Mammals of Africa by Bloomsbury Pty Ltd

mammals of africa volume I

Volume I: Introductory Chapters and Afrotheria (352 pages)

mammals of africa volume II prImates

Introductory chapters and afrotherIa edited by jonathan kingdon, david happold, michael hoffmann, thomas butynski, meredith happold and jan kalina

edited by thomas butynski, jonathan kingdon and jan kalina

Volume II: Primates (560 pages)

mammals of africa

Volume III: Rodents, Hares and Rabbits (784 pages)

volume III

mammals of africa volume Iv

rodents, hares and rabbIts

hedgehogs, shrews and bats edited by meredith happold and david happold

edited by david happold

Volume IV: Hedgehogs, Shrews and Bats (800 pages)

mammals of africa

Volume V: Carnivores, Pangolins, Equids and Rhinoceroses (560 pages)

mammals of africa volume vI

volume v

carnIvores, pangolIns, equIds and rhInoceroses

hIppopotamuses, pIgs, deer, gIraffe and bovIds

edited by jonathan kingdon and michael hoffmann

Volume VI: Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer and Bovids (704 pages)

Contents Volume I: Introductory Chapters and Afrotheria Edited by Jonathan Kingdon, David Happold, Michael Hoffman,Thomas Butynski, Meredith Happold and Jan Kalina Series Acknowledgements Acknowledgements for Volume I Chapter 1. Mammals of Africa: An Introduction and Guide Chapter 2. Thinking Mammals: An Introduction to African Mammals in Science, Natural History and Culture Chapter 3. The Evolution of a Continent: Geography and Geology Chapter 4. Africaâ&#x20AC;&#x2122;s Environmental and Climatic Past Chapter 5. The Biotic Zones of Africa: A Mammalian Perspective Chapter 6. Mammalian Evolution in Africa Chapter 7. Classification Chapter 8. Behaviour and Morphology ORDER HYRACOIDEA Hyraxes ORDER PROBOSCIDEA Elephants ORDER SIRENIA Dugong, Manatees ORDER AFROSORICIDA Otter-shrews and Golden-moles ORDER MACROSCELIDEA Sengis (Elephant-shrews) ORDER TUBULIDENTATA Aardvark Glossary Bibliography Authors of Volume I Indexes Volume II: Primates Edited by Thomas Butynski, Jonathan Kingdon and Jan Kalina Series Acknowledgements Acknowledgements for Volume II Mammals of Africa: An Introduction and Guide ORDER PRIMATES Primates Glossary Bibliography Authors of Volume II Indexes Volume III: Rodents, Hares and Rabbits Edited by David Happold Series Acknowledgements Acknowledgements for Volume III Mammals of Africa: An Introduction and Guide

ORDER RODENTIA Rodents ORDER LAGAMORPHA Hares and Rabbits Glossary Bibliography Authors of Volume III Indexes Volume IV: Hedgehogs, Shrews and Bats Edited by Meredith Happold and David Happold Series Acknowledgements Acknowledgements for Volume IV Mammals of Africa: An Introduction and Guide ORDER ERINACEOMORPHA Hedgehogs ORDER SORICOMORPHA Shrews ORDER CHIROPTERA Bats Glossary Bibliography Authors of Volume IV Indexes Volume V: Carnivores, Pangolins, Equids and Rhinoceroses Edited by Jonathan Kingdon and Michael Hoffmann Series Acknowledgements Acknowledgements for Volume V Mammals of Africa: An Introduction and Guide ORDER CARNIVORA Carnivores ORDER PHOLIDOTA Pangolins ORDER PERISSODACTYLA Equids and Rhinoceroses Glossary Bibliography Authors of Volume V Indexes Volume VI: Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer and Bovids Edited by Jonathan Kingdon and Michael Hoffmann Series Acknowledgements Acknowledgements for Volume VI Mammals of Africa: An Introduction and Guide ORDER CETARTIODACTYLA Pigs, Hippopotamuses and Ruminants Glossary Bibliography Authors of Volume VI Indexes 1

Series Editors Jonathan Kingdon was born in Tanganyika and has spent the better part of his life in Africa. He has been acclaimed as both a leading academic and a prominent artist, awarded several prizes and medals, and is Vice-President of Fauna & Flora International. The millennium issue of American Scientist named his seven-volume East African Mammals: an Atlas of Evolution in Africa as one of the ‘One Hundred Books that Shaped a Century of Science’. Richard Dawkins describes him as “a world class zoologist, ecologist and writer....a Living World Treasure...an artist with words and a poet with images”. His Island Africa won the1990 Sir Peter Kent Conservation Book Prize, and was described in New Scientist as “a rare and extraordinary view of the natural world, a dynamic and imaginative vision”. Many of his 16 books (totalling 5000 pages) have been translated into languages other than English and he has published numerous papers, articles and chapters. Exhibitions of his work have been shown in museums and galleries in Africa, Australia, Europe, Asia and the USA. David Happold was educated at Peterhouse, Cambridge and the University of Alberta. After completing his PhD, he held academic positions at the University of Khartoum (Sudan), University of Ibadan (Nigeria), University of Malawi, and The Australian National University (formerly Reader in Zoology; currently Emeritus Fellow). His primary research interests are ecology, demography, biogeography and conservation of African small mammals, and his fieldwork was conducted over 16 years in African deserts, savannas, rainforests and mountains. He has published over 70 papers on African mammals in international journals (often with his wife Meredith as co-author), and has contributed to or written several books including Large Mammals of West Africa, Ecology of African Mammals (with M. J. Delany), The Mammals of Nigeria, and African Naturalist. For his research on African mammals, he was awarded the degree of Doctor of Science at the University of Cambridge in 1997, and was elected an Honorary Member of the American Society of Mammalogists. Tom Butynski is a conservationist and ecologist who has worked in Africa for 35 years, mostly in Botswana, Kenya, Uganda and Equatorial Guinea. He has published more than 150 papers and articles and is currently a member of four IUCN/SSC Specialist Groups (Primates, Antelopes, Afrotheria, Wild Pigs). Dr Butynski has served as Director of the Institute of Tropical Forest Conservation in Uganda, Director of Conservation International’s Eastern Africa Biodiversity Hotspots Program, Vice-Chair of the Africa Section of the IUCN/SSC Specialist Group, and Senior Editor of the journal African Primates. He now works for the Zoological Society of London as Director of the King Khalid Wildlife Research Centre in Saudi Arabia.

