 1991, Simmons 2005). Le Berre (1990) mentions the presence of E. bottae in Morocco based on a record of E. isabellinus from Heim de Balsac (1948), but this record probably represents E. serotinus isabellinus, which occurs in NW Africa. Habitat In SW Asia, apparently restricted to the deserts and semi-desert regions, but probably only occurs in localized areas, such as farmlands near water, where insects and suitable day-roosts are available. In Israel, has been observed ‘flying over cultivated fields lined by eucalyptus and tamarisk trees, surrounded by desert’ (Harrison & Bates 1991). Feldman et al. (2000) mention edges of vegetation and cliffs as the preferred habitat in Israel. In Iraq, found in gardens and date-groves near rivers, but not encountered in desert localities (Harrison & Bates 1991). In Oman, recorded flying over rocky mountain slopes at 1880–2100 m (Harrison & Bates 1991). Abundance Apparently not very common anywhere. Based on museum records and the literature, the African subspecies is probably rare or very rare. Adaptations No information for Africa. In Arabia, E. bottae is reported to fly strongly and to be noisy in flight (Harrison & Bates 1991). In Israel, flight-speed of 5.70 m/sec was recorded by Holderied et al. (2005). In Iraq, has been found roosting by day in buildings, including in crevices beneath the corrugated iron roofing of verandahs (Holderied et al. 2005).

No information.

Reproduction and Population Structure No information for Africa. From Israel, there are records of two !!, each with twin embryos (crown–rump length ca. 14 mm), in Apr (Harrison & Bates 1991). Predators, Parasites and Diseases Nader & Kock (1989) were unable to find ectoparasites and, although they do not exclude the possible loss of parasites during transport, they suggest that the lack of parasites might be the result of specific nesting sites or habits. This invites investigation. Conservation extralimitally).

IUCN Category: Least Concern (Africa and also

Measurements Eptesicus bottae FA: 42.2 (38–47) mm, n = 54 WS: 281.6 ± 7 mm, n = 6* TL: n. d. T: 39.5 (27–50) mm, n = 43 E: 15.6 (13–18) mm, n = 60 Tr: 6.5 (3.9–8.8) mm, n = 50 Tib: 16.7 (13–20) mm, n = 51 HF: 8.8 (7–10) mm, n = 24 WT: 14.0 (14.0–14.0) g, n = 3 GLS: 16.6 (15.3–18.0) mm, n = 46 GWS: 10.9 (10.0–12.7) mm, n = 43 C–M3 (alv.): 6.1 (5.6–6.8) mm, n = 47 Throughout geographic range (BMNH, ZMB) *Mean ± S.D.; Israel (Holderied et al. 2005) Key References Feldman et al. 2000; Harrison & Bates 1991; Nader & Kock 1989. Victor Van Cakenberghe & Meredith Happold

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Eptesicus floweri HORN-SKINNED SEROTINE Fr. Sérotine de Flower; Ger. Warzen-Breitflügelfledermaus Eptesicus floweri (de Winton, 1901). Ann. Mag. Nat. Hist., ser. 7, 7: 46. Wad Marium, Khartoum, Sudan.

Taxonomy Originally Glauconycteris floweri. This species has unique horny warts on the forearms and consequently has been placed in a separate subgenus, Rhinopterus, by some authors, including Hayman & Hill (1971), Hill & Harrison (1987) and Koopman (1993, 1994). Synonyms: lowei. Subspecies: none recognized but see Geographic Variation. Chromosome number: not known. Description Very small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; four upper and five lower cheekteeth and two upper incisors on each side; ears separated and short (ca. 10 mm); the smallest African Eptesicus (FA: 34–38 mm); ventral hairs unicoloured; inner upper incisor bicuspid; unique horny warts on forearms, legs and tail. A poorly known species. Dorsal pelage yellowish-fawn to pale rusty-brown; hairs unicoloured. Ventral pelage yellowish-brown on throat, becoming paler or pure white on abdomen; hairs unicoloured. Muzzle and cheeks, naked, dark brown. Ears dark brown, relatively short, subtriangular with tip rounded. Tragus length ca. 40% of E; anterior margin straight; posterior margin convex with lobe at base; broadest at mid-height. Wings and interfemoral membrane pale brown, wings of holotype of floweri (and one other specimen) with very faint pale hind-border, a third specimen with a conspicuous white hind-border (no other information); horny warts (denticles) on arm-wings and anterior three-quarters of interfemoral membrane give the membranes a spotted appearance. The horny warts on the dorsal surfaces of the forearms, tibiae and tail are characteristic of the monotypic subgenus Rhinopterus. Postcalcareal lobe fairly well developed. Tail ca. 76% of HB. Baculum short, flat, roughly triangular (Figure 123c, Hill & Harrison 1987). Skull small for an African Eptesicus. Braincase relatively high (42, 43% of GLS, n = 2) and broad (47, 50% of GLS, n = 2). Rostrum short and broad; dorsal surface almost completely flat with angular margins (therefore clearly distinguishable from the lateral surfaces). Profile of forehead region (viewed laterally) almost straight. No occipital helmet; sagittal crest moderate, low or absent; lambdoid crests moderate to weakly developed. Inner upper incisor bicuspid. Outer upper incisor very small, barely exceeding cingulum of inner incisor. Anterior upper premolar absent; canine and posterior premolar in contact. Lower molars myotodont (G. Bennell pers. comm.). Geographic Variation No subspecies recognized by Koopman (1994) and Simmons (2005). Braestrup (1935) regarded E. floweri as a variable species, there being a smaller form with ventral pelage buff (floweri) and a larger form with ventral pelage whitish (formerly lowei). Both forms occur sympatrically in some parts of Sudan, so the variation is apparently not geographic. See also Distribution. Similar Species No other African vespertilionid has horny warts. The South African notius was described as a bat with warts and referred to Rhinopterus, but it is probable that the warts of this individual were caused by nematodes and this form is now considered a subspecies of Pipistrellus capensis (Koopman 1975).

Eptesicus floweri

Distribution Endemic to Africa. Known with certainty only from the Sahel Savanna and Sudan Savanna BZs in Sudan (Kock 1969a, Koopman 1975). A specimen from Timbuktu, Mali, is either a small representative of E. floweri or a separate form (Braestrup 1935). In this Malinese specimen, the inner upper incisor is unicuspid and the outer upper incisor is two-thirds of the height of the inner (cf. the Sudanese E. floweri described above), suggesting this specimen represents a separate form (E. Thorn pers. comm.). Recently, two specimens (EBD 26179-80) were reported from Mahmûdé L.,Wilaya Hodh Ech, Mauritania (16° 29.969' N, 07° 42.911'W) by Padial & Ibáñez (2005); not mapped. The FA lengths of these specimens are 33.5 mm (!) and 33.2 mm ("); therefore they are slightly smaller than specimens from Sudan and Mali (see Measurements). Habitat Recorded from Acacia wooded grassland and deciduous bushland, semi-desert grassland and shrubland, and mosaics of edaphic grassland with Acacia wooded grassland (terminology:White 1983). Two were shot beside a pool in Sudan (NMW). Possibly limited to habitats with Acacia spp. (Koopman 1975). The recent specimens from Mauritania were also found in ‘Acacia formations of the Sahel’ (Padial & Ibáñez 2005). Abundance Uncertain. De Winton (1901) reported it common at Shendi in N Sudan. Rarely collected. Remarks Flies slowly and apparently forages near the ground amongst the dense and thorny Acacia bushes. Has also been observed flying low over ponds (von Wettstein 1917). Roosts by day in Acacia

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Eptesicus hottentotus

thickets – low down near the roots of these trees (de Winton 1901). At dusk, individuals crawl up the branches and take flight, uttering a very characteristic squeak, which they continue to make during flight. The horny warts on the wings are also characteristic of the African molossid Platymops setiger, and the non-African Eptesicus nasutus, suggesting an ecological parallel with these species (E. Thorn pers. comm.). Conservation IUCN Category: Least Concern. Acacia senegalensis, the widely distributed tree that provides the roosting sites of this species in Sudan, is a valuable source of Gum Arabic, but the extent to which harvesting of Gum Arabic has affected the abundance of E. floweri is not known (D. Kock pers. comm.).

TL: 79 or 83* T: 34 or 36* E: 10 mm, n = 1 Tr: 4.1 mm, n = 1 Tib: 12 mm, n = 1 HF: n. d. WT: n. d. GLS: 12.0, 13.5 mm, n = 2 GWS: 8.4 mm, n = 1 C–M3 (alv.): 4.4, 4.9 mm, n = 2 Sudan (AMNH, BMNH and literature) *de Winton 1901 and Thomas 1915b Key References de Winton 1901; Kock 1969a; Padial & Ibáñez 2005;Thomas 1915b.

Measurements Eptesicus floweri FA: 36.5 (34–38) mm, n = 5 WS: n. d.

Victor Van Cakenberghe & Meredith Happold

Eptesicus hottentotus LONG-TAILED SEROTINE Fr. Sérotine hottentote; Ger. Hottentotten-Breitflügelfledermaus Eptesicus hottentotus (A. Smith, 1833). S. Afr. Quart. J. 2: 59. Uitenhage, Cape Province, South Africa.

Taxonomy Originally Vespertilio hottentota. Synonyms: augusticeps, bensoni, megalurus, pallidior, portavernus, smithi. Subspecies: three or four (see Geographic Variation). Chromosome number (Zimbabwe, South Africa): 2n = 50; aFN = 48; X is a medium-sized submetacentric and Y a small acrocentric (Peterson & Nagorsen 1975, Rautenbach et al. 1993, Kearney et al. 2002).

[42–49]% of GLS, n = 43). Rostrum broad. Profile of forehead region (viewed laterally) straight. Occipital helmet present; sagittal crest slight; lambdoid crest well developed. Inner upper incisor large, unicuspid. Outer upper incisor minute. Anterior upper premolar absent; canine and posterior premolar in contact (Figure 122f). Lower molars myotodont.

Description Small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; four upper and five lower cheekteeth and two upper incisors on each side; ears separated and short for a vespertilionid (14–20 mm); one of the three large African Eptesicus (FA: 45–54 mm); ventral hairs bicoloured; inner upper incisor unicuspid (cf. bicuspid in other African Eptesicus). Males on average with slightly shorter FA than !!, at least in some parts of geographic range (Namibia). Pelage sleek, soft and dense; mid-dorsal hairs 7–9 mm. Dorsal pelage varying geographically from pale brown to almost black, often with silky sheen; hairs dark at base with paler tips. Ventral pelage paler, often with creamy sheen; hairs with basal two-thirds blackishbrown, terminal third pale greyish-brown with cream or whitish tips. Ears dark brown to blackish, relatively short, tip rounded. Tragus length ca. one-third of E; broadest at base; posterior margin mostly straight but smoothly convex near tip; tip rounded (Figure 122c). Wings and interfemoral membrane brown (when pelage is comparatively pale) to blackish-brown; no white hind-border. Tail 73 (58–84)% of HB, therefore (on average) not relatively longer than in other African Eptesicus. Tail with terminal vertebra slightly protruding beyond interfemoral membrane. Postcalcareal lobe well developed. Baculum short, roughly triangular but with concave sides, flat but with proximal end thicker than distal end (Figure 123b) (Hill & Harrison 1987). Skull (Figure 122d and e) large for an African Eptesicus. Braincase relatively low (38.8 [37–42]% of GLS, n = 19) and narrow (45.7

Geographic Variation Based on measurements and/or pelage colouration, the following taxa have been given subspecific status by some authors, but more information and material are needed to confirm their status: E. h. portavernus: SW Kenya. According to Schlitter & Aggundey (1986), this subspecies is larger and heavier than the other subspecies, and has a shorter palate and deeper braincase. Dorsal pelage pale brown; ventral pelage pale tan. However, only two specimens are known. E. h. bensoni: E Zambia, Malawi and NW Mozambique and across Zimbabwe to E South Africa (Rautenbach et al. 1984, Schlitter & Aggundey 1986, Koopman 1994). Dorsal pelage dark brown to almost black; ventral pelage buff. E. h. pallidior: Namibia to NW Cape Province (Meester et al. 1986). Dorsal pelage pale brown to fawn; ventral pelage whitish-buff with irregular darker patches. Considered a synonym of hottentotus by Schlitter & Aggundey (1986). E. h. hottentotus: S and SW Cape Province (Meester et al. 1986), not extending far inland. Dorsal pelage deep reddish-brown (Smith 1833) or brown, sometimes with yellowish tinge (specimens from Algeria State Forest; T. Kearney pers. comm.). Similar Species Eptesicus serotinus. Inner upper incisor bicuspid. North Africa. E. bottae. Inner upper incisor bicuspid. Egypt. E. platyops. Inner upper incisor bicuspid. Senegal, Nigeria and Bioko I. 555

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Family VESPERTILIONIDAE

in clutter, faster while foraging in semi-open areas (Norberg & Rayner 1987). Day-roosts include dry caves, one very wet cave, abandoned mines, a small hollow in rocks at the entrance of an old mine inhabited by Miniopterus natalensis, and the outside wall of a building (Smithers 1983, Herselman & Norton 1985). Individuals hang from the walls in very loose formations. Has been mist-netted over water. Foraging and Food No information. Based on wing morphology, predicted to forage by slow-hawking in both moderately cluttered areas and open spaces. One was mist-netted flying low over a marsh (Herselman & Norton 1985). Six were captured in mist-nets set partly over water and partly over land at the edge of the water; all were caught over the land (Seamark 2005). Diet not known. Echolocation Calls of one individual (flying in a room) were broad bandwidth calls (20–54 kHz) of short duration (2–4 ms), with peak-frequency 28–32 kHz and usually a strong harmonic component (Taylor 2000). See also Monadjem et al. (2010). Eptesicus hottentotus

E. floweri. Much smaller (FA: 34–38 mm; GLS: 12.0, 13.5 mm). Horny warts on forearms, legs and tail. Sudan. Pipistrellus (28 African spp.). All much smaller (FA: 50 m above ground (Iberia): steep/shallow FM; fundamental harmonic only (Figure 132a). Start-frequency 25.1 kHz; end-frequency 17.6 kHz; bandwidth 7.5 kHz; peak-frequency 18.8 kHz; call-duration 12.3 ms; inter-call interval 319.7 ms (Ibáñez et al. 2001). Uses consecutive calls of alternate frequency as does Nyctalus noctula (not found in Africa) although more rarely and only during flights at low altitude ( 50 m): successive calls have same frequency; only the fundamental harmonic is present; bandwidth comparatively narrow; inter-call interval comparatively long. (b) In flight at low altitude (< 20 m) a few minutes after leaving roost: successive calls differ in frequency; second harmonic present; bandwidth wider; inter-call interval shorter.