Mike Hoffmann trained as a mammalogist at the Mammal Research Institute, University of Pretoria, South Africa with the late John Skinner, and at the Wildlife Conservation Research Unit at the University of Oxford, UK. He has since moved into the field of international biodiversity conservation, working with Conservation International and IUCN – the International Union for Conservation of Nature. Currently, Mike serves as Senior Scientist to IUCN’s Species Survival Commission – the largest network of conservation professionals in the world – helping to ensure uptake of Commissionbased research and expertise in appropriate decision-making and policy arenas. A member of the IUCN SSC Afrotheria, Antelope and Canid Specialist Groups, he has co-edited six books and authored 21 peer-reviewed papers, six in the journals Nature and Science. Meredith Happold read Zoology at Monash University in Australia and obtained her PhD on the evolution and adaptive significance of social behaviour in ten endemic rodents from very diverse Australian habitats. She then spent several years in Nigeria and published observations on the fruit bats of western Nigeria, with David Happold as co-author. Since 1977, Meredith has held a Visiting Fellowship at the Australian National University where, although primarily involved in research, she also taught human and vertebrate biology, animal behaviour and human ecology. Since 1982, her research has focused on African bats (especially those of Malawi) and, to a lesser extent, on African rodents. She has published, often with David Happold as co-author, 35 papers including 19 on bats (distribution and zoogeography, reproduction and reproductive strategies, social behaviour, renal form and function, taxonomy, bat-banding and parasites). She has also researched echolocation, wing-morphology and flight, foraging behaviour and community structures of Malawian bats. Jan Kalina is a conservation biologist. She studied the ecology and behaviour of the Black-and-white Casqued Hornbill in Kibale Forest, Uganda for her doctorate in wildlife ecology from Michigan State University. As a research wildlife biologist for the US Forest Service she worked on the recovery of the Critically Endangered Puerto Rican Parrot. Her work with mammals began at Wildlife Conservation Society’s Bronx Zoo in New York and at Colorado State University. As a CARE employee in Uganda, she lobbied for the establishment of the Bwindi Impenetrable National Park and Mgahinga Gorilla National Park. Living in the Impenetrable Forest for six years, she helped build and fund park infrastructure, including the Institute of Tropical Forest Conservation. As a lecturer at Makerere University, she taught today’s leaders in conservation and science in Uganda. Her publications mainly concern gorillas, rainforests and threatened species. Jan is currently joint-owner of Soita Nyiro Conservancy in Laikipia, Kenya.

Family Giraffidae

Giraffe Giraffa camelopardalis.

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Behaviour and Morphology

The morphology of heads and their evolutionary origins The concentration of disparate activities that are welded together in vertebrate skulls would seem extraordinary were it not so familiar. The further elaboration of head shapes, especially by horns, antlers, tusks and ossicones, has provided humans with symbols and trophies from the earliest times, yet remain a poorly explored aspect of morphology. It is partly our human fascination with heads and partly the superior survivorship of teeth that has ensured that fossil heads dominate palaeontological collections and provide the data for much of our knowledge of the evolution of heads. Because the fore-ends of primitive chordates and vertebrates were the first to encounter both food and obstacles, basic sense organs and a ‘mouth’ developed there. The polarity of organisms and their symmetrical organization crosses all the higher taxa (Wainwright 1988). The need to respond to light, chemical, electric or vibrational signals led to the differentiation of cells until they developed into eyes, nose and ears, each encapsulated in compartments that shared the upper part of what became the skull. The lower section of this structure became a hinged mandible and eventually both jaws developed teeth. In many predatory vertebrates, from the most primitive to the most advanced, the size of jaws closely reflects the size and toughness of the prey. Furthermore, a gross expression of animal proportions is the size of heads relative to the size of bodies. Apart from whales, the most extreme examples of disproportionately large or small heads are to be found in bats. This is partly because the head and jaws get little or no heavy-duty help from the limbs in the behaviour patterns of capturing and processing prey. Thus Moloney’s Mimic Bat, a species that only takes small, soft-bodied prey, has no need for a large head while the Heart-nosed Bat Cardioderma cor seizes robust, often vertebrate, prey with its large jaws and must quickly subdue it with deep, damaging bites powered by massive jaw muscles. The head of the former is about one-seventh of the combined head–body volume, whereas Cardioderma heads are closer to a quarter! One of the most fundamental expressions of cranial morphology concerns the sizes of sensory activities in the head. It is in the relative

sizes of compartments and in permutations of connecting bridges, struts and welds that the species-specific morphology of mammal heads becomes obvious.Wherever sufficient data exist on the ecology and behaviour of species, we find superb and detailed examples of forms evolving morphological modifications that serve very precise behavioural functions. This is particularly obvious in the relative sizes of, say, orbits, olfactory equipment and auditory bullae in mammal skulls. Even more explicit are the functional shapes of eyes, ears, noses and sensory whiskers in mammals as different as a Galago, a Bat-eared Fox Otocyon megalotis, an Aardvark Orycteropus afer or an Aquatic Genet Genetta piscivora.

Thomas’s Galago Galagoides thomasi

Bat-eared ‘Fox’ Otocyon megalotis

Aardvark Orycteropus afer

Aquatic Genet Genetta piscivora Head/body proportions in Mimic Bat Mimetillus moloneyi and Heart-nosed Bat Cardioderma cor (after Kingdon 1974).

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Heads to illustrate super-development of different sensory faculties.

Head-shape and chewing

Skull outlines of four pairs of related mammal species, showing the role of chewing muscles (masseter and temporalis) influencing skull shape. Weak chewers (top row) have shallow muscle attachments on relatively gracile skulls; powerful chewers (lower row) have robust skulls with deep muscle attachments. a. African Elephant Loxodonta africana and Pleistocene Elephant Elephas recki. b. Human Homo sapiens and Western Gorilla Gorilla gorilla. c. Link Rat Deomys ferrugineus and Giant Squirrel Protoxerus stangeri. d. Bushbuck Tragelaphus scriptus and Mountain Reedbuck Redunca fulvorufula.

Head-shape and chewing The mechanics of jaw function are nowhere more simple and explicit than in the Savanna Elephant Loxodonta africana, which has a backand-forth mode of chewing with very little sideways action. The short, heavy mandible hangs from the temporalis muscle as if it were the chair of a swing. Here the ‘ropes’ of the temporalis are at rightangles to the lower tooth rows that resemble the chair’s seat. The swing seat is pulled back towards the chest, in a loose, easy arc, by the digastric muscle. Coming forward again it closes over the mouthful of food and, clamping hard against the upper toothrow, grinds forward under the combined force of masseter and temporalis muscles. In one of those confounding inversions of expectation, modern African elephants are, in terms of dentition and diet, relatively primitive. Until quite recently (about 19,000 years ago) the commonest and most widespread elephant species in Africa was Elephas recki, a close relative of the Indian elephant. Its fossils are abundant and widespread, from South Africa to the Sahara. Elephas recki (like its Indian congener E. indicus and its Nordic relative, the mammoth, Mammutus) was a dentally advanced species with very deep, multi-plated molars that could masticate coarse, abrasive foods more efficiently than Loxodonta africana. However, it is not just teeth and their bony buttressing that take the strain of chewing. More force must be exerted on tougher foods and that force must be exerted by larger, longer muscles. And increased muscle forces reshape the bones onto which they are attached, such is the adaptability of bone form and function. In the case of elephant skulls, this resulted in the phylogenetic elevation of paired, honeycombed arches that absorb all the forces of chewing and cowl the braincase like a bonnet (tool language again). Although E. recki had a rather small skull relative to its body, its forehead was immense and a large part of this