Social and Reproductive Behaviour Very little information. Establishes maternity colonies of up to 80 !! (no ""). Vocalizations in the maternity colonies are frequent even during the day. These social calls have high intensities; each sequence comprises 25 calls with tonal structure, which are slightly modulated with endfrequencies between 9 and 15 kHz; therefore they are audible to humans and have a characteristic metallic tone (Ibáñez et al. 2004). As yet, there is no information about the roosting behaviour of "". Reproduction and Population Structure Litter-size (Europe): one or two (frequencies not known). Extralimitally, in southern Iberia, the reproductive chronology is restricted seasonal monoestry with "" showing well-developed scrotal testes from early Aug to Oct (mid-summer to mid-autumn), and !! giving birth at end of May or early Jun (late spring) and lactating until early Aug (Ibáñez et al. 2004). A neonate born in captivity weighed 9.9 g and had forearm 26.5 mm (Topál 1976).Young are volant by ca. 40 days (Alekseeva 1988). Predators, Parasites and Diseases Several cases of predation by the Barn Owl Tyto alba are known in Europe. A predation by Beech Marten Martes foina is reported from Crimea (Ibáñez et al. 2004). Conservation IUCN Category: Near Threatened (assessed mainly from extralimital data). Ongoing reduction of Mediterranean forests in North Africa as result of human activities and desertification is likely to severely reduce population numbers. Almost qualifies for Vulnerable.

591

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Measurements Nyctalus lasiopterus FA: 61.4 (59–64) mm, n = 6 WS: n. d. TL: 145.2 (138–152) mm, n = 6 T: 55.2 (48–62) mm, n = 6 E: 21.0 (19–23) mm, n = 6 Tr: n. d. Tib: n. d. HF: 14.2 (13–16) mm, n = 6

WT: 48.8 (37.3–58.5) g, n = 60* GLS: 22.4 (22.1–22.7) mm, n = 6 GWS: 15.5 (15.1–16.0) mm, n = 6 C–M3: 8.8 (8.6–9.0) mm, n = 6 Libya (Qumsiyeh & Schlitter 1982) *Southern Iberia (C. Ibáñez unpubl.) Key Reference

Ibáñez et al. 2004. C. Ibáñez

Nyctalus leisleri LEISLER’S NOCTULE (LEISLER’S BAT) Fr. Noctule de Leisler; Ger. Kleinabendsegler Nyctalus leisleri (Kuhl, 1817). Die Deutschen Fledermäuse. Hanau, pp. 14, 46. Hanau, Hessen, Germany.

Taxonomy Originally Vespertilio leisleri. Synonyms: four; none in Africa. Subspecies: two; only the nominate occurs in Africa. Chromosome number (Poland, Czechoslovakia): 2n = 46; aFN = 50 (Fedyk & Fedyk 1970, Zima 1978). Description Small to very small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; five upper and five lower cheekteeth and two upper incisors on each side; ears separated, short for a vespertilionid (13–16 mm) and also relatively short; fifth metacarpal notably shorter than third and fourth (cf. Pipistrellus); FA: 41–44 mm. Sexes similar. Pelage soft, sleek, dense; mid-dorsal hairs 7–9 mm; ventral pelage extending on to wings and interfemoral membrane. Dorsal pelage golden-brown; hairs dark brown at base becoming golden-brown towards tip.Ventral pelage slightly paler (yellowish); hairs blackish-brown with brown tip. Muzzle dark brown. Ears dark brown, widely separated, relatively short, subtriangular but with very rounded tip; outer margin with 4–5 folds. Tragus kidney-shaped, short, blunt. Wings dark brown, long and narrow; fifth metacarpal notably shorter than third and fourth metacarpals (ca. 80–85% of third metacarpal, n = 3). Wingmembranes attaching to ankle. Interfemoral membrane dark brown. Calcar reaching half-way to tip of tail. Postcalcareal lobe wide with visible T-shaped piece of cartilage (Figure 131). Skull small (cf. N. lasiopterus); rostrum broad; viewed laterally, the dorsal profile is a smooth convex curve from front to back. No sagittal crest; lambdoid crest weakly developed; no occipital helmet. Dentition moderately weak. Incisors relatively small. Anterior upper premolar visible from the outside. Height of mandibular ramus: 2.9–3.6 mm. Geographic Variation None recorded in Africa. Similar Species Only two other vespertilionids occurring (or perhaps occurring) in North Africa have five upper and five lower cheekteeth and two upper incisors on each side; ears not joined at base; FA: >40 mm: Nyctalus lasiopterus. Much larger (FA: 59–69 mm throughout geographic range). Height of mandibular ramus >5.0 mm. N. noctula. Larger (FA: 45–57 mm). Height of mandibular ramus 4.3– 4.8 mm. Pelage unicoloured. Occurrence in Africa uncertain.

Nyctalus leisleri

Distribution In Africa, recorded in the Mediterranean Coastal and Afromontane–Afroalpine BZs from the mountains of Morocco (Rif, High Atlas), Algeria (Tell Atlas) and Libya (Cyrenaica) (Hanák & Gaisler 1983, Ibáñez 1988, Kowalski & Rzebik-Kowalska 1991, Benda et al. 2004d). Presumably also occurs in N Tunisia although not yet recorded there. Extralimitally: this Palaearctic species ranges from western Europe (and Madeira I.) through Urals and Caucasus to the W Himalayas. Habitat All of the North African specimens were mist-netted in, or near, forests at 400–900 m; mainly oak forests (Quercus suber, Q. ilex, Q. faginea, Q. canariensis), and also juniper-cypress forest (Juniperus excelsa, Cupressus sempervirens), where they often foraged over streams (Hanák & Gaisler 1983, Gaisler & Kowalski 1986, Ibáñez 1988, Benda et al. 2004d).

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Abundance Uncertain. As for nearly all the tree-dwelling species that require specific surveys, most of the records in North Africa are recent. Particularly abundant in mist-net captures in Djurdjura (NE Algeria) (Gaisler & Kowalski 1986). Population size limited by the reduced distribution of suitable habitats. Adaptations Aspect ratio medium; wing-loading very high (Norberg & Rayner 1987). Flight fast; manoeuvrability predicted to be poor. In summer, roosts by day mainly in tree-holes, under bark (including for breeding colonies) and occasionally in buildings. In winter, hibernates in hollow trees and sometimes in rock-crevices (Schober & Grimmberger 1987). Hibernation period in Africa shorter than in Europe – in Algeria, "" were active in mid-Oct (Gaisler & Kowalski 1986). Migrates: longest recorded movement 1567 km, from Germany to Spain (Ohlendorf et al. 2000). Foraging and Food Forages by fast-hawking over open areas and above canopies; foraging flights fast, straight, often with dives. Foraging mainly occurs during two periods, the first after dusk and the second ending ca. 30 min before dawn. Occasionally flies during the day. Foraging areas average 7.4 km2, but are up to 18.4 km2 in southern Britain (Waters et al. 1999). Diet: no information for Africa. In Ireland and England, Diptera (Nematocera and Cyclorrhapha) are the predominant prey, but Coleoptera, Trichoptera and Lepidoptera are also important (Sullivan et al. 1993,Waters et al. 1995, 1999,Vaughan 1997). In Germany and Switzerland, Lepidoptera, Nematocera and Neuroptera may, at least locally, be important items of food (Beck 1995, Shiel et al. 1998). Echolocation Search-phase calls (above level of treetops) are of two types. (a) Call-shape QCF; start-frequency 26–28 kHz; endfrequency 22–24 kHz; peak-frequency 22–24 kHz; call-duration 15– 25 ms. (b) Call-shape shallow FM; start-frequency 26–30 kHz; endfrequency 25–26 kHz; peak-frequency 25–26 kHz; call-duration 7–15 ms; intercall-interval 220–550 ms (Waters et al. 1995,Tupinier 1996). Social and Reproductive Behaviour Mating takes place in late summer. Males are very territorial at this time; each " is assumed to attract !! by means of an ‘advertisement song’ (von Helversen & von Helversen 1994) and occupies a mating-roost with a harem of up to nine !!. During winter, "" and !! hibernate together in large groups. In summer, !! form maternity colonies of 10–50 individuals (exceptionally several hundred in attics in Ireland; Shiel &

Fairley 1999); adult "" and immature bats roost elsewhere. In Poland, maternity colonies in trees comprised 20.7 (7–38) individuals; roost-fidelity was low (Ruczynski & Ruczynska 1999). Maternity roosts are vacated by late Aug to early Sep. Reproduction and Population Structure Litter-size (Western Europe): usually two. In Europe, the reproductive chronology is restricted seasonal monoestry with mating in late summer (late Aug– Sep); births in mid-summer (Jun–Jul); young are volant one month later (Pottier 1993, Shiel & Fairley 1999). In Morocco, births occur in May and by mid-Jul juveniles are almost the same size as adults (Ibáñez 1988). Maximum life-span: nine years (Schober & Grimmberger 1987). Predators, Parasites and Diseases

No information.

Conservation IUCN Category: Least Concern (assessed mainly from extralimital data). Loss of habitat because of logging and overgrazing is the main threat in North Africa. Measurements Nyctalus leisleri FA: 42.4 (41–44) mm, n = 14 WS (d): 279.2 (267–286) mm, n = 5 TL: 101, 104 mm, n = 2 T: 40.9 (38–44) mm, n = 7 E: 14.1 (13–16) mm, n = 7 Tr: 6.6 (5.5–7.5) mm, n = 10 Tib: 15.3 (15–16) mm, n = 5 HF: 9.2 (9–10) mm, n = 6 WT: 12.1 (11–15) g, n = 11 GLS: 15.2 (14.8–15.7) mm, n = 27 GWS: 10.3 (10.0–10.8) mm, n = 24 C–M3: 5.8 (5.6–6.1) mm, n = 24 Morocco, Algeria, Libya (Hanák & Gaisler 1983, Ibáñez 1988, Romero Zarco 1990, Kowalski & Rzebik-Kowalska 1991, Benda et al. 2004d) Key References Benda et al. 2004d; Gaisler & Kowalski 1986; Hanák & Gaisler 1983; Ibáñez 1988; Kowalski & Rzebik-Kowalska 1991. Stéphane Aulagnier

593

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Family VESPERTILIONIDAE

GENUS Nycticeinops Schlieffen’s Twilight Bat Nycticeinops Hill & Harrison, 1987. Bull. Br. Mus. Nat. Hist. 52: 254. Type species: Nycticeius schlieffeni Peters, 1859.

Nycticeinops schlieffeni.

A monotypic genus endemic to Africa south of the Sahara. Diagnostic combination of characters: four upper and five lower cheekteeth and one upper incisor on each side (as in Otonycteris, Scotoecus and Scotophilus); ears relatively short (cf. Otonycteris); penis short (cf. Scotoecus); FA: 28–35 mm (cf. 41–88 mm in Scotophilus); dental

formula: 1113/3123 = 30. Selected characters of Nycticeinops are illustrated in Figure 133. Based on bacular morphology, Hill & Harrison (1987) removed schlieffeni (the only African member of the genus Nycticeus) into a new genus, Nycticeinops. Nycticeinops is considered a subgenus of Nycticeus by Koopman (1994), but chromosomal evidence from one specimen from Somalia provides additional support for the generic status of Nycticeinops (Ruedas et al. 1990): the chromosome numbers for the Somali specimen are 2n = 34, aFN = 52 compared with 2n = 46, aFN = 48 for Nycticeus humeralis, a species from North America. This difference would represent one of the largest intra-generic differences in diploid number within the Vespertilionidae, which strongly indicates that Nycticeinops and Nycticeus are not congeneric. Mitochondrial ribosomal sequences provide further support for the generic distinction between Nycticeus and Nycticeinops (Hoofer & Van Den Bussche 2001). However, Rautenbach et al. (1993) found the karyotypes of 22 specimens of N. schlieffeni from southern Africa to be 2n = 42, aFN = 50 (i.e. different from that of the specimen from Somalia). The difference between the karyotype of these southern African specimens and that of Nycticeus humeralis is not so marked and therefore Rautenbach et al. (1993) retained schlieffeni in Nycticeus. It is possible that the material from Somalia and that from South Africa represents two distinct species (G. N. Bronner pers. comm.). Alternatively, the identity of the Somali specimen

Figure 133. Nycticeinops schlieffeni. Flight membranes, and bones of wing, hindlimb and tail. Left ear and tragus. Details of tragus. Skull (MNHN CG 1997-2104).

594

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Nycticeinops schlieffeni

might be erroneous. Nycticeinops is placed in the tribe Vespertilionini by Hoofer & Van Den Bussche (2001) and in tribe Nycticeiini (with Scotoecus and Scotophilus) by Simmons (2005). The only described species is Nycticeinops schlieffeni. Meredith Happold

Nycticeinops schlieffeni SCHLIEFFEN’S TWILIGHT BAT (SCHLIEFFEN’S BAT) Fr. Nycticinope de Schlieffen; Ger. Schlieffens Abendfledermaus Nycticeinops schlieffeni (Peters, 1859). Monatsber. K. Preuss. Akad. Wiss. Berlin 1859: 223. Cairo, Egypt.

Taxonomy Originally Nycticejus schlieffeni. Synonyms: adovanus, africanus, albiventer, australis, bedouin, cinnamomeus, fitzsimonsi, minimus. Subspecies: uncertain (see Geographic Variation). Chromosome number (South Africa): 2n = 42; aFN = 50 (Rautenbach et al. 1993). This differs markedly from the 2n = 34; aFN = 52 karyotype reported for a specimen from Somalia (Ruedas et al. 1990) (see Genus Nycticeinops). Description Very small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; four upper and five lower cheekteeth and only one upper incisor on each side; ears well separated, short for a vespertilionid (9–13 mm); FA: 28–35 mm; wings dark brown; penis 5.6–6.2 mm; anterior lower premolar ca. half height and half crown area of posterior premolar. Sexes similar in colour and size. Pelage dense, soft, slightly fluffy, with no sheen; mid-dorsal hairs 4–6 mm. There is considerable variation in colour. Dorsal pelage cinnamon, fawn or greyish-fawn; hairs unicoloured. Ventral pelage slightly to considerably paler, whitish or pure white. Head slightly flattened; muzzle short with prominent lateral glandular swellings. Ears dark brown; comparatively and relatively short for a vespertilionid (36.1 [31–42]% of FA); inner margin slightly convex, outer margin slightly concave; tip rounded. Tragus length 43.7 (40– 50)% of E; shape as in Figure 133. Eyes very small. Wings and interfemoral membrane dark brown with blackish-brown venation. Free edge of interfemoral membrane with faint white border in some individuals. Penis comparatively short (5.6–6.2 mm, n = 9) (cf. Scotoecus albofuscus, S. hirundo). Baculum with expanded bifid base and long fluted shaft with pointed tip (Hill & Harrison 1987). Skull (Figure 133) somewhat flattened and fairly broad; dorsal profile (viewed laterally) rising at gentle slope from incisors to lambdoid crest; frontal area slightly concave. Zygomatic arches slender and weak but seldom lost during preparation of skulls (cf. Scotoecus). Sagittal crest absent or very low and only over posterior of braincase; lambdoid crest weakly developed; no occipital helmet. Upper incisor fairly long, unicuspid. Canines (upper and lower) robust; upper canine with anterior surface rounded and not grooved. Upper premolar sharply pointed, reaching to two-thirds to three-quarters height of canine. Posterior upper molar with three ridges (cf. Scotophilus). Lower incisors tricuspid. Anterior lower premolar ca. half height and half crown area of posterior lower premolar (cf. Scotoecus). Geographic Variation Four subspecies (albiventer, australis, fitzsimonsi and schlieffeni) were recognized by Hayman & Hill (1971), but they are based largely on colour and are of dubious validity