enlargement can be confidently correlated with the behavioural need to enlarge the temporal muscles in order to exert more force on a formidable battery of chewing teeth. Comparing the skulls and teeth of L. africana and E. recki provides a vivid illustration of how changed diets (often triggered by climate change) drive modifications to tooth and skull structure. New stresses and strains on teeth call for changed mechanics in the skull, the external expressions of which are distinctive head shapes and silhouettes. Understanding the mutability of bones is, in turn, impossible without taking into account changes in the muscle forces exerted upon the bones. The interplay between behaviour, diet, teeth, the mechanics of mastication and the gross morphology of bones and muscles are evident in comparison between a dissection of an extant Savanna Elephant’s head with a reconstruction of the same muscles on an E. recki skull. Another simple demonstration of ‘function and form’ emerges from comparing the head proportions of two similar-sized antelopes: the Bushbuck Tragelaphus scriptus and Mountain Reedbuck Redunca fulvorufula.The former has a slender head with long-lipped mouth; the latter’s head is short-mouthed and wedge-shaped. These differences begin with tooth proportions. The soft-foliage-eating Bushbuck has modest, shallowly rooted masticatory teeth set in equally shallow jaws, but the broad-leaved shrubs and herbs that it eats need a wide maw. The tough-grass-eating Mountain Reedbuck instead has heavily crenellated cheekteeth that are firmly rooted in a well-reinforced, deep-jawed skull and mandible. It is the expansion of teeth and their rooting, together with massive masseter and temporalis muscles, that open up that mandibular wedge. It is the nibbling of tough grasses that calls for that tight, short-lipped, muscular mouth. The external expressions of such dietary differences are an elongated sleek face in the Bushbuck, a relatively short, chunky face for the Mountain Reedbuck (see drawings). 127

Family CERCOPITHECIDAE

Cercocebus torquatus Red-capped Mangabey (White-collared Mangabey) Fr. Mangabé à collier blanc; Ger. Rotkopfmangabe Cercocebus torquatus (Kerr, 1792). Animal Kingdom, p. 67. West Africa.

Red-capped Mangabey Cercocebus torquatus adult male.

Taxonomy Monotypic species (Kingdon 1997, Groves 2001, 2005b). Previously classified at the subspecies level, Cercocebus torquatus torquatus, with the Sooty Mangabey C. t. atys and the Whitenaped Mangabey C. t. lunulatus comprising the other two subspecies (Dorst & Dandelot 1970, Groves 1978). Synonyms: collaris, crossi. Chromosome number: 2n = 42 (Dutrillaux et al. 1979). Description Large, slate-grey, moderately long-muzzled, semiterrestrial monkey with red crown and white collar. Sexes alike except markedly dimorphic in size. Adult // weigh about 55% as much as adult ??. Face skin dark grey to black with eyelids bright white and sharply contrasting with face skin. Ears naked, dark greyish-black, extending above white collar. Crown russet to dark maroon-red, outlined by the white collar and broad temporal line, and may show whirl or slight medial partition. Chin, sides of the neck, cheeks and throat white. Cheek stripe grey, of variable thickness, separating white cheeks from white throat. Whiskers white, short, directed backward. Nape with white patch. Dorsum chestnut brown to dark grey with darker dorsal stripe. Outer limbs darker grey at distal end and on feet. Ventrum and inner limbs sharply-demarcated white. Hands and feet with brownish-black skin.Tail darker grey than dorsum, whitens distally with white tuft. Callosities brownish-pink, fused in ?, separate in /. Skin and pelage colour different in infants: skin of face, hands and feet pink; eyelids lighter than face skin. Head auburn. Dorsum brownish-grey. Crown red by four months of age; full adult colouration by five months (Field 1995a). Suborbital fossae shallow relative to other Cercocebus spp., whereas the paranasal ridges are the most pronounced of the Cercocebus spp. (McGraw & Fleagle 2006).These features, and the presence of straight temporal lines are, among the Cercocebus species, the most similar to 186

Lateral, palatal and dorsal views of skull of Red-capped Mangabey Cercocebus torquatus adult male.

Mandrillus.This suggests that C. torquatus is the sister taxon to Mandrillus within the Cercocebus–Mandrillus clade (McGraw & Fleagle 2006). Geographic Variation Skull size and breadth substantially variable across populations, although not in a pattern justifying distinct subspecies (Groves 1978). Length of white on tail also variable (90–235 mm) across populations (Hill 1974, Groves 1978). Similar Species None within geographic range. Distribution Endemic to West Africa. Rainforest BZ. From W Nigeria (from west of Niger R.) through Cameroon (east as far as E Dja Reserve), Equatorial Guinea (Rio Muni), and the Gabon coast (to south of Nyanga R. in Mayumba N. P.) into S Congo (ConkouatiDouli N. P.) (Rosevear 1953, Gartlan & Struhsaker 1972, Happold 1973, Schlitter et al. 1973, Blom et al. 1992, Groves 2001, Oates 2011). Distribution patchy across range. Almost all populations within 350 km of the coast (Maisels et al. 2007). Western limit uncertain. Happold (1973) stated that the Redcapped Mangabey occurs in forest relics in the derived savanna zone of W Nigeria, near the Dahomey Gap (e.g. within the Igangan F. R.).

Cercocebus torquatus

Schlitter et al. (1973) state that the species might occur in remnant forests west of the Nigerian border in Bénin. This is supported by Campbell et al. (2008), who obtained verbal reports of Red-capped Mangabeys in SE Bénin. Habitat Occupies a variety of habitat types, although principally relatively wet habitats: swamp forest or seasonally/intermittently inundated forests, mangroves, lowland riverine and coastal forest. Also in primary and secondary terra firma dry forest, and utilize agricultural areas such as rice fields and gardens (Sabater Pi & Jones 1967, Jones & Sabater Pi 1968, Gartlan & Struhsaker 1972, Hill 1974, Napier 1981, Garcia & Mba 1997, Gautier-Hion et al. 1999, Cooke 2005). Garcia & Mba (1997) observed Red-capped Mangabeys frequently associated with nipa palm trees Raphia spp. in riparian forests along the Uolo R. and Laña R. in Monte Alen N. P., Equatorial Guinea. The majority of sightings by Sabater Pi & Jones (1967) were in lowland forests (<350 m asl) of the coastal region, in riverine forest a confluences of rivers and the ocean, or in swamps characterized by tangles of red mangrove Rhizophora mangle roots and stands of Pandanus sp., with dense closed canopies 10–12 m high composed of plants such as Raphia spp., Sclerosperma manni, Mytragine ciliata, Oxytenanthera abissinica and Cytosperma senegalense. Vegetation near the coast of Rio Muni where Red-capped Mangabeys were observed was 10–25 m plants such as Phoenix reclinata, Cocos nucifera, Canavalia ensiformis, Urena lobata, Pennisetum purpureum and Terminalia cattapa (Jones & Sabater Pi 1968). Annual rainfall across the geographic range of Red-capped Mangabeys ca. 1400–3200 mm with wet seasons from Mar to May (minor) and Sep to Dec (major) (Mitani 1991, Blom et al. 1992, Garcia & Mba 1997). Found at altitudes from sea level along the Atlantic coast to <1000 m, with most reported locations below 500 m. Abundance Abundance decreases from west to east both in Cameroon and Gabon (Matthews & Matthews 2002); rarely encountered east of Campo Forest in Campo-Ma’an N. P., S