(Rosevear 1965, Koopman 1994) and are not recognized by Simmons (2005). Specimens from Mozambique and KwaZulu–Natal, South Africa (the form australis) are darker than those from elsewhere. Similar Species Only one other dark-winged vespertilionid in Africa has only one upper incisor on each side and FA 28 mm): Otonycteris hemprichii. Ears with inner margins not joined at base. Only four upper and five lower cheekteeth and one upper incisor on each side. Distribution  In Africa, only known from the Sahara Arid BZ. Confirmed records are from the oases of Siwa and Al Jaghbub near the Libyan–Egyptian border, from the Nile Valley (from the Delta in Egypt to the Fifth Cataract in N Sudan), and from oases in Sinai

Plecotus christii

663

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Family Vespertilionidae

(Kock 1969a, Gaisler et al. 1972, Qumsiyeh 1985, P. Benda et al. unpubl.). Old records from Eritrea and Ethiopia are now assigned to P. balensis. Extralimitally: SW Jordan and S Palestine. Harrison & Bates (1991) provisionally identified specimens from the Arabian Peninsula as P. austriacus christii, but this bat lives only in the deserts along the Rift Valley (Jordan, Palestine); in N Arabia, only Plecotus macrobullaris has been confirmed (Benda et al. 2004c), and the population provisionally assigned to P. christii from Yemen and SW Saudi Arabia needs revision. Habitat  All records are from open areas of oases in very dry deserts, and from mesic habitats along the Nile R., where suitable day-roosts are available. Abundance  Probably rare. However, in Sinai, it was one of the bats that was recorded comparatively frequently. Remarks  Wings broad; this species is well adapted for hovering and forages by gleaning, often picking insects off leaves and blossoms. In Israel, feeds predominantly on medium-sized and large Lepidoptera (Noctuidae and Hepialidae) and occasionally on Coleoptera (Scarabaeidae), Diptera and Trichoptera (Whitaker et al. 1994, Feldman et al. 2000). By day, roosts mainly in rocky cavities, but also roosts in dark areas of pyramids, old monuments, ruins, caverns and abandoned mines where individuals hang from the walls. Sedentary; the homerange is restricted to the oasis. Usually found to be solitary. Males become territorial in autumn, when mating occurs. Litter-size: one (n = 1). Reproductive chronology not known. Extralimitally, in the Negev Desert (Israel), one / in early pregnancy was reported at the beginning of Mar and, in the Dead Sea area, 50% of 48 // were

lactating in mid-Apr, and one / was still lactating in a cave nearby as late as mid-Jun (Yom-Tov et al. 1992). Conservation  IUCN Category: Data Deficient. Population probably small; suitable habitat might be fragmented, but needs more research to be properly evaluated. Measurements Plecotus christii FA: 38.4 (36–41) mm, n = 17 WS: n. d. TL: 93.3 (92–96) mm, n = 6 HB: 48.5 (42–54) mm, n = 9 T: 45.0 (42–48) mm, n = 6 E: 36.0 (32–39) mm, n = 10 Tr: 17.8 (17.4–18.5) mm, n = 4 Tib: n. d. HF: 8.3 (8–9) mm, n = 7 WT: 5.8, 7.8 g, n = 2 GLS: 16.6 (16.0–17.1) mm, n = 15 GWS: 8.6 (8.1–8.9) mm, n = 14 C–M3: 5.4 (5.2–5.7) mm, n = 15 Egypt and Libya (Gaisler et al. 1972, Qumsiyeh 1985, Benda et al. 2004c) Key References  Benda et al. 2004c; Gaisler et al. 1972; Kock 1969a; Qumsiyeh 1985. Petr Benda & Stéphane Aulagnier

Plecotus gaisleri  Gaisler’s Long-eared Bat Fr. Oreillard du Maghreb; Ger. Libysches Langohr Plecotus gaisleri Benda, Kiefer, Hanak and Veith, 2004. Folia Zool. 53, Monograph 1: 28. Wadi al Kuf, SW Massah, Cyrenaica, Libya.

Taxonomy  Populations of Plecotus from the Maghreb in NW Africa and Cyrenaica in NE Libya were identified as P. auritus and then considered to represent a subspecies of P. austriacus. Subsequently, based on morphological evidence, Benda et al. (2004c) revised African populations of Plecotus and concluded that the Maghrebian and Cyrenaican populations represented subspecies of P. teneriffae. They described a new subspecies, P. t. gaisleri, based on specimens from Cyrenaica and, pending genetic evidence, tentatively included the Maghrebian population in P. t. gaisleri although some differences in colour were observed. In their Appendix 2, however, they referred to the Maghrebian population as P.t.cf. gaisleri. Genetic evidence confirmed that the Maghrebian and Cyenaican populations are distinct (Juste et al. 2004). Subsequently, based on both morphological and genetic evidence, Spitzenberger et al. (2006) revised the genus Plecotus, and concluded that P. kolombatovici contained four subspecies of which two occurred in Africa: P. k. gaisleri in Cyrenaica and an as yet undescribed subspecies in the Maghreb. More recently, based only on material from Cyrenaica, Mayer et al. (2007) raised gaisleri to specific status, but did not discuss the relationship of P. gaisleri to the Maghrebian population. Therefore, for convenience and pending resolution of this relationship,

in this profile, both populations are treated as P.gaisleri, with presumably two subspecies. Synonyms: aegyptius, christiei. Chromosome number (Tunisia): 2n = 32; aFN = 50 (Baker et al. 1974). Description  A very small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; five upper and six lower cheekteeth and two upper incisors on each side; ears joined at base, large for a vespertilionid (E: 33–40 mm) and relatively enormous (ca. 92% of FA); dorsal pelage greyish-brown, hairs with basal half dark; face, ears and wings dark; muzzle not inflated behind nostrils. Sexes similar. Pelage dense, soft, slightly woolly; mid-dorsal hairs 7–9 mm. Dorsal pelage greyish-brown; hairs tricoloured with basal half dark rusty-brown, terminal half pale or dark umber depending on humidity of habitat (animals from dry regions are paler) with brown or greyish-brown tip.Ventral pelage pale yellowish-brown or pale greyish-brown; hairs dark greyish-brown with pale yellowishbrown or pale greyish-brown at tip. Face dark brown to dark greyishbrown. Muzzle narrow, not inflated behind nostrils, protuberance above upper eyelid (supraorbital gland) comparatively large, ca. 1.2 mm in diameter (cf. P. christii). Ears dark brown to dark greyish-