Cameroon (Mitani 1990), and apparently absent in SE Cameroon (Lobéké N. P.) (Davenport & Usongo 1997). Although Gartlan & Struhsaker (1972) found Red-capped Mangabeys in E Dja Reserve, later surveys here did not locate them (Ngandjui 1997). Struhsaker (1972) reported this species as ‘common’ in both Douala-Edéa R. and Korup N. P. Densities have since declined: 0.23 groups/km2 in 1999; 0.14 groups/km2 in 2000 (Waltert et al. 2002); 0.01 groups/km2 in 2004–05 (Linder 2008); 0.02 groups/km2 in 2006–08 (Astaras et al. 2011). Matthews & Matthews (2002) reported 0.51 groups/km2 in Campo Forest, and significantly higher densities in areas of lower levels of human disturbance (0.71 groups/km2 on Dipikar Island, with minimal disturbance, vs. 0.36 groups/km2 in a heavily exploited logging concession area; p<0.01). These censuses indicated Redcapped Mangabeys were largely restricted to the coastal areas, only reaching 80–100 km inland. In Gabon, early reports also suggested that Red-capped Mangabeys are less abundant in the interior (Malbrant & Maclatchy 1949). Non-systematic surveys confirmed this observation in the protected areas of Moukalaba and Sette Cama; Red-capped Mangabeys were reported to be ‘abundant’ although restricted in distribution to areas near the coast (Blom et al. 1992). Cooke (2005) found Red-capped Mangabeys to be the most abundant cercopithecid near the village of Sette Cama, south of Loango N. P., Gabon. Morgan (2007) encountered 36 groups in Loango N. P. and estimated mean group density at 0.97 groups/km2 in this coastal protected area. Generally rare in the interior of Equatorial Guinea (Jones & Sabater Pi 1968, Garcia & Mba 1997), but locally abundant. Declining in all areas of Equatorial Guinea (C. Jones pers. comm. in Wolfheim 1983). In Nigeria, rare or not abundant west of the Niger R. (Happold 1973, Schlitter et al. 1973). In Okomu N. P., SW Nigeria, White, L. (1988) found ca. 85 groups, although the area was threatened by hunting and habitat alteration, worsened by the negative consequences of a programme of agricultural development assistance to migrant farmers (Oates 1995). Here, in 2008–09, Akinsorotan et al. (2011) found 0.1 groups/km2 and 2.2 groups/ km2. Surveys (since 1995) in S Nigeria indicate that the Red-capped Mangabey is locally extinct or rare in a number of areas of the Niger Delta and Akwa Ibom (Z. Tooze pers. comm.). These observations support Kingdon’s (1997) evaluation that the Red-capped Mangabey was once a widespread, successful species, which is declining from all areas where habitat loss to agricultural expansion and intensive hunting occur. Adaptations Diurnal and semi-terrestrial (Jones & Sabater Pi 1968, Mitani 1989, Gautier-Hion et al. 1999). Red-capped Mangabeys spend considerable time on the ground, frequently descend to the ground to feed or to flee when alarmed, and utilize forest strata from the ground to 40 m high in the canopy (Jones & Sabater Pi 1968, Mitani 1989, 1991). In Campo-Ma’an N. P., the vertical distribution of Red-capped Mangabeys differed by activity patterns: ‘escape’ (fleeing observers) occurred on the ground in 85% of 21 observed instances; ‘move’ (locomotion) and ‘socio-rest’ (including both grooming and resting) were recorded most frequently at 0–5 m (ca. 58% of 185 observations and ca. 63% of 161 observations, respectively); whereas ‘feed’ (search, approach, manipulation, consumption of food) correlated positively with distribution of fruit production, with a mode height of 25–30 m (Mitani 1989). 187

Family NESOMYIDAE

Prionomys batesi Bates’s African Climbing Mouse (Dollman’s African Tree Mouse) Fr. Souris arboricole de Bates; Ger. Bates Klettermaus Prionomys batesi Dollman, 1910. Two new African mammals. Ann. Mag. Nat. Hist., ser. 8, 6: 226. Dja River, Bitye, Cameroon. 2000 ft (618 m).

Taxonomy Closely related to Dendromus and Dendroprionomys (see genus profiles). Synonyms: none. Chromosome number: not known. Description Very small mouse with long tail. Pelage short, dense, soft and woolly. Dorsal pelage brownish-grey; hairs dark grey at base, brown at tips. Mid-dorsal stripe absent. Flanks pale grey; hairs dark grey at base, pale grey at tips. Ventral pelage pale grey, similar to flanks; hairs grey at base, paler at tips. Head similar in colour to dorsal pelage. Conspicuous brown eye-ring. Long stiff vibrissae. Ears moderate, darkly pigmented, with sparse brown hairs. Chin, throat and chest grey. Limbs short, with few short hairs. Forefoot with four digits: Digit 1 absent, Digits 2, 3, 4 and 5 thin, each with small pale claw. Large carpal tuberosity on ‘hand’. Hindfoot with five digits: Digit 1 shorter than other digits, with ‘nail-like’ claw, Digits 2, 3, 4 and 5 long and thin, each with small claw; Digit 1 opposable (cf. Digit 5 opposable in Dendromus and Dendroprionomys) (Figure 30); three large callosities at base of digits on hindfoot. Tail long (ca. 122% of HB), brown, with scales arranged in rings; few (if any) small bristles. Skull: upper incisors pro-odont without longitudinal grooves, braincase globular-shaped, molar teeth with tall pointed cusps (Figure 34). Nipples: not known. Geographic Variation None recorded.

Figure 34. Skull and mandible of Prionomys batesi (BMNH 11.5.5.9, holotype).

190

Similar Species Dendromus spp. Cheekteeth molariform; incisor teeth orthodont. Dendroprionomys rousseloti. Cheekteeth with tall pointed cusps; incisor teeth orthodont. Distribution Endemic to Africa. Rainforest BZ (West Central Region, Gabon Sub-region). Recorded from Cameroon, S Central African Republic and W Congo (Odzalla). Habitat Rainforest, especially where there are small shrubs and lianas. Abundance Uncertain; known only from a few specimens at five localities. Adaptations Terrestrial and arboreal. Bates’s African Climbing Mice climb on thin branches, twigs and lianas using the prehensile tail, long digits, callosity on the forefoot and opposable Digit 1 of hindfoot (in a similar way to Dendromus spp.). Burrows are made by digging in forest soil using the pro-odont upper and lower incisor teeth; each burrow is indicated by a little hillock of soil, ca. 30 cm diameter, without a visible entrance hole.The burrow contains a small nest chamber lined with dry leaves. Activity is entirely nocturnal: one individual emerged at 21:00h (three hours after nightfall) and climbed rapidly into nearby shrubs and lianas. Activity alternated between climbing (and foraging) in the forest and 15-minute ‘rest’ periods in the nest (F. Petter unpubl.). Bates’s African Climbing Mice emit a strong odour, not unlike that of some species of shrews.

Prionomys batesi

Foraging and Food Insectivorous. The diet consists (almost exclusively) of leaf-nesting ants, as well as spiders. The ants are captured with the tongue and crushed by the sectorial cusped cheekteeth. Large quantities of prey are consumed (F. Petter unpubl., Genest-Villard 1980). Social and Reproductive Behaviour No information. Reproduction and Population Structure Young are born in a nest chamber, lined with leaves, under the soil (see above).

Measurements Prionomys batesi HB: 69 (54–78) mm, n = 11 T: 93.1 (67–103) mm, n = 10 HF: 14.9 (13–16) mm, n = 11 E: 10.3 (8–11) mm, n = 10 WT: n. d. GLS: 24.4 (21.7–25.8) mm, n = 9 GWS: 13.9 (11.0–15.2) mm, n = 12 M1–M3: 4.7 (4.3–5.0) mm, n = 9 Central African Republic (MNHN)

Predators, Parasites and Diseases No information. Conservation IUCN Category: Data Deficient. Considered as Rare by Schlitter (1989).The very small geographic range and presumed rarity are cause for concern.

Key References Denys et al. 1995; Genest-Villard 1980; Petter 1964, 1966b. Christiane Denys

Genus Steatomys Fat Mice Steatomys Peters, 1846. Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin 11: 258. Tete, Mozambique. Type species: Steatomys pratensis Peters, 1846.

Steatomys sp.

This exclusively African genus is widespread on the continent south of the Sahara and currently eight species are recognized. Four species are confined to West and sub-Saharan Africa, while the other four species occur in southern, central and East Africa. Species in the genus inhabit a variety of savanna habitats. Species of the genus are small with soft shiny unpatterned dorsal pelage ranging in colour from pale grey-brown to brownish-buff. Ventral pelage is pure white. Tail is short, usually 50–60% of headbody length. Muzzle is pointed and ears are rounded. Limbs are short with white feet. Forefeet have four well-developed digits (Digit 1 absent), and hindfeet have five well-developed digits. All digits have long, sharp claws, which may be employed in the digging of burrows. Characteristics of the skull include orthodont incisors; each upper incisor with a single shallow groove on the outer face (usually paler in colour than the ungrooved part); zygomatic plate small and vertical (as in Dendromus) with a masseteric tubercle at lower anterior corner (also as in Dendromus); auditory bullae relatively large; palate wide, widest at level with upper M1 and narrowest at level with upper M3; and ungrooved lower incisors. M1 is the largest molar. M3 is greatly

Figure 35. Skull and mandible of Steatomys jacksoni (HC 2797).

reduced (Figure 35). Species of this genus are readily identified by their rotund shape, short tail and grooved upper incisors. A well-known character of all species in the genus is the ability to lay down substantial amounts of subcutaneous fat (hence the common name ‘fat mouse’), a feature which allows them to enter torpor readily during cold conditions. Hibernation, however, does not occur, and individuals may be active throughout the year. All members of this genus are terrestrial and nocturnal. They construct their own burrows, which may be simple or relatively complex. 191

Family VESPERTILIONIDAE

GENUS Nycticeinops 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

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).

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.

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).

Scotoecus hirundo. Usually heavier (10.2 [8–15] g). FA sometimes longer (28–40 mm). Dorsal pelage chocolate brown, or medium to pale sepia brown; ventral pelage grey, beige, greyish-white or pale beige. Tragus less pointed (Figure 144b). Penis extremely long (14–16 mm). Upper canine with anterior surface flat and shallowly grooved. Anterior lower premolar at least two-thirds height of posterior premolar, usually only slightly shorter.

Geographic Variation Four subspecies (albiventer, australis, fitzsimonsi and schlieffeni) were recognized by Hayman & Hill (1971),

Similar Species Only one other dark-winged vespertilionid in Africa has only one upper incisor on each side and FA <41 mm:

Distribution In Africa, recorded from the Sahara Arid BZ near the Nile R. in N Egypt, and in almost all biotic zones and mosaics south of the Sahara except the Rainforest, South-West Arid and Highveld BZs. Recorded disjunctly from Mauritania to Sudan, Eritrea, Djibouti and Somalia (with outlying localities near the Nile

Nycticeinops schlieffeni

595

Family FELIDAE

Caracal caracal Caracal Fr. Caracal; Ger. Karakal (Wüstenluchs) Caracal caracal (Schreber, 1776). Die Säugethiere 3 (16): pl. 110 [1776]; text 3 (24): 413, 587 [1777]. ‘Vorgebirge der guten Hofnung’, restricted by J. A. Allen (1924: 281) to ‘Table Mountain, near Cape Town, South Africa’.

Caracal Caracal caracal.

Taxonomy Seven subspecies have been recognized (Smithers 1975), but their validity seems questionable and they are perhaps best considered as geographical variants. Synonyms: aharonii, algira, bengalensis, berberorum, coloniae, corylinus, damarensis, limpopoensis, lucani, medjerdae, melanotis, melanotix, michaelis, nubica, poecilotis, roothi, schmitzi, spatzi. Chromosome number: 2n  = 38 (Hsu & Arrighi 1966). Description Medium-sized, robustly built, uniformly coloured cat with dark-backed ears with distinct terminal tufts averaging 45 mm in length. Head typically cat-like with short face prominently marked with black and white patches, notably around eyes and mouth. Chin and throat are white, with variable extent of white on cheeks.The large ears pointed with a tuft of longish black hair at the tip. Back surfaces of the ears predominantly black but liberally sprinkled with white hairs; inner surfaces white. Hindquarters slightly higher than shoulders due to powerfully muscled and elongated hindlegs. Pelage thick but short and soft, the colour varying from pale reddish-fawn with a greyish infusion, to rich brick-red. Underparts, including inner leg surfaces, paler than rest of pelage and faint spotting or blotching may or may not be present. Extent of white on chest and belly very variable. Tail is similar in colour to the dorsal aspect and approximately one-third of head–body length. In temperate regions, guard hairs of the winter coat up to 30 mm long but usually less than 20 mm in the summer coat. Colour of guard hairs variable but pale at the base, with either broad annulations in off-white, dark brown or black. In temperate regions underfur in winter is thick but in summer greatly diminished. Paws proportionally large, with five digits on the forefeet and four on the hindfeet. The first digit on the forefoot is higher than the other four and does not make contact with the ground but is equipped with a substantial claw (dew claw). Claws of the forefeet are sharp, retractable and curved (about 25 mm along the upper curve). The claws on the hindfeet are less curved but larger (exceeding 30 mm). Males are consistently larger than // in all respects (Stuart & Stuart 1992). Skull high and rounded in profile, with particularly short, blunt rostrum. Supraoccipital crest well developed and in older animals 174

Caracal Caracal caracal.

joined at right-angles by sagittal crest. Ear bullae large and well developed. An overall heavily built skull, with stout mandible and tall, narrow coronoid process (Skinner & Chimimba 2005). Dentition is typically felid; the second upper premolar is usually absent (present in only eight of 100 skulls examined by Stuart & Stuart [1985]), unlike the Serval Leptailurus serval in which they are usually present. GeographicVariation Separating the seven described subspecies occurring in Africa is based in most cases on small samples and minor differences in pelage colouration (Smithers 1975). Animals from arid low rainfall areas tend to be paler than those from higher rainfall regions. Within the southern African populations, from where the largest samples are available, there is considerable variation in overall pelage colour. Melanic individuals have been recorded from Kenya and Uganda (Rosevear 1974). Similar Species Leptailurus serval. Sympatric in most parts of the range, except, most notably, in the drier western and south-western parts of southern

Caracal caracal

the Middle East and Turkey, and then eastwards to C India and northwards to Kazakhstan and Turkmenistan (Stuart 1984, Nowell & Jackson 1996, Sunquist & Sunquist 2002).

Lateral view of skull of Caracal Caracal caracal.