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brown, relatively long (94 [86–99]% of FA and therefore look enormous), inner margins joined by low band at base. Tragus opaque, of medium relative length (46 [42–50]% of E), 6 mm at widest, tapering to blunt point. Eyes relatively large for a microbat. Wing-membranes dark brown to dark greyish-brown; skin over wing-bones moderately darker. Thumb comparatively long (6.1 [5.2–6.8] mm) (cf. P. christii). Baculum Y-shaped; lateral arms long, broad, with acute angle between them (Figure 143c). Skull medium-large for a Plecotus, with braincase relatively high, short and broad (braincase width 47–51% of GLS) (Benda et al. 2004c). Dorsal profile of forehead region strongly concave for an African Plecotus (Figure 142c) (cf. P.balensis, P.christii). Rostrum relatively long (C–M3: 33–35% of GLS), medium in height and relatively broad (width between canines 68–71% of C–M3). Dentition more robust than in P. balensis and P. christii. Anterior upper premolar comparatively large (length 0.44–0.54 mm; breadth 0.52–0.62 mm); within toothrow; canine and posterior premolar well separated. Geographic Variation  There are no profound morphological differences between P. g. gaisleri from Cyrenaica and P. g. cf. gaisleri from the Maghreb (Benda et al. 2004c).These authors observed colour variation in the Maghrebian populations: specimens from more arid desert and semi-desert habitats are paler than those from coastal and montane habitats and, similarly, they are paler than P. g. gaisleri from Cyrenaica. Spitzenberger et al. (2006) also reported differences in colour but also observed differences in cranial characters. Mean CbL in ?? and // respectively: 15.8 and 16.0 mm in P. gaisleri; 15.9 and 16.2 mm in P. g. cf. gaisleri. Similar Species  The only other vespertilionids in Africa that have ears with inner margins joined are: Plecotus balensis. Dorsal hairs with basal two-thirds dark brown. Skull with frontal region weakly concave. Rostrum usually relatively shorter and narrower (details in Description). Baculum Y-shaped with obtuse angle between lateral arms. Ethiopia, possibly Eritrea. P. christii. Ears unpigmented except around tip. Wing-membranes pale, semi-translucent. Muzzle inflated behind nostrils. Skull with frontal region weakly concave; rostrum relatively narrower and usually relatively shorter (details in Description). BaculumY-shaped with comparatively short lateral arms. NE Sahara (Libya, Egypt, N Sudan). Barbastella (2 spp.). Ears much shorter (13–18 mm). Only five lower cheekteeth and one upper incisor on each side. Only one other vespertilionid in Africa has long ears (>28 mm): Otonycteris hemprichii. Ears with inner margins not joined at base. Only four upper and five lower cheekteeth and one upper incisor on each side. Distribution  In Africa, recorded from the Mediterranean Coastal and Afromontane–Afroalpine BZs in Morocco, Algeria, Tunisia and N Cyrenaica in Libya, and marginally from the Saharan Arid BZ in Morocco, Algeria, Tunisia and Tripolitania (NW Libya) (Aulagnier & Thévenot 1986, Kowalski & Rzebik-Kowalska 1991,

Plecotus gaisleri

Benda et al. 2004c, d). Extends from the coast to the NW border of the Sahara (Anti-Atlas, Saharan Atlas, Jebel Nafusa). Old records from Senegal require confirmation and have not been mapped (most probably they have been mislocated). Habitat  Inhabits a variety of open and semi-covered areas – mainly steppes but also agricultural landscapes, lowland basins and mountains. Recorded from sea level to 2630 m in the High Atlas. In the Maghreb, mostly recorded from Morocco and the geographic range covers all the mountain ridges of this country. In Algeria, recorded from the coast to the southern slopes of the Saharan Atlas Mts. In Tripolitania, recorded from three arid places on the Jebel Nafusa. In Cyrenaica, recorded from at least nine localities in an area of ca. 10,000 km2 of coastal and montane vegetation. Abundance  Comparatively numerous, although only found in small colonies. Common in both small settlements and large towns. Adaptations  Wing morphology and flight characteristics similar to other Plecotus. In summer, roosts by day mainly in rocky cavities and caves, but also in dark areas of minarets, old monuments, ruins, caverns and old houses. Most frequently clings to sides of deep cracks, but also hangs from ceilings. In winter, roosts by day in various spaces in buildings, caves, mines, wells and holes in trees. Found in mixed colonies with Rhinolophus blasii. Active all yearround in Algeria, although daily torpor has often been recorded in summer, and undoubtedly occurs in all months. Foraging and Food  Forages at night (and even before sunset in autumn in Algeria; Kowalski & Rzebik-Kowalska 1991), by slowhawking in parks, over meadows, among rocks with sparse subdesert vegetation, and over water. Feeds predominantly on mediumsized and large Lepidoptera (mostly Noctuidae), and occasionally on Coleoptera and Diptera (Trujillo 2002). 665

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Echolocation  No information.

Conservation  IUCN Category: Not Evaluated. Pesticides threaten populations in agricultural areas.

Social and Reproductive Behaviour  In winter, roosts singly or in tight clusters of ca. ten individuals of both sexes. In summer, the sexes segregate; adult // roost in maternity colonies of 10–30 adults; they hang separately or in small groups. Mating occurs in autumn; ?? become territorial at this time. Reproduction and Population Structure  Litter-size: one (n = 2). Reproductive chronology not known, probably restricted seasonal monoestry as in P. teneriffae (Trujillo 2002). Births occur from end of Apr to mid-May in Cyrenaica, and from May to early Jun in Tunisia (Baker et al. 1974). In the Canary Is., 70% of // give birth in the main known colony, and most of the young in this colony can fly by mid-Jul (Trujillo 2002). The size of testes is maximal in Sep, which is probably the main mating period. Predators, Parasites and Diseases  Predators occasionally include Barn Owls Tyto alba and Pharaoh Eagle-owls Bubo ascalaphus (Aulagnier 1989). Ectoparasites include a bat-fly Nycteribia schmidli (Diptera: Nycteribiidae), a flea Nycteridopsylla pentactena (Siphonaptera: Ischnopsyllidae) and a tick Carios boueti (Acari: Argasidae) (Anciaux de Faveaux 1984, Beaucournu & Kowalski 1985).

Measurements Plecotus gaisleri FA: 39.5 (37–42) mm, n = 44 WS: n. d. TL: n. d. HB: 49.8 (45–55) mm, n = 24 T: 47.6 (44–52) mm, n = 24 E: 36.1 (33–40) mm, n = 28 Tr: 16.7 (14.9–18.5) mm, n = 25 Tib: n. d. HF: 8.6 (8–9) mm, n = 14 WT: 8.1 (7.0–10.0) g, n = 24 GLS: 17.0 (16.2–17.4) mm, n = 43 GWS: 9.0 (8.6–9.2) mm, n = 39 C–M3: 5.8 (5.6–5.9) mm, n = 42 Morocco, Algeria, Tunisia, Libya (mainly from Benda et al. 2004c) Key References  Aulagnier & Thévenot 1986; Benda et al. 2004c; Gaisler 1984; Hanák & Elgadi 1984; Kowalski & Rzebik-Kowalska, 1991. Petr Benda & Stéphane Aulagnier

GENUS Scotoecus Lesser House Bats Scotoecus Thomas, 1901. Ann. Mag. Nat. Hist., ser. 7, 7: 263. Type species: Scotophilus albofuscus Thomas, 1890.

A polytypic genus with, controversially, two, three or four species endemic to sub-Saharan Africa and one endemic to Pakistan and N India (Hill 1974b, Koopman 1994, Simmons 2005). Diagnostic combination of characters (African species only): four or five upper and five lower cheekteeth and one upper incisor on each side (as in Nycticeinops, Otonycteris and Scotophilus); ears relatively short; FA: 28–40 mm; penis very long (9–16 mm); upper canine with anterior surface flattened and grooved; posterior upper molar with three ridges; anterior lower premolar at least two-thirds height of posterior premolar, usually subequal. Skull somewhat flattened; braincase broad; rostrum very broad across lachrymals; dorsal profile (viewed laterally) rising at gentle slope from incisors to lambdoid crest; frontal area slightly convex. Interorbital region broad. Zygomatic arches very slender and weak; usually lost during preparation of skulls. Sagittal and lambdoid crests very weakly developed; helmet slight. Rostral and anterior palatal emarginations very deep. Selected characters of Scotoecus are illustrated in Figure 144. Treated as subgenus of Nycticeus (e.g. Ellerman et al. 1953, Hayman & Hill 1971), reinstated to generic status by Rosevear (1965) and

retained as a genus by Hill (1974b), Koopman (1993, 1994) and Simmons (2005). In Africa, there are light-winged and dark-winged forms. All light-winged Scotoecus are placed in S. albofuscus (with S. a. woodi as a subspecies) and there are no taxonomical problems associated with this species. In contrast, five dark-winged forms have been named but their status is controversial. Simmons (2005) recognizes three dark-winged species. In contrast, pending resolution of the problems, all dark-winged forms are tentatively considered here to represent one species, S. hirundo. The controversies are discussed in the profile of this species. Both African species are found in woodland savannas; S. albofuscus also occurs in some forest habitats. The African species recognized here are: S. albofuscus.Wings white, semi-translucent, becoming brownish around edges and near body. S. hirundo. Wings dark-brown, slightly translucent. Meredith Happold

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Scotoecus albofuscus

e

a

f

b

c

d

g

Figure 144. African Scotoecus. (a) Flight membranes, and bones of wing, hindlimb and tail of Scotoecus albofuscus from Malawi. (b) Left ear and tragus of S. hirundo from Malawi. (c) Details of left tragus of the same individual. (d) Left tragus of Scotoecus albofuscus from Côte d’Ivoire. (e), (f) and (g) Lateral, dorsal and ventral views of skull of Scotoecus albofuscus woodi (holotype, BMNH 17.2.1.1, from Malawi).