Africa. Distinctive spotting and barring, a ringed tail and a large white spot at back of each ear, with terminal tuft absent. Profelis aurata. Largely allopatric, occurring in the forest zone of equatorial Africa. Dark-backed ears that lack a terminal hair tuft, have variably dark spotted upper- and underparts, the latter pelage being white to off-white. Distribution The Caracal is widely distributed on the African continent, being absent only from the equatorial forest belt and from much of the central Sahara, but they are present in the montane massifs of that desert and its fringes, including the Adrar des Iforas in NE Mali (Sidiyène & Tranier 1990), Hoggar and Tassili mountains of SE Algeria and the Saharan Atlas (De Smet 1989, Kowalski & RzebikKowalska 1991), the Aïr of Niger (Dragesco Joffé 1993), and edges of the great sand areas of the Eastern Great Erg. The presence of Caracal in the savanna regions of the Congo basin, north of the Congo R., is equivocal.The form lucani is described from Landana, which is in Cabinda, Angola. Coetzee (1977) gave the range of this subspecies as N Angola, Gabon, and south-western DR Congo (it is also mentioned by Malbrant & Maclatchy 1949); however, the species is not mentioned in N Angola by Crawford-Cabral (1989a) or Hill & Carter (1941). Extralimital to the African continent, the Caracal has a very wide distribution, occurring in suitable habitats to the Arabian Peninsula,

Habitat The Caracal occupies a wide variety of habitats from semidesert to relatively open savanna and scrubland to moist woodland and thicket or evergreen/montane forest (as in the Western Cape of South Africa), but favours drier woodland and savanna regions with lower rainfall and some cover (Stuart & Wilson 1988). They range from sea level in many parts of their range, to 2,260 m in the Moroccan Atlas Mts (Cuzin 2003) and 3300 m in the Ethiopian Highlands (Yalden et al. 1996). In KwaZulu–Natal (South Africa), recorded in commercial forests (Eucalyptus spp. and Pinus spp.) (Rowe-Rowe 1992a), and in areas of intensive wheat cultivation in the Western Cape they hide up during the day in adjacent scrubland, moving at night on open fields. Particularly common in the Cape fynbos, an area of low and often dense scrub (Stuart 1981). On the Pro-Namib plains of Namibia Caracal are closely associated with lightly wooded dry river courses and rocky inselbergs, and on South Africa’s central Karoo plains they are strongly associated with rugged mountain and hill ranges. Abundance The Caracal is common in parts of its range, especially in South Africa and S Namibia where it is expanding into new, and recolonizing vacant, areas (Stuart 1984, Stuart & Wilson 1988). Found in all major national parks and many other conservation areas within their range (Stuart 1984, C. Stuart & T. Stuart pers. obs.). In central and West Africa, where they are largely absent, densities are apparently lower, possibly due to finer partitioning of resources in a more diverse carnivore community (Kingdon 1977, Stuart 1984, Nowell & Jackson 1996), while north of Sahara densities are very low, and Caracal could be on the verge of extinction at least in Morocco (F. Cuzin pers. comm.). In South Africa, Grobler (1981) estimated the population density of Caracal in Mountain Zebra N. P. in the Eastern Cape as 1/2.6 km2, while Avenant & Nel (1998) recorded a density of 0.23–0.47/km2 in West Coast N. P. in the Western Cape. Adaptations Large pinnae and well-developed ear bullae reflect the important role played by hearing in hunting, but colouring and tufted tips appear to be elaborated to serve communication by means of frequent ritualized ear movements (Kingdon 1977). Skull and dentition (in particular the sharp canine teeth) are stoutly structured, an adaptation to taking relatively large prey in proportion to its size.The masseter and temporalis muscles are well developed to deliver the killing bite. As in other felids, the teeth of the carnassial shear are adapted to cutting, although the teeth in the lower jaw show no adaptation to crushing. Paws notably large and the well-developed claws also indicate an ability to take larger prey. Normally terrestrial, they are adept tree climbers, using their powerful dew claws in this pursuit. Caracals are principally nocturnal, although in protected areas they may occasionally hunt during the day, especially in the early morning and late afternoon, and particularly on cooler days or if overcast.

Caracal caracal

Foraging and Food Caracal prey mainly on small- to mediumsized mammals, from small murids to antelope up to the size of around 50 kg, but they will also take birds, reptiles, invertebrates, fish and some plant matter. The most detailed studies of diet have 175

Family Bovidae

Tragelaphus angasii Nyala Fr. Nyala; Ger. Tiefland-Nyala Tragelaphus angasii Angas, 1849. Proc. Zool. Soc. Lond. 1848: 89 [1849]. South Africa, KwaZulu–Natal, ‘Hills that border upon the northern shores of St. Lucia Bay, in the Zulu country, lat. 28° south’. [Attribution to Angas, 1849, and not Gray, follows Grubb 2004, 2005].

Taxonomy There are no recognized subspecies. Genetic variation, on the basis of microsatellite, mitochondrial control region and allozyme markers, occurs between the Malawi, Mozambique and KwaZulu–Natal populations and it is postulated that this geographic variation is a function of a distribution pattern stemming from habitat specificity (Grobler et al. 2005). Synonyms: none. Chromosome number: 2n = 55 for ? and 2n = 56 for / (Wurster & Benirschke 1968, Wallace 1980). Description No other antelope shows such extreme sexual dimorphism as the Nyala, with adult ?? almost twice the size of // (Anderson 1976). Females, and young of both sexes, russet brown, with up to 18 vertical white stripes running from shoulders to hindquarters. At one year of age general coat colour of ?? is still same as //, but hair on neck has darkened while that on throat, sides of belly and dorsal ridge has grown noticeably longer than in //. Change in coat colour and length occurs rapidly between 14 and 16 months of age, the age at which spermatogenesis begins. This is accompanied by rapid increases in testis weight and seminiferous tubule diameter. Adult ?? are predominantly charcoal grey; lower legs remain russet brown, stripes and spots are retained and animals develop a white chevron on top of the muzzle. Up to three white spots on the cheeks and white spots on the upper hindleg. Bases of backs of ears white. Dorsal crest of long hair (dark on neck), and a heavy fringe of long hair (slightly darker than hair on body) on underparts of the neck to the belly. Long hair on tail dark, but underside white; this is very obvious in the male dominance ritual. In older individuals the stripes are reduced to three or four, or are absent entirely.Very rarely adult ?? occur that have retained their juvenile pelage (those that have been examined exhibited either cryptorchidism or had undeveloped testes). Adult // may or may not have a white chevron present between the eyes, also have a dorsal crest (though they lack the fringe of hairs on the throat and belly) and the lower parts of the legs do not contrast with the overall body colouration as is the case in ??. Both sexes have glands at the base of the false hooves, but inguinal and preorbital glands are absent. Females have two pairs of inguinal nipples. Aberrant-coloured Nyalas have been recorded. Dixon (1964) reported a ? from Mkuzi G. R., KwaZulu–Natal, which was pale beige in colour, and Lobão Tello & Van Gelder (1975) recorded a pale yellowish-coloured / and a palomino-coloured ? from Zinave N. P. in Mozambique. Only ?? have horns, which are keeled and grow in a shallow spiral. Horns first develop between five and six months (Anderson 1986). In adult ?? the tips of the horns are whitish partly as a result of the frequent ‘horning’ of moist earth when animals visit a waterhole. Age determination is discussed by Anderson (1986): adult dentition is attained at two years by which time the horns are half-grown (reaching mature size at 4.5 years). top: Adult male Nyala Tragelaphus angasii; middle: Adult female Nyala Tragelaphus angasii; bottom: Subadult Nyala Tragelaphus angasii.

148

Geographic Variation None recorded, but see Taxonomy.

Tragelaphus angasii

Dorsal view of skull of Nyala Tragelaphus angasii.

both within and outside of their former distribution range (including even the farming districts in N Namibia). Where habitats are suitable and the areas selected do not experience cold wet winters, the species has adapted well to artificial dispersion. Animals from NW South Africa have spread into neighbouring parts of Botswana, such as the Tuli Block.

Tragelaphus angasii

Similar Species Tragelaphus spekii. Not sympatric. Similar in body size, but less strikingly marked and has typically elongated hooves as an adaptation to its swamp habitat. The male’s horns are very similar in size and shape to those of the Nyala. T. scriptus. Sympatric throughout the range of the Nyala. Smaller, and without long hair on the underside of adult ??; comparatively straighter and shorter horns and more dependent on closed habitats and a dicotyledonous diet.A similar though less pronounced sexually dimorphic colour change is shown, particularly in the southern subspecies T. s. sylvaticus. Distribution Endemic to Africa. Inhabits dense thickets and open thicket woodland mosaics in Malawi, Mozambique, Zimbabwe, South Africa and Swaziland. Historical Distribution Originally confined to the hot, low-altitude areas of south-eastern Africa, below the 18° isotherm. North of the Zambezi R., found only in the Lower Shire valley in Malawi (Sidney 1965,Ansell 1981). South of the Zambezi R., Nyalas were widespread in Mozambique and occurred as far south as the Hluhluwe R. (about 28° 20' S) in KwaZulu–Natal, South Africa. In Zimbabwe, NE South Africa (KwaZulu–Natal) and NE Swaziland they were found in scattered pockets in suitable habitats, mainly along the major river valleys. In South Africa, Rautenbach (1982) remarks that Nyala occurred naturally as far west as Ellis Ras in what is now Limpopo Province. Nyalas suffered severe mortality during the 1895/96 rinderpest epizootic and this may account for the discontinuous distribution and low numbers encountered at the end of the nineteenth century (Selous 1909). Current Distribution Currently more widespread, because since the early 1960s animals have been translocated from game reserves in KwaZulu–Natal to other protected areas and private ranches in areas

Habitat Thicket and forest are essential habitat requirements for Nyalas. They use these when resting during the heat of the day and seek shelter in them during cold and inclement weather.When alarmed, Nyalas always escape to the nearest cover. At least 10% of the homeranges of animals studied in Hluhluwe G. R. consisted of forest (Anderson 1976) and, in Mozambique, Lobão Tello & Van Gelder (1975) showed that in one year 90% of Nyalas observed were in association with thickets.They frequent both flat and hilly landscapes, naturally occurring up to an altitude of about 400 m, and, although it is not essential, prefer to be in the vicinity of permanent water. At night, Nyalas use open habitats more readily than in daylight, but they are seldom far from cover. In the dry season they regularly use the river flood-plains in Ndumo G. R. Abundance The demand for Nyalas on private ranches has increased their distribution and numbers in South Africa, Swaziland and Zimbabwe, and also in Namibia, where the species has never occurred naturally. East (1999) estimated the total population of Nyalas to exceed around 32,000 individuals. However, more recent estimates suggest South Africa has at least 30,000 (a slight increase from the estimate of East [1999]), with the largest populations (25,000) in KwaZulu–Natal (P. Thomson pers. comm.). The major populations are in the Game Reserves of Ndumo (4000), Mkhuze (7000) and Hluhluwe-iMfolozi (7000) (Rowe-Rowe 1994). Kruger N. P. has in excess of 1000 animals. In Swaziland, the species was extinct by the 1950s, but they have been successfully reintroduced (and introduced to other parts of the country) and there are now more than 1000 on protected areas and ranches (Monadjem 1998). Nyalas are still widespread in Mozambique but, because of uncontrolled hunting, at very low densities and numbers probably do not exceed 3000 (J. Anderson, unpubl.). Zimbabwe has more than 1000, the majority in Gonarezhou N. P. and on game ranches in the south-east. Numbers in Malawi have declined from 3000 (East 1999) to about 1500, most notably in the population in Lengwe N. P. (which was originally created especially for this species) and in Mwabvi G. R. where a relict population is thought still to occur (S. Munthali pers. comm.). There are about 350 in two populations on private property. Extralimital to the species’ natural range, Namibia has about 250, all on private ranches. 149

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Volume I: Introductory Chapters and Afrotheria Edited by Jonathan Kingdon, David Happold, Michael Hoffman, Thomas Butynski, Meredith Happold and Jan Kalina 352pp This first volume in the series comprises eight introductory chapters covering topics such as evolution, geography and geology, biotic zones, classification, behaviour and morphology. The rest of the book is devoted to the Afrotheria, a grouping that comprises six orders and 49 species; these are the hyraxes, elephants, Dugong, manatees, otter-shrews, golden-moles, sengis (elephant-shrews) and Aardvark.