Scotoecus albofuscus  Light-winged Lesser House Bat (Gambian Lesser House Bat) Fr. Scotèce à ailes blanche; Ger. Weißflügel-Hausfledermaus Scotoecus albofuscus (Thomas, 1890). Ann. Mus. Civ. Stor. Nat. Genova 29: 84. Bathurst, Gambia.

Taxonomy  Originally Scotophilus albofuscus. Synonyms: woodi. Sub­species: two of dubious validity (see Geographic Variation). Chromosome number: not known. Description  Very small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; four upper and five lower cheekteeth and only one upper incisor on each side; ears well separated, short for a vespertilionid (9–15 mm); FA: 29– 34 mm; wings white and semi-translucent; penis ca. 9 mm. Sexes apparently similar in colour and size, but data are limited. Pelage soft and dense; mid-dorsal hairs 4–6 mm. Dorsal pelage dark brown, golden-brown or fawn; hairs unicoloured.Ventral pelage slightly paler than dorsal pelage. Head somewhat flattened, muzzle broad, flat with

nostrils opening sideways from small transverse pad. Ears widely separated, comparatively and relatively short for a vespertilionid (40.8 [32–52]% of FA), oval; outer margin with semi-circular antitragus; tip rounded. Tragus short (just over one-third of E), rounded (Figure 144d). Eyes small. Wings white, semi-translucent, becoming brownish with brown venation around edges and close to body; forearms and first, second and third fingers brown, fourth and fifth fingers white. Interfemoral membrane blackish-brown. Calcars long but weak. Penis long (ca. 9 mm). Skull (Figure 144) as described in Genus Scotoecus. Upper incisor fairly long, unicuspid, directed inwards. Upper canine with anterior face flat and conspicuously grooved. Only one upper premolar – anterior premolar apparently always absent (Hill 1974b, Taylor & 667

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van der Merwe 1998). Posterior premolar pointed; in contact with canine; reaching two-thirds to three-quarters height of canine. Posterior upper molar with three ridges (cf. Scotophilus). Anterior lower premolar only slightly shorter, and slightly stouter, than the tall, slender posterior lower premolar (cf. Nycticeinops). Dental formula: 1113/3123 = 30. Geographic Variation  Two subspecies were recognized, mainly on basis of body and skull measurements, by Hill (1974b) and retained by Koopman (1994) and Simmons (2005). However, analysis of more data than Hill (1974b) had access to suggests that the morphometric differences are slight, but the geographic distributions of the two taxa appear to be very widely separated by the Rainforest BZ in the Congo Basin, and this supports their subspecific status: S. a. albofuscus: West Africa. FA: 29.7 (29–31) mm, n = 7; GLS: 13.9 (13.1–14.6) mm, n = 4; C–M3: 4.9 (4.7–5.1) mm, n = 8. S. a. woodi: eastern part of geographical range. FA: 30.1 (29–34) mm, n = 27; GLS: 13.6 (12.7–14.9) mm, n = 21; C–M3: 5.2 (4.9– 5.6) mm, n = 21. Similar Species  Three other African vespertilionids have white wings: Pipistrellus rendalli.Two upper incisors on each side. Penis not extremely long. P. tenuipinnis. Two upper incisors on each side. Penis not extremely long. Glauconycteris gleni.Two upper incisors on each side. Brown reticulation lines on arm-wing. Penis not extremely long. Note: the wings of some other species, e.g. Mimetillus moloneyi, can become very pale after long preservation in alcohol, but these species have two upper incisors on each side. Distribution  Endemic to Africa. Known only from a few scattered localities, mainly in the Sudan Savanna, Guinea Savanna and ZambezianWoodland BZs but with some records in the Rainforest BZ and the Rainforest–Savanna Mosaic in West Africa, and in the Coastal Forest Mosaic BZ. Apparently absent from the Congolian rainforest. Distributed from Gambia to Cameroon, and from Uganda and Kenya to KwaZulu–Natal, but not yet recorded from all countries within this range. Mapped from country checklists (see order Chiroptera), Hill (1974b), Kearney & Taylor (1997), other literature (see below) and museum records. Habitat  In West Africa, recorded from Sudanian woodland savannas including those dominated by Isoberlinia; Yola Marsh in Nigeria; and mosaic of grassland and thicket in Shai Hills, Ghana (Rosevear 1965, Grubb et al. 1998). No evidence of occurrence in forests in West Africa. Elsewhere, recorded in Zoka Forest, Uganda (Kityo & Kerbis 1996); East African coastal mosaic in Kenya (Whitaker & Mumford 1978); riverine forest with Hyphaene palms in S Malawi (R. C. Wood, in Kershaw 1922) and remnants of this forest ca. 80 years later (Happold & Happold 1997); mopane woodland in Mozambique (Smithers & Lobão Tello 1976), and Dune Forest in KwaZulu–Natal (Kearney & Taylor 1997).

Scotoecus albofuscus

Abundance  Very rarely collected and considered rare throughout its geographic range, but the use of elevated mist-nets might result in an increase in records (see below). Adaptations  Aspect ratio low; wing-loading medium. Flight fast with very poor manoeuvrability. One bat whose flying abilities were tested could not fly around a 3.5×3.5 m room by banking, and did not attempt to turn by stalling-and-twisting (M. Happold unpubl.). Mr R. C. Wood (in Kershaw 1922) noted that these bats roosted ‘among the leaves of low Hyphaene palms in forest, where they appear to be moderately numerous, though rarely seen’. Foraging and Food  Based on wing morphology and the flying characteristics of one individual, predicted to forage by moderately fast-hawking mainly in open spaces moderately high above ground and above canopies of trees. This, and the difficulty of catching this species at its day-roosts, might explain why S. albofuscus is rarely collected: it might be caught more often in elevated mist-nets in the future. The stomachs of three individuals in Kenya contained mainly Hemiptera and Coleoptera, with some Lepidoptera, Diptera and other insects (Whitaker & Mumford 1978). Echolocation  No comparable information available but one call is illustrated in Monadjem et al. (2010). Social and Reproductive Behaviour  Several may be taken together nestling down among the leaves (R. C. Wood in Kershaw 1922). Reproduction and Population Structure  At 16° 32'S (Chiromo, Malawi), one / was post-lactating in early Apr (M. Happold unpubl.). At 28° 17' S (St Lucia, KwaZulu–Natal, South Africa), a newly caught / gave birth to twins in late Nov (Kearney & Taylor 1997).