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FULLY ILLUSTRATE D Contents

Series Acknowle

dgements

Acknowledgem

ents for Volum

1. Mammals of

Genus Dugon g Dugo Dugong dugon Dugo ng – H. Marsh ng – H. Marsh & P. Dutto

173 n 174 iDae Manatees – J.e. Reynolds, iii 179 Genus Trichec us Manatees – J.a. Powell Trichecus Senega lensis West africa 180 n Manatee – J.a. Powell, 180 oRDeR HyRa CoiDea Hyrax 19 3. The Evolution es – J. shoshani, & e.R. seiffert of a Continent P. Bloomer Geology – D. : Geography Livingstone & and J. Kingdon 183 FaMiLy PRoC aViiD ae Hyraxes – 25 4. Africa’s Envir & e.R. seiffert J. shoshani, P. Bloom onmental and er Climatic Past R. J. Morley & J. Kingdon – 185 Genus Dendrohyrax Tree Hyraxes – 41 5. The Biotic Zone Dendrohyrax arbore P. Bloomer us southern Tree Perspective – s of Africa: A Mammalian 187 Hyrax (southern Dassie) – J.M. D. C. D. Happo Milner & a. Gayla Tree ld & J. M. Lock Dendrohyrax dorsali rd s Western Tree 55 6. Mammalian 187 Hyrax Hyrax ) (Beecroft’s Tree – s. schult Evolution in Africa – J. Kingd Dendrohyrax validu z, D. Rober ts on s eastern Tree Hyrax 73 7. Classification 190 e. Topp-Jørgen – D. Rober ts, – C. P. Groves sen & D. Moye & D. C. D. Happo r ld 99 Genus Hetero 8. Behaviour and 193 hyrax Bush Hyrax Morphology – J. Kingdon & – P. Bloomer Heterohyrax brucei F. Vollrath 107 Bush 196 R. e. Barry, H.n. Hyrax (yellow-spotted Hyrax )– Hoeck CLass MaMMaLi 196 Genus Procavia a – J. Kingdon Rock Hyrax – P. Bloom Procavia capensis Rock Hyrax (Klipd er H.n. Hoeck 129 suPeRCoHoRT 200 assie) – H.n. Hoeck & P. Bloomer aFRoTHeRia – J. Kingdon, e.R. seiffert, s.B. Hedges & G. Rathb 201 CoHoRT aFRo un inseCTiPHiLLia 137 CoHoRT Paen – e. R. seiffert unGuLaTa – e.R. seiffert 207 oRDeR TuBu LiDenTaTa aardv 141 suPeRoRDeR ark – T. Lehmann TeTHyTHeRia – e.R. seiffert 208 FaMiLy oRyC TeRoPoDiDae 142 oRDeR PRoB aardvark– T. Lehm osCiDea eleph ann ants – J. shoshani 209 Genus Orycte & P. Tassy ropus aardvark– 143 FaMiLy eLeP T. Lehmann & Orycteropus afer HanTiDae eleph a. Taylor aardvark (antb ants – J. shoshani 209 ear)– a. Taylor & P. Tassy 145 Genus Loxodo 210 Bibliography nta african eleph ants – Loxodonta african a savanna eleph P. Tassy & J. shoshani 148 Authors ant (african Bush elephant) – J. H. 216 of Volume I Poole, P. Kahum Loxodonta cycloti bu & i. Whyte s Forest elephant 151 – a. Turkalo & 246 R. Barnes 165 Indexes oRDeR siRen French ia Dugong, Manat ees – D. P. Domn 248 German ing 171 FaMiLy DuG 00 english onGiDae Dugo ng – J.e. Reyno 00 scientific lds iii 173 00 00 Africa: An Intro duction and D. C. D. Happo Guid ld, M. Hoffmann, T. Butynski & J. e – Kingdon mals: An Intro Mammals in duction to Afric Scien an Culture – J. Kingd ce, Natural History and on

2. Thinking Mam

FaMiLy TRiCHeCH

SERIES EDITORS Jonathan Kingdon Department of Zoology, University of Oxford Jonathan Kingdon was born in Tanzania and has spent much of his life in Africa. Over the course of a long career he has achieved acclaim as both a leading academic and a prominent artist. Professor Kingdon is also the author of The Kingdon Field Guide to African Mammals and The Kingdon Pocket Guide to African Mammals. David C. D. Happold Research School of Biology, Australian National University David Happold has held academic positions at the University of Khartoum (Sudan), University of Ibadan (Nigeria), University of Malawi, and The Australian National University. Thomas M. Butynski Zoological Society of London/King Khalid Wildlife Research Centre Tom Butynski is a conservationist and ecologist who has worked in Africa for 35 years, mostly in Botswana, Kenya, Uganda and Equatorial Guinea.

Michael Hoffmann International Union for Conservation of Nature â&#x20AC;&#x201C; Species Survival Commission Mike Hoffmann trained as a mammalogist at the University of Pretoria, South Africa and the University of Oxford, UK. He has since moved into the field of international biodiversity conservation. Meredith Happold Research School of Biology, Australian National University Meredith Happold spent several years studying bats in Nigeria. Since 1977 she has held a Visiting Fellowship at the Australian National University and her research has focused on African bats. Jan Kalina Soita Nyiro Conservancy, Kenya Jan Kalina is a conservation biologist. Her work with mammals began at Wildlife Conservation Societyâ&#x20AC;&#x2122;s Bronx Zoo in New York and at Colorado State University. She is currently joint-owner of Soita Nyiro Conservancy in Laikipia, Kenya.

Volume IV: Hedgehogs, Shrews and Bats Edited by Meredith Happold and David Happold 800pp Profiles of 156 species of insectivores, comprising the hedgehogs and shrews. The rest of the volume is devoted to the 224 species of African bats. The latter are divided into nine families, namely fruit bats, horseshoe bats, leaf-nosed bats, false vampire bats, mousetailed bats, sheath-tailed bats, slit-faced bats, free-tailed bats and vesper bats.

Volume V: Carnivores, Pangolins, Equids and Rhinoceroses Edited by Jonathan Kingdon and Michael Hoffmann 560pp Comprises 83 species of carnivores, and includes jackals, wolves, dogs, foxes, weasels, polecats, striped weasels, Zorilla, otters, Ratel, fur seals, monk seals, Palm Civet, cats, genets, linsangs, African Civet, hyaenas, Aardwolf and mongooses. The volume is completed with profiles of four pangolins, four zebras and two rhinoceroses.

Volume VI: Pigs, Hippopotamuses, Chevrotain, Giraffes, Deer and Bovids Edited by Jonathan Kingdon and Michael Hoffmann 704pp Comprises a single order, currently subdivided into three suborders, containing the pigs, hippopotamuses, chevrotains, Giraffe, Okapi, deer, buffalos, spiral-horned antelopes, dwarf antelopes, duikers, grysboks, Beira, dik-diks, gazelles, Klipspringer, Oribi, reduncines, Impala, alcelaphines, horse-like antelopes, sheep and goats; the volume contains 98 species profiles.

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This is the eighth and final volume in the authoritative Birds of Africa series. It covers the Malagasy region, including Madagascar and various islands and archipelagos of the Indian Ocean including Seychelles, Comoros, Mauritius and Réunion. Every resident and migrant species is covered in full detail, comparable to other volumes in the series, and with a colour map for each species. Vagrants are treated in less detail. All species are illustrated on a beautiful series of 64 colour plates, with original artwork from John Gale and Brian Small. Hardback | 960pp | 300x234mm | Illustrations: 64 colour plates | May 2013 | ISBN 9780713665321 | Price (AUD$250* until 31 August 2013) $300 | ePDF: 9781408190494

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A major new multi-volume reference work on African mammals and a landmark publishing event. Covers all 1,160 species of African mammal in unprecedented detail. Mammals of Africa is a splendid rare blend of natural history and science. It takes handbooks to a new level and shows what professionals and amateurs alike can expect if they wish to carry with them the very best. E. O. Wilson, Harvard University

Mammals of Africa is the magnum opus of Kingdon’s team - except that it seems more magnum than any opus has any right to be. It is up-to-date, comprehensive and, above all, beautiful, in both the writing and the pictures. Richard Dawkins, New College, University of Oxford

mammals of africa volume I

introductory chapters and afrotheria edited by jonathan kingdon, david happold, michael hoffmann, thomas butynski, meredith happold and jan kalina

Art in science, the science of art, and the art of science: say these words to informed biologists worldwide, and with one voice they will acclaim: Jonathan Kingdon. This marvellous book, compendious, meticulous, beautiful and inspirational is the crescendo of a lifetime’s work in and for Africa. No other continent can boast such a tome as Mammals of Africa, and Africa and her magnificent mammals owe it to Jonathan Kingdon and his remarkable team. David W. Macdonald CBE, Professor of Wildlife Conservation, University of Oxford

Mammals of Africa represents a high-water mark - not just in mammalogy, but in scientific publishing overall. Tim Flannery, Macquarie University, Sydney

Format: 279 x 217mm Publication: February 2013 Binding: Hardback, in 6 volumes Extent: 3,760 pages Illustrations: 660 colour paintings, 2,000+ line drawings, 1,100 maps

Mammals new.indd 1

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12/09/2012 15:13