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Scotoecus hirundo

Predators, Parasites and Diseases  Ectoparasites include a bat-fly Basilia robusta (Diptera: Nycteribiidae) and mites Spinturnix walkeri (Acari: Spinturnicidae) and Notoedres sp. (Acari: Sarcoptidae) (Anciaux de Faveaux 1984). Conservation  IUCN Category: Data Deficient. Measurements Scotoecus albofuscus FA: 30.0 (29–34) mm, n = 35 WS (a): 228.6 (210–240) mm, n = 7 TL: 88.3 (76–95) mm, n = 29 T: 33.1 (27–41) mm, n = 30 E: 12.1 (9–15) mm, n = 29

Tr: 4.0, 4.4 mm, n = 2 Tib: 11.5 (10–13) mm, n = 3 HF: 8.5 (8–9) mm, n = 5 WT: 7.1 (5.0–9.5) g, n = 11 GLS: 13.6 (12.7–14.9) mm, n = 25 GWS: 10.5 (10.0–11.2) mm, n = 10 C–M3: 5.1 (4.7–5.8) mm, n = 30 Throughout geographic range (BMNH, HC, HZM, NMZB, ROM, SMF and literature) Key References  Hill 1974b; Kearney & Taylor 1997; R. C.Wood in Kershaw 1922. Meredith Happold

Scotoecus hirundo  Dark-winged Lesser House Bat (Swallow-like Lesser House Bat) Fr. Scotèce hirondelle; Ger. Schwarzflügel-Hausfledermaus Scotoecus hirundo (de Winton, 1899). Ann. Mag. Nat. Hist., ser. 7, 4: 355. Gambaga, Ghana.

Taxonomy  Originally Scotophilus hirundo. Synonyms: albigula, artinii, falabae, hindei. Subspecies: uncertain (see Geographic Variation). The taxonomy of dark-winged Scotoecus is controversial. Ellerman et al. (1953) placed all dark-winged Scotoecus in S. hirundo. Hill (1974b) revised the genus and divided African dark-winged forms into S. hirundo with no subspecies, and S. hindei with subspecies S. h. hindei and S. h. albigula (the latter having more massive canines and cheekteeth). A collection of one S. h. hindei and one S. h. albigula from Malawi (identified by J. E. Hill on basis of craniodental measurements) appeared to provide evidence of sympatry and, consequently, albigula was raised to specific status (Happold et al. 1987), a conclusion followed by Ansell & Dowsett (1988). However, a larger collection of eight ?? and 11 // from Malawi showed that differences in C–M3 and M3–M3 were correlated with sex, the ?? having more massive teeth than the //: consequently, Happold & Happold (1997) followed Koopman (1965, 1975, 1993) and Robbins (1980) who placed all dark-winged Scotoecus in S. hirundo. Robbins (1980) recorded sexual dimorphism in dark-winged Scotoecus from Benin and Ghana (?? larger than //), and also analysed Hill’s measurements (Hill 1974b) to show that his sample of S. hirundo was comprised mostly of // (14 of 17) and his sample of S. hindei was mostly ?? (eight of nine). Taylor & van der Merwe (1998) conducted a principal component analysis of seven cranial characters of 19 dark-winged Scotoecus to test Koopman’s (1993) hypothesis that hindei, albigula and hirundo are synonymous. They found that five hirundo from Cameroon, Uganda and Ethiopia, four falabae from Cameroon, four hindei from Ethiopia and Uganda plus one from Malawi, four other specimens referred to as S. cf. hindei from Malawi, and one albigula from Malawi, separated into five clusters, suggesting that S. hirundo, S. hindei and S. albigula merited specific status, and that the specimens referred to as S. cf. hindei could not be grouped with any of the above species. Sex was not taken into consideration in this study. Cotterill (2001e) measured FA and eight craniodental parameters of 11 female and two male darkwinged specimens from Zambia and, based on differences in skull proportions, concluded that one / represented S. hindei and the

others S. albigula. The specimen he referred to S. hindei was larger and outside (albeit sometimes very marginally) the ranges of specimens referred to S. albigula, in the following measurements and ratios: C1–C1, C1–C1 as percentage of GLS, M3–M3, M3–M3 as percentage of GLS and C–M3. Although these differences indicate the possible presence of two species, the sample size is too small for statistical analysis. Pending a revision of dark-winged forms that takes sexual dimorphism into consideration, all dark-winged forms are provisionally treated here as S. hirundo. Chromosome number (Kenya, as S. hindei): 2n = 30; aFN = 50 (Nagorsen et al. 1976). Description  Very small microbat without noseleaf and with tail more or less fully enclosed in interfemoral membrane; four or five upper cheekteeth, five lower cheekteeth and only one upper incisor on each side; ears well separated and short for a vespertilionid (9– 14 mm); FA: 28–40 mm; wings blackish-brown; penis extremely long (14–16 mm); anterior lower premolar at least two-thirds height of posterior premolar, usually only slightly shorter. Sexes similar in colour; ?? on average slightly larger in some body measurements and with more massive teeth, longer upper toothrow and wider M3– M3 (Robbins 1980, M. Happold unpubl.). Pelage soft, dense, with no sheen; mid-dorsal hairs 4–6 mm. Dorsal pelage chocolate brown, or medium to pale sepia brown; hairs unicoloured (Malawi) or with basal half off-white, terminal half sepia brown (West Africa). Ventral pelage grey, beige, greyish-white or pale beige: chest sometimes paler. Head somewhat flattened; muzzle broad, flat, dark brown, almost naked. Nostrils opening sideways from small transverse pad. Ears widely separated, comparatively and relatively short for a vespertilionid, oval; tip rounded. Tragus relatively short (41.9 [38–46]% of E); somewhat hatchet-shaped, anterior margin strongly concave; posterior margin with smooth, obtuse angle just above midheight, tip rounded (Figure 144c; Malawi). Eyes very small. Wings and interfemoral membrane dark brown, slightly translucent. Penis extremely long, 14–16 mm, ca. 25–30% of HB. Skull as described in Genus Scotoecus. Less flattened than in S. albofuscus. Upper canine with anterior surface flat and shallowly 669

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grooved. Anterior upper premolar usually present (at least on one side) but minute, fully displaced lingually, sometimes loose and possibly easily lost. Posterior upper premolar pointed, in contact with canine and reaching two-thirds to three-quarters height of canine. Posterior upper molar with three ridges (cf. Scotophilus). Anterior lower premolar at least two-thirds height of posterior lower premolar, usually only slightly shorter (cf. Nycticeinops). Sexual dimorphism in craniodental characters is illustrated by differences in M3–M3 – in Benin and Ghana, ??: 6.6 (6.3–6.8) mm, n = 11; //: 6.4 (5.8–6.6) mm, n = 19 (Hill 1974b in Robbins 1980). In Malawi, ?? 7.7 (7.6–7.8) mm, n = 6; // 7.3 (7.1–7.6) mm, n = 10 (M. Happold unpubl.). Dental formula: 1113/3123 = 30 or 1123 /3123 = 32. Geographic Variation  Koopman (1994) recognized four subspecies, S. h. hirundo (Senegal to Benin, and possibly to Ethiopia), S. h. falabae (Nigeria and Cameroon), S. h. hindei (Southern Sudan and Somalia to SE DR Congo) and S. h. albigula (Kenya to Angola). However, the status and distribution of these forms is uncertain (see Taxonomy). A preliminary survey of FA, C–M3 and M3–M3 measurements (made by different people) suggests that there is geographic variation, with an increase in size (in both sexes) from Ghana/Benin (Robbins 1980), to Sudan/Ethiopia, to Kenya/ Tanzania, to Malawi. For example, in ??, mean FA for these areas (in the above order) are 33.4 (n = 11); 33.6 (n = 5); n. d.; 35.4 (n = 18). Similarly, mean C–M3: 5.1 (n = 11); 5.2 (n = 6); 5.5 (n = 9), 6.0 (n = 8). Similarly, mean M3–M3: 6.6 (n = 11), 6.6 (n = 4); Kenya/Tanzania n. d.; 7.7 (n =  6). West African specimens have bicoloured dorsal pelage, cf. unicoloured in Malawi. Similar Species  Only one other dark-winged vespertilionid has only one upper incisor on each side and FA: 25 